This paper examines the following questions: (1) Do early pioneer species have a greater impact on the survival and growth of different successional native trees compared to the shrub Rhodomyrtus tomentosa? (2) Do canopy treatments affect soil nutrients and light availability? (3) What is the mechanism underlying the interaction between nurse species and target species? Degraded shrubland sites (Heshan, Guangdong, China), under moist subtropical conditions were studied. About 1-year-old seedlings of Pinus massoniana, Schima wallichii, Schefflera heptaphylla, Castanopsis hystrix, Cryptocarya chinensis and Castanea chinensis were transplanted under the canopy of R. tomentosa, Dicranopteris dichotoma and in open interspaces without vegetation. Survival and growth were recorded from the first growing season after planting. Leaf gas exchange, water potential, soil physicochemical characters and irradiation were then measured. Canonical redundancy analyses (RDA) were used to evaluate the relationships between environmental factors and seedling survival and growth conditions. Both R. tomentosa and D. dichotoma canopy treatment facilitated seedling survival and growth either directly or indirectly. Irradiance/radiation was considered the most important resource (factor) for seedling growth in subtropical regions, however, soil nutrients and species are yet to be examined simultaneously with irradiance/radiation under field conditions. We conclude that early successional species facilitates the survival and growth of late successional species in subtropical shrublands. However, further predictions of successional trajectories remain elusive and are influenced by stochastic processes, including arrival order, shade tolerance, physiological character of the colonizing species and their competitive interactions.
Anthelmea, F. and R. Michalet. 2009. Grass-to-tree facilitation in an arid grazed environment (Aïr Mountains, Sahara). Basic Appl. Ecol. 10: 437–446.
Bainbridge, D. 1994. Tree shelters improve establishment on dry sites. Tree Planters Note 45: 13–16.
Bertness, M.D. and R.M. Callaway. 1994. Positive associations in communities. Trends Ecol. Evol. 9: 191–193.
Brooker, R.W., F.T. Maestre, R.M. Callaway, C.L. Lortie, L.A. Cavieres, G. Kunstler, P. Liancourt, K. Tielboerger, J. Travis, F. Anthelme, C. Armas, L. Coll, E. Corcket, S. Delzon, E. Forey, Z. Kikvidze, J. Olofsson, F. Pugnaire, C.L. Quiroz, P. Saccone, K. Schiffers, M. Seifan, B. Touzard and R. Michalet. 2008. Facilitation in plant communities: the past, the present, and the future. J. Ecol. 96: 18–34.
Bruno, J.F., J.J. Stachowicz and M.D. Bertness. 2003. Inclusion of facilitation into ecological theory. Trends Ecol. Evol. 18: 119–125.
Callaway, R.M. 1995. Positive interactions among plants. Bot. Rev. 61: 306–349.
Callaway, R.M. and L.R. Walker. 1997. Competition and facilitation: a synthetic approach to interactions in plant communities. Ecology 78: 1958–1965.
Castro, J., R. Zamora and J.A. Hódar. 2004. Benefits of using shrubs as nurse plants for reforestation in Mediterranean mountains: a 4-year study. Restor. Ecol. 12: 352–358.
Chaar, H., T. Mechergui, A. Khouaja and H. Abid. 2008. Effects of treeshelters and polyethylene mulch sheets on survival and growth of cork oak (Quercus suber L.) seedlings planted in northwestern Tunisia. Forest Ecol. Manag. 256: 722–731.
Cushman, J.H., J.C. Waller and D.R. Hoak. 2010. Shrubs as ecosystem engineers in a coastal dune: influences on plant populations, communities and ecosystems. J. Veg. Sci. 21: 821–831.
Drake, D.R. and L.W. Pratt. 2001. Seedling mortality in Hawaiian rain forests: the role of small-scale physical disturbance. Biotropica 33: 319–323.
Drezner, T.D. 2007. An analysis of winter temperature and dew point under the canopy of a common Sonoran Desert nurse and the implications for positive plant interactions. J. Arid. Environ. 69: 554–568.
Duan, W.J., H. Ren, S.L. Fu, Q.F. Guo and J. Wang. 2008. Pathways and determinants of early spontaneous vegetation succession in degraded lowland of South China. J. Integr. Plant Biol. 50: 147–156.
Fahey, T.J., J.J. Battles and G.F. Wilson. 1998. Responses of early successional northern hardwood forests to changes in nutrient availability. Ecol. Monogr. 68: 183–212.
Francisco, M.P. and F.I. Pugnaire. 2010. Species identity and water availability determine establishment success under the canopy of, Retama sphaerocarpa shrubs in a dry environment. Restor. Ecol. 17: 900–907.
Gómez-Aparicio, L., R. Zamora, J.M. Gómez, J.A. Hódar, J. Castro and E. Baraza. 2004. Applying plant facilitation to forest restoration: A meta-analysis of the use of shrubs as nurse plants. Ecol. Appl. 14: 1128–1138.
Holmgren, M. and M. Scheffer. 2010. Strong facilitation in mild environments: the stress gradient hypothesis revisited. J. Ecol. 98: 1269–1275.
Holste, E.K., R.K. Kobe and C.F. Vriesendorp. 2011. Seedling growth responses to soil resources in the understory of a wet tropical forest. Ecology 92: 1828–1838.
Holzapfel, C., K. Tielborger, H.A. Parag, J. Kigel and M. Sternberg. 2006. Annual plant-shrub interactions along an aridity gradient. Basic Appl. Ecol. 7: 268–279.
Kitzberger, T., D.F. Steinaker and T.V. Veblen. 2000. Effects of climatic variability on facilitation of tree establishment in northern Patagonia. Ecology 81: 1914–1924.
Kobe, R.K. 1999. Light gradient partitioning among tropical tree species through differential seedling mortality and growth. Ecology 80: 187–201.
Kobe, R.K., J.A. Silander and C.D. Canham. 1995. Juvenile tree survivorship as a component of shade tolerance. Ecol. Appl. 5: 517–532.
Lawrence, D. 2003. The response of tropical tree seedlings to nutrient supply: meta-analysis for understanding a changing tropical landscape. J. Trop. Ecol. 19: 239–250.
Lepš, J. and P. Šmilauer. 2003. Multivariate Analysis of Ecological Data Using CANOCO. Cambridge Univ. Press, Cambridge.
Liu, G.S. 1996. Soil physical and chemical analysis and description of soil profiles. Standards Press of China, Beijing, China.
Oesterheld, M. and M. Oyarzabal. 2004. Grass-to-grass protection from grazing in a semi-arid steppe. Facilitation, competition, and mass effect. Oikos 107: 576–582.
Padilla, F.M., J.d.D. Miranda, R. Ortega, M. Hervás, J. Sánchez and F.I. Pugnaire. 2011. Does shelter enhance early seedling survival in dry environments? A test with eight Mediterranean species. Appl. Veg. Sci. 14: 31–39.
Peace, W.J.H. and P.J. Grubb. 1982. Interaction of light and mineral nutrient supply in the growth of, Impatiens parviflora. New Phytol. 90: 127–150.
Pearcy, R.W. 2007. Responses of plants to heterogeneous light environments. In: Pugnaire F. and F. Valladares (Eds.), Functional Plant Ecology. Taylor & Francis, New York, NY, USA, pp. 213–258.
Peng, S.L., J. Liu and H.F. Lu. 2005. Characteristics and role of, Acacia auriculiformis on vegetation restoration in lower subtropics of China. J. Trop. For. Sci. 17: 508–525.
Pugnaire, F.I., C. Armas and F. Valladares. 2004. Soil as a mediator in plant-plant interactions in a semi-arid community. J. Veg. Sci. 15: 85–92.
Pugnaire, F.I., P. Haase, J. Puigdefabregas, M. Cueto, S.C. Clark and L. Incoll. 1996. Facilitation and succession under the canopy of a leguminous shrub, Retama sphaerocarpa, in a semi-arid environment in south-east Spain. Oikos 76: 455–464.
Ren, H., X.A. Cai, C.H. Li and Y.S. Ye. 2010. Atlas on Tool Species of Vegetation Recovery in South China. Huazhong University of Science & Technology Press, Wuhan.
Rey-Benayas, J.M. 1998. Growth and survival in, Quercus ilex L. seedlings after irrigation and artificial shading on Mediterranean set-aside agricultural land. Ann. Sci. Forest. 55: 801–807.
Rodríguez-Calcerrada, J., J.A. Pardo, L. Gil and I. Aranda. 2008. Ability to avoid water stress in seedlings of two oak species is lower in a dense forest understory than in a medium canopy gap. Forest Ecol. Manag. 255: 421–430.
Russell, A.E., J.W. Raich and P.M. Vitousek. 1998. The ecology of the climbing fern, Dicranopteris linearis on windward Mauna Loa, Hawaii. J. Ecol. 86: 765–779.
Russo, S.E., P. Brown, S. Tan and S.J. Davies. 2008. Interspecific demographic trade-offs and soil-related habitat associations of tree species along resource gradients. J. Ecol. 96: 192–203.
Scholes, R.J. and S.R. Archer. 1997. Tree-grass interactions in savannas. Annu. Rev. Ecol. Syst. 28: 517–544.
Shachak, M., B. Boeken, E. Groner, R. Kadmon, Y. Lubin, E. Meron, G. Ne'Eman, A. Perevolotsky, Y. Shkedy and E.D. Ungar. 2008. Woody species as landscape modulators and their effects on biodiversity patterns. BioScience 58: 209–221.
Sharrow, S.H. 2001. Effects of shelter tubes on hardwood tree establishment in western Oregon silvopastures. Agroforest. Syst. 53: 283–290.
Shiels, A.B., L.R. Walker and D.B. Thompson. 2006. Organic matter inputs create variable resource patches on Puerto Rican landslides. Plant Ecol. 184: 223–236.
Smit, C., J. DenHouden and H. Müller-Schärer. 2006. Unpalatable plants facilitate tree sapling survival in wooded pastures. J. Appl. Ecol. 43: 305–312.
Walker, L.R., F.H. Landau, E. Velázquez, A.B. Shiels and A.D. Sparrow. 2010. Early successional woody plants facilitate and ferns inhibit forest development on Puerto Rican landslides. J. Ecol. 98: 625–635.
Walker, L.R. and A.B. Shiels. 2008. Post-disturbance erosion impacts carbon fluxes and plant succession on recent tropical landslides. Plant Soil. 313: 205–216.
Walker, L.R., D.J. Zarin, N. Fetcher, R.M. Myster and A.H. Johnson. 1996. Ecosystem development and plant succession on landslides in the Caribbean. Biotropica 28: 566–576.
Wright, J.P., C.G. Jones, B. Boeken and M. Shachak. 2006. Predictability of ecosystem engineering effects on species richness across environmental variability and spatial scales. J. Ecol. 94: 815–824.
Yang, L., N. Liu, H. Ren and J. Wang. 2009. Facilitation by two exotic, Acacia: Acacia auriculiformis and Acacia mangium as nurse plants in South China. Forest. Ecol. Manag. 257: 1786–1793.
Yang, L., H. Ren, N. Liu and J. Wang. 2010. The shrub, Rhodomyrtus tomentosa acts as a nurse plant for seedlings differing in shade tolerance in degraded land of South China. J. Veg. Sci. 21: 262–272.