Temporal changes of fine root overyielding and foraging strategies in planted monoculture and mixed forests

Weiwei Shu, Xiaoxiao Shen, Pifeng Lei, Wenhua Xiang, Shuai Ouyang, Wende Yan

Mixed forests are believed to enhance ecosystem functioning and sustainability due to complementary resource use, environmental benefits and improved soil properties. The facilitation between different species may induce overyielding. Meanwhile, the species-specific fine root foraging strategies and tradeoffs would determine the structure and dynamics of plant communities. Here the aim was to investigate the admixing effects of fine-root biomass, vertical distribution and morphology in Pinus massoniana-Cinnamomum camphora mixed plantations and corresponding monocultures at 10-, 24- and 45-year old stands. The fine root biomass in the Pinus-Cinnamomum mixed forest exerted a certain degree of overyielding effect. These positive admixing effects, however, did not enhance with forest stand development. The overall relative yield total ranged from 1.83 and 1.51 to 1.33 in 10-, 24- and 45-year-old stand, respectively. The overyielding was mainly attributed to the over-performance of late successional species, Cinnamomum, in mixed stands. The vertical fine root biomass distribution model showed fine roots of pioneer species, Pinus, shifted to the superficial layer when mixed with Cinnamomum. Furthermore, the specific root length (SRL) of Pinus was significantly higher in Pinus-Cinnamomum mixed stands than that in monocultures, and the magnitude of differences increased over time. However, the vertical fine-root distribution and SRL for Cinnamomum did not show significant differences between monoculture and mixtures. Our results indicated that the magnitude of fine root overyielding in mixed forests showed a high degree of consistency with the total amount of fine root biomass itself, suggesting the overyielding effects in mixed forests were correlated with the degree of belowground interaction and competition degree involved. The late successional species, Cinnamomum, invested more carbon to belowground by increasing the fine root biomass in mixtures. While the pioneer species, Pinus, adapted to the presence of the species Cinnamomum by modification of vertical distribution and root morphological plasticity in the mixtures. These species-specific fine root foraging strategies might imply the differences of forest growth strategies of co-occurring species and contribute to the success and failure of particular species during the succession over time.