Abstract
The impact of biodiversity loss on the functioning of forest ecosystems has become a central issue in ecology. Most reports of the positive effects of tree mixture on the biodiversity–productivity relationship focus on mixtures that combine tree species with contrasting traits. Nevertheless, little is known about how coniferous mixtures of the same genus affect forest productivity, what mechanisms are involved, and how the understory is affected. Here, we assessed the effect of mixed versus monospecific stands of Pinus sylvestris L. and P. pinaster Ait. on productivity, its impact on the understory, and its relationship with soil water and fertility, based on research with six triplets (6 triplets × 3 forest stands × 1 plot = 18 plots) in North-Central Spain. Each triplet consisted of two plots dominated either by P. sylvestris or P. pinaster and of one mixed plot that contained both species. Productivity, at the stand and neighborhood levels, and the understory richness, and soil water and fertility at the stand level were analyzed. A positive effect of pine mixture on productivity was observed at the smaller spatial scale, and it had no negative effect on the understory richness. The greater space-use efficiency (higher tree density and basal area) of both Pinus species in the admixtures was related to soil water and fertility niche complementarity. The fundamental role of scale in determining the relationship between species richness and ecosystem functioning in forests is highlighted.
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Acknowledgements
We would like to thank Luis Alfonso Ramos Calvo for his invaluable help with soil sampling, Carmen Blanco and Juan Carlos Arranz for their advice in the laboratory analyses, José Riofrío and Cristóbal Ordoñez for their assistance in the characterization and location of plots in the field, and the Regional Forest Service of Castilla and León to facilitate the triplet’s installation and monitoring. We also thank Josu G. Alday for his assistance with structural equations modeling and Juan Manuel Diez Clivillé (deceased) for his assistance with English.
Funding
This research was funded by a predoctoral grant to DLM (BES-2015-072852) and the Project FORMIXING (AGL2014-51964-C2-1-R) from the Ministry of Economy and Competitiveness of the Spanish Government.
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DLM carried out the field and laboratory work, ran the data analysis, and discussed the results. DLM and CMR discussed data analysis and commented on the results and discussion. CMR supported DLM with the statistical analysis. MBT supported DLM with the laboratory analysis. DLM, CMR, MBT, and FB edited the manuscript. FB coordinated the research project.
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Appendices
Appendix 1: Data analyses of overstory and soil properties
Basal area of Pinus sylvestris (G-PS)
G-PS = ∑ gPSi /Si | gPSi: section of Pinus sylvestris stem (m2) Si: surface (ha) |
Section of Pinus sylvestris stem (gPSi)
gPSi = π/4 dnPSi2 | dnPSi: normal diameter of every Pinus sylvestris stem > 7.5 cm at the breast height (m) |
Basal area of Pinus pinaster (G-PP)
G-PP = ∑ gPPi /Si | gPPi: section of Pinus pinaster stem (m2) Si: surface (ha) |
Section of Pinus pinaster stem (gPPi)
gPPi = π/4 dnPPi2 | dnPPi: normal diameter of every Pinus pinaster stem > 7.5 cm at the breast height (m) |
Total basal area (G-T)
G-T = G-PS + G-PP | G-PS: basal area of Pinus sylvestris G-PP: basal area of Pinus pinaster |
Water holding capacity
Water holding capacity of each horizon (WHCHi)
WHCHi = UWHi⋅bDHi⋅%EFHi THi | UWHi: useful water of each horizon bDHi: bulk density of each horizon %EFHi: % of earth fraction of each horizon THi: thickness of each horizon |
Water holding capacity in the whole mineral soil profile (0–50 cm; WHC)
WHC = ∑ WHCHi. |
Sum of bases stock
Sum of bases stock of each horizon (SBstockHi)
SBstockHi = SBHi⋅bDHi⋅%EFHi THi | SBHi: sum of bases of each horizon bDHi: bulk density of each horizon %EFHi: % of earth fraction of each horizon THi: thickness of each horizon |
Sum of bases stock in the whole mineral soil profile (0–50 cm; SBstock)
SBstock = ∑ SBstockHi. |
Appendix 2
See Table 3.
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López-Marcos, D., Turrión, MB., Bravo, F. et al. Overyielding in mixed pine forests with belowground complementarity: impacts on understory. Eur J Forest Res 140, 777–791 (2021). https://doi.org/10.1007/s10342-021-01365-0
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DOI: https://doi.org/10.1007/s10342-021-01365-0