Partitioning changes in ecosystem productivity by effects of species interactions in biodiversity experiments
Figures

Under the competitive hypothesis, more competitive species gain size (biomass or volume) and less competitive species lose size in mixture relative to their full density monocultures.
The magnitude of competitive growth responses (i.e. proportional changes in individual size) increases with relative competitive ability (proportional deviations of species competitive ability from community average) from a minimum of 0 (at community average or null expectation) to a maximum of Wip/Wif – 1 for more competitive species and –1 for less competitive species at competitive exclusion. The changes may not be linear, greater near community average and smaller with deviations of competitive ability (Gaudet and Keddy, 1988) or resource availability (e.g. light, see Brüllhardt et al., 2020) away from community averages. Wif represents individual size in full density monocultures, and Wip represents individual size in partial density monocultures.

Partitioning net biodiversity effects with competitive and additive partitioning models.
Partitioning net biodiversity effects (changes in stand volume in mixtures relative to full density monocultures) with competitive (a and b) and additive (c and d) partitioning models for 20- to 80-year-old mixed trembling aspen and white spruce with varying species compositions from nearly pure aspen (90% aspen) to nearly pure spruce (90% spruce) on average medium productivity sites in western Canada. The growth and yield data (Table 1) are generated with GYPSY (Huang et al., 2009), and calculations of biodiversity components with this figure are detailed in Supplementary file 1A.
Tables
Simulated and experimental species mixtures.
Biodiversity effects with partial density monocultures (BPM) are calculated from the differences between the partial density monoculture yield of more competitive species and total mixture yield expected from species relative yields and full density monoculture yields (see competitive exclusion in Supplementary file 1A and B for detailed calculations).
Species mixture (species1:species2) | Age | Total density | Mixture composition | Full density monoculture yields | Observed yields in mixtures | Partial density monoculture yields | |||||
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Species1 | Species2 | Species1 | Species2 | Species1 | Species2 | Species1 | Species2 | BPM | |||
Simulated mixed trembling aspen and white spruce in western Canada generated with GYPSY model (Huang et al., 2009) (density and stand volume yield, stems and m3 per hectare) | |||||||||||
Populus:Picea | 20 years | 11,000 | 0.9 | 0.1 | 23.7 | 0.7 | 22.3 | 0.1 | 23.3 | 0.5 | 1.9 |
Populus:Picea | 20 years | 11,000 | 0.7 | 0.3 | 23.7 | 0.7 | 19.0 | 0.2 | 22.5 | 0.6 | 5.7 |
Populus:Picea | 20 years | 11,000 | 0.5 | 0.5 | 23.7 | 0.7 | 15.1 | 0.3 | 21.4 | 0.6 | 9.2 |
Populus:Picea | 20 years | 11,000 | 0.3 | 0.7 | 23.7 | 0.7 | 10.6 | 0.4 | 19.8 | 0.7 | 12.2 |
Populus:Picea | 20 years | 11,000 | 0.1 | 0.9 | 23.7 | 0.7 | 5.0 | 0.5 | 16.9 | 0.7 | 13.9 |
Populus:Picea | 40 years | 3,600 | 0.9 | 0.1 | 110.4 | 27.2 | 104.5 | 3.4 | 108.3 | 15.5 | 6.3 |
Populus:Picea | 40 years | 3,600 | 0.7 | 0.3 | 110.4 | 27.2 | 90.3 | 7.6 | 103.5 | 20.7 | 18.1 |
Populus:Picea | 40 years | 3,600 | 0.5 | 0.5 | 110.4 | 27.2 | 72.4 | 12.1 | 97.0 | 23.4 | 28.2 |
Populus:Picea | 40 years | 3,600 | 0.3 | 0.7 | 110.4 | 27.2 | 49.6 | 17.5 | 87.4 | 25.3 | 35.2 |
Populus:Picea | 40 years | 3,600 | 0.1 | 0.9 | 110.4 | 27.2 | 20.6 | 23.9 | 70.6 | 26.7 | 35.1 |
Populus:Picea | 60 years | 1,800 | 0.9 | 0.1 | 189.7 | 88.3 | 180.5 | 9.6 | 186.1 | 47.0 | 6.6 |
Populus:Picea | 60 years | 1,800 | 0.7 | 0.3 | 189.7 | 88.3 | 158.1 | 22.4 | 177.3 | 64.3 | 18.0 |
Populus:Picea | 60 years | 1,800 | 0.5 | 0.5 | 189.7 | 88.3 | 128.1 | 38.1 | 164.8 | 73.7 | 25.8 |
Populus:Picea | 60 years | 1,800 | 0.3 | 0.7 | 189.7 | 88.3 | 85.4 | 58.5 | 144.8 | 80.5 | 26.1 |
Populus:Picea | 60 years | 1,800 | 0.1 | 0.9 | 189.7 | 88.3 | 28.6 | 83.0 | 109.2 | 85.9 | 10.8 |
Populus:Picea | 80 years | 1,100 | 0.9 | 0.1 | 240.2 | 143.1 | 229.6 | 13.7 | 235.6 | 73.3 | 5.1 |
Populus:Picea | 80 years | 1,100 | 0.7 | 0.3 | 240.2 | 143.1 | 203.7 | 33.0 | 224.1 | 102.0 | 13.1 |
Populus:Picea | 80 years | 1,100 | 0.5 | 0.5 | 240.2 | 143.1 | 167.3 | 58.2 | 207.6 | 117.8 | 15.9 |
Populus:Picea | 80 years | 1,100 | 0.3 | 0.7 | 240.2 | 143.1 | 110.3 | 94.0 | 179.1 | 129.4 | 6.8 |
Populus:Picea | 80 years | 1,100 | 0.1 | 0.9 | 240.2 | 143.1 | 26.2 | 139.6 | 124.5 | 138.9 | –28.4 |
Experimental grassland mixtures (density and aboveground biomass yield, stems, and grams per pot) (Mahaut et al., 2020) | |||||||||||
Bromus:Dactylis | 13 weeks | 6 | 0.5 | 0.5 | 5.5 | 9.8 | 2.6 | 8.6 | 5.3 | 11.5 | 3.8 |
Bromus:Lotus | 13 weeks | 6 | 0.5 | 0.5 | 5.5 | 6.3 | 3.8 | 3.2 | 5.3 | 4.7 | –1.2 |
Bromus:Plantago | 13 weeks | 6 | 0.5 | 0.5 | 5.5 | 8.6 | 1.9 | 10.9 | 5.3 | 16.1 | 9.0 |
Bromus:Sanguisorba | 13 weeks | 6 | 0.5 | 0.5 | 5.5 | 4.4 | 2.9 | 2.2 | 5.3 | 3.6 | 0.3 |
Bromus:Trifolium | 13 weeks | 6 | 0.5 | 0.5 | 5.5 | 13.3 | 3.8 | 9.3 | 5.3 | 14.2 | 4.8 |
Dactylis:Lotus | 13 weeks | 6 | 0.5 | 0.5 | 9.8 | 6.3 | 11.8 | 2.0 | 11.5 | 4.7 | 3.4 |
Dactylis:Plantago | 13 weeks | 6 | 0.5 | 0.5 | 9.8 | 8.6 | 4.4 | 9.5 | 11.5 | 16.1 | 2.3 |
Dactylis:Sanguisorba | 13 weeks | 6 | 0.5 | 0.5 | 9.8 | 4.4 | 6.0 | 1.9 | 11.5 | 3.6 | 4.4 |
Dactylis:Trifolium | 13 weeks | 6 | 0.5 | 0.5 | 9.8 | 13.3 | 7.5 | 7.7 | 11.5 | 14.2 | 2.6 |
Lotus:Plantago | 13 weeks | 6 | 0.5 | 0.5 | 6.3 | 8.6 | 1.0 | 16.2 | 4.7 | 16.1 | 8.6 |
Lotus:Sanguisorba | 13 weeks | 6 | 0.5 | 0.5 | 6.3 | 4.4 | 1.9 | 2.6 | 4.7 | 3.6 | –0.6 |
Lotus:Trifolium | 13 weeks | 6 | 0.5 | 0.5 | 6.3 | 13.3 | 2.4 | 9.2 | 4.7 | 14.2 | 4.4 |
Plantago:Sanguisorba | 13 weeks | 6 | 0.5 | 0.5 | 8.6 | 4.4 | 11.6 | 1.1 | 16.1 | 3.6 | 9.6 |
Plantago:Trifolium | 13 weeks | 6 | 0.5 | 0.5 | 8.6 | 13.3 | 10.2 | 6.9 | 16.1 | 14.2 | 3.3 |
Sanguisorba:Trifolium | 13 weeks | 6 | 0.5 | 0.5 | 4.4 | 13.3 | 1.9 | 13.4 | 3.6 | 14.2 | 5.3 |
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Species codes: Populus - Populus tremuloides, Picea - Picea glauca, Bromus - Bromus erectus, Dactylis - Dactylis glomerata, Lotus - Lotus corniculatus, Plantago - Plantago lanceolata, Sanguisorba - Sanguisorba minor, and Trifolium - Trifolium repens.
Partitioning net biodiversity effects (changes in aboveground biomass, g) with competitive and additive partitioning models using experimental data (Table 1) from a grassland diversity-productivity study (Mahaut et al., 2020).
Calculations are detailed in Supplementary file 1B.
Species mixture | Competitive partitioning | Additive partitioning | ||
---|---|---|---|---|
0.50Species1: 0.50Species2 | Positive effect | Competitive/ negative effect | Complementarity effect (CE) | Selection effect (SE) |
Bromus:Dactylis | 1.8 | 1.7 | 2.7 | 0.9 |
Bromus:Lotus | 1.2 | –0.1 | 1.1 | –0.1 |
Bromus:Plantago | 2.5 | 3.2 | 4.3 | 1.4 |
Bromus:Sanguisorba | 0.1 | 0.1 | 0.1 | 0.0 |
Bromus:Trifolium | 0.9 | 2.8 | 3.6 | 0.0 |
Dactylis:Lotus | 4.5 | 1.2 | 4.2 | 1.6 |
Dactylis:Plantago | 4.3 | 0.3 | 5.1 | –0.4 |
Dactylis:Sanguisorba | 0.0 | 0.8 | 0.3 | 0.5 |
Dactylis:Trifolium | 3.0 | 0.7 | 4.0 | –0.3 |
Lotus:Plantago | 7.6 | 2.3 | 7.9 | 1.9 |
Lotus:Sanguisorba | 0.0 | –0.9 | –0.6 | –0.3 |
Lotus:Trifolium | 0.0 | 1.8 | 0.8 | 1.1 |
Plantago:Sanguisorba | 1.5 | 4.7 | 3.9 | 2.3 |
Plantago:Trifolium | 5.0 | 1.2 | 7.8 | –1.6 |
Sanguisorba:Trifolium | 2.9 | 3.6 | 3.9 | 2.6 |
Average | 2.3 | Competitive: 1.7 Negative: –0.2 | 3.3 | 0.6 |
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Species codes: see Table 1.
Additional files
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MDAR checklist
- https://cdn.elifesciences.org/articles/98073/elife-98073-mdarchecklist1-v1.pdf
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Supplementary file 1
Simulated mixed trembling aspen and white spruce and experimental grassland mixtures.
(A) Simulated growth and yield of mixed trembling aspen (Populus tremuloides Michx.) and white spruce (Picea glauca [Moench] Voss) by different ages and stand compositions in western Canada. (B) Greenhouse experiment with grassland mixtures conducted by Mahaut et al., 2020.
- https://cdn.elifesciences.org/articles/98073/elife-98073-supp1-v1.xlsx