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Compensatory Effects Between Pinus Massoniana and Broadleaved Tree Species

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Compensatory Effects Between Pinus Massoniana and Broadleaved Tree Species Journal of Plant Ecology Compensatory effects between VOLUME 3, NUMBER 3, PAGES 183–189 Pinus massoniana and broadleaved SEPTEMBER 2010 doi: 10.1093/jpe/rtq020 tree species Advanced Access published on 4 August 2010 1,2,3 1 1,2 1 1, available online at Qian Li ,YuLiang, Bo Tong , Xiaojun Du and Keping Ma * www.jpe.oxfordjournals.org 1 State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 10093, China 2 Graduate School of Chinese Academy of Sciences, Beijing 100049, China Downloaded from https://academic.oup.com/jpe/article/3/3/183/960486 by guest on 30 September 2021 3 College of Life Science and Technology, Central South University of Forestry and Technology, Changsha 410004, China *Correspondence address. State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China. Tel: +86-10-6283-6223; Fax: +86-10-8259-9518; E-mail: [email protected] Abstract Aims P. massoniana. Pinus massoniana had competitive effects on Litho- Evaluation of the interaction intensity between Masson pine (Pinus carpus glaber, Cyclobalanopsis glauca and Elaeocarpus japonicus. massoniana) and broadleaved trees will facilitate selecting tree com- Significantly negative relations were found between biomass of binations suitable for reforestation in abandoned sites in subtropical P. massoniana and the broadleaved trees in the third year of the ex- areas. periment, and the compensatory effects between P. massoniana and the broadleaved trees may be involved in stability maintenance in the Methods multi-species forests in the subtropical area. The results of homoge- Pinus massoniana and seven broadleaved trees species were grown neity test of variances also showed that the biomass per pot in the either in monoculture or in two-species mixture. Biomass of tree spe- mixture had significantly lower variances than that in the monocul- cies was measured and inter-specific interactions were estimated us- ture, suggesting that total biomass is more stable in the mixture than ing log response ratio. Test of homogeneity of variances was the monoculture. performed to compare the stability of biomass in the monoculture of the broadleaved trees with that in the mixture. Keywords: competitive effects d plant–plant interactions Important Findings d facilitative effects d subtropical forest Our results showed that the direction and intensity of interactions Received: 6 May 2010 Revised: 16 June 2010 Accepted: 16 June between P. massoniana and the broadleaved trees varied from year 2010 to year and the identity of the broadleaved species. Facilitative inter- actions were found between Camellia oleifera, Rhus chinensis and INTRODUCTION ative effects of neighbor plants may result from inter-specific competition for nutrients, water or light. In contrast, the accu- Inter-specific interactions between plants are the essential mulation of nutrients, provision of shade, amelioration of dis- drivers in structuring plant communities (Callaway and turbance or protection from herbivores by some species may Walker 1997; Fowler 2003). The nature of plant–plant inter- lead to facilitative effects of neighbor plants (Fernando et al. actions is diverse, with the potential net outcome of these 2004). interactions ranging from positive to negative (Goldberg Evergreen broad-leaved forests formerly covered ;25% 1990). Both positive and negative effects may exist between area in China (Wang et al. 2007). However, most of the sub- two plant species and the net effect of a plant–plant interaction tropical forests have been destroyed by clear logging in the depends on the balance of negative and positive effects 1950’s and the 1960’s due to policy and economic reasons. (Callaway and Walker 1997; Chapin et al. 1994; Maestre In recent years, China has made great efforts in reforestation et al. 2004; Maestre et al. 2009; Niu and Wan 2008). The neg- and ecological restoration in the disturbed areas. It has been Ó The Author 2010. Published by Oxford University Press on behalf of the Institute of Botany, Chinese Academy of Sciences and the Botanical Society of China. All rights reserved. For permissions, please email: [email protected] 184 Journal of Plant Ecology a long-term debate in forestry of which kinds of forests, pure or Experimental design and sampling mixed species, should be used in reforestation (Berkowitz et al. One conifer species, P. massoniana, and seven broadleaved tree 1995; Brenton and Facelli 2005; Knoke et al. 2008). Owing to species, i.e. Quercus variabilis (Fagaceae), Q. glandulifera (Faga- the advantages of mixed-species forests in fireproofing, path- ceae), Cyclobalanopsis glauca (Fagaceae), Lithocarpus glaber (Faga- ogen resistance and higher productivity in many countries in ceae), Rhus chinensis (Anacardiaceae), Elaeocarpus japonicus the world, it is currently a major objective of forest manage- (Elaeocarpaceae) and Camellia oleifera (Theaceae), were used ment and policy to convert mono-species coniferous forests in- in our study. Quercus variabilis, Q. glandulifera and R. chinensis to multi-species forests (Baumgarten and von Teuffel 2005; are deciduous, and C. glauca, L. glaber, E. japonicus and C. oleifera Fritz 2006; Knoke et al. 2008). How to select suitable plant spe- are evergreen trees. Plant seeds were sterilized with a 0.1% cies or species combinations from many candidate plant spe- hypochloric acid solution for 3 min and sown in March cies is essential for the success of reforestation in terms of 2003. Tree seedlings were either grown as monoculture or biomass accumulation and ecosystem stability. two-species mixture of P. massoniana and one broadleaved Pinus massoniana (Masson pine) is a tree species commonly species in pots (38 cm in diameter and 40 cm in height). Four Downloaded from https://academic.oup.com/jpe/article/3/3/183/960486 by guest on 30 September 2021 used in reforestation of disturbed sites in subtropical area of individuals of single species were kept in each pot as the mono- China, and has been documented to have positive effects on culture treatment and two individuals of each species were kept soil and water conservation in forest plantations (Wang et al. in each pot as the mixture treatment. Fifteen replicates per spe- 2007). Similar to the situation in other countries, mixed for- cies combination were used in the experiment, resulting in a to- ests are more and more popular in reforestation in China. tal of 225 pots (15 species combinations 3 15 replicates). During However, lack of information on the interactions between the experiment duration, the pots were kept under conditions pine and broadleaved tree species makes it difficult to select with ambient temperature and precipitation. suitable species combinations in the establishment of mixed Plants were harvested in the end of October in 2003, 2004 forests. Evaluation on the performances of both pine and and 2005. Three replicates of each species combination were broadleaved species in species mixtures will be therefore harvested in each year. Roots and shoots were separated for helpful in selecting suitable species combinations in refores- all seedlings, oven dried at 70°C for 48 h and weighed. tation (Wang et al. 2002; White et al. 1999; Zavala and Zea 2004). Statistical analyses In the present study, P. massoniana and seven common Shoot, root and total plant biomass of P. massoniana and broadleaved trees species of the local forest were grown in broadleaved trees in the mixture were compared with those monoculture or two-species mixture. Growth of tree species in the monoculture using Student’s t-test in SPSS for win- were determined and inter-specific interactions between dows (version 13.0, SPSS Inc., USA). We used log response P. massoniana and the broadleaved trees were measured. ratio (lnRR) to evaluate the response of one species to the The specific objectives of this study are (i) to evaluate the in- presence of the other. LnRR = ln(Xmix/Xmono), where X is teraction intensity between P. massoniana and the broadleaved an estimation of plant performance in the presence (mix) trees and (ii) to select species combinations suitable for refor- or in the absence (mono) of neighbors (Oksanen et al. estation in the abandoned sites in subtropical area. 2006; Weigelt and Jolliffe 2003). Positive values of lnRR in- dicate facilitative effects of heterospecific neighbor species on target species and negative values indicate competitive inhi- MATERIALS AND METHODS bition of neighbor species on target species. Test of homoge- neity of variances was performed using Levene statistic in Site descriptions SPSS for windows (version 13.0, SPSS Inc.) to compare the # The experiment was conducted in Dujiangyan city (30°45 to stability of biomass in the monoculture of broadleaved trees # # # 31°22 N, 103°25 to 103°47 E), Sichuan Province, China. The with that in the mixture. Correlation between biomass of site has a mean annual precipitation of 1 244 mm, a mean an- P. massoniana and broadleaved trees were analyzed by linear nual temperature of 15.2°C and an altitude of ;780 m above regression using SPSS for windows. the sea level. The typical vegetation of this region is subtropical broad-leaved evergreen forest and the dominant tree species RESULTS belong to families of Fagaceae, Lauraceae, Theaceae, Magno- liaceae and Pinaceae (Du et al. 2007). Soil was taken from an Responses of P. massoniana to broadleaved trees abandoned site where clear-cut logging was carried out at the In the first year, no broad-leaved trees showed significantly end of 1950s and early of 1960s (sees the descriptions in negative effects on seedling growth of P. massoniana Du et al. 2008). The abandoned site was dominated by a variety (Fig. 1a). Two species, L. glaber and R. chinensis, showed signif- of annual and perennial ferns, grasses, herbs and some shrubs icantly facilitative effects on shoot biomass of P. massoniana. (Liang et al. 2004). Soil organic matter (SOM) and nitrogen A significantly higher root biomass of P.
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