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Tree diversity effect on dominant height in temperate forest Patrick Vallet, Thomas Pérot Irstea Nogent-sur-Vernisson CAQSIS, 28 29 March 2017, Bordeaux

2 Overyielding in mixed forest Context For many species associations, a mixture effect on productivity Mixture effect often studied on basal area Is height growth influenced by tree mixture? Is overyielding found on basal area correspond to: A real-overyielding? A different allocation of produced matter Study at the species level Species dominant height growth

3 Analysis method using NFI data GENERAL PRINCIPLE Step 1: Selection of a pure stand dataset in NFI data Selection of a mixed stand dataset in NFI data Step 2: Development of specific dominant height growth model in pure stand Hdom = f(age, climatic & edaphic variables) Step 3: Model applied in mixed stand dataset Expected dominant height (control) Calculation of mixture effects Mixture effect = Hdom observed - Hdom expected Hdom expected Analyses of the mixture effects

4 Species studied PURE STANDS Dominant height growth model developed for 5 target species Pure stand plots location

5 Species studied MIXED STANDS Selection in the same geographical area (SylvoEcoRégions) A SER is selected in mixed stand database only if at least 5 pure plots were used for pure stand model calibration

6 Species studied MIXED STANDS Mixture effect of a companion species on the target species Abies alba Mill. Acer pseudoplatanus L. Betula pendula Roth Carpinus betulus L. Castanea sativa Mill. Fagus sylvatica L. Fraxinus Excelsior L. Picea abies (L.) Karst. Pinus nigra Arn. laricio Pinus nigra Arn. nigra Pinus sylvestris L. Prunus avium L. Pseudotsuga menziesii (Mirb.) Franco Quercus robur L. Quercus petraea (Matt.) Liebl. Quercus pubescens Willd. Sessile oak - - 12 21 12 233 25 - - - 42 - - 123 - - Common beech 79 19-29 - - 55 35 - - 35 10-94 216 - Scots pine - - - - - 31-16 12 27 - - - 26 32 12 Silver fir - - - - - 64-89 - - 10 - - - 10 - Norway spruce 109 - - - - 43 - - - - 23-15 - - - Number of plots in mixed stands

7 Species studied MIXED STANDS Seven mixture associations Species association Number of plots Sessile oak Common beech 195 Sessile oak Scots pine 26 Common beech Silver fir 63 Common beech Norway spruce 35 Common beech Scots pine 27 Scots pine Norway spruce 14 Silver fir Norway spruce 87

8 Analysis method using NFI data GENERAL PRINCIPLE Step 1: Selection of a pure stand dataset in NFI data Selection of a mixed stand dataset in NFI data Step 2: Development of specific dominant height growth model in pure stand Hdom = f(age, climatic & edaphic variables) Step 3: Model applied in mixed stand dataset Expected dominant height (control) Calculation of mixture effects Mixture effect = Hdom observed - Hdom expected Hdom expected Analyses of the mixture effects

9 Development of dominant height growth model PURE STAND No height increment data in French NFI : only transversal data Use of a standard growth curve Need to include biophysical factors to take into account site fertility Asymptotic value a = a Hossfeld II model type Standard curve in literature Parcimonious: 3 parameters Flexible: can be adapted to many situations n 0,i + ak, i X k k = 1 hdom ~ a 1+ 1 b age d d a a/2 X k are environmental factors b

10 Example for sessile oak PURE STAND Environmental factors induced variations

11 Biophysical factors in the height models Factors Parameter Sessile oak European beech Intercept a 0 36.87 *** 33.81 *** 23.57 *** 23.28 *** 6.29 Scots pine Silver fir Norway Spruce Climate and water availability Soil characteristics Elevation (km) -8.69 *** -9.06 *** -8.17 *** 7.99 *** Elevation 2 (km 2 ) 3.62 *** -7.59 *** March min. temperature ( C) 1.17 *** Slope (%) -0.10 *** -0.0317 ** Soil depth (cm) 0.0496 *** Soil water holding capacity (mm) 0.27 ** 0.0490 *** 0.0317 *** 0.0303 *** July water balance (mm) 0.076 *** July water balance 2 (mm 2 ) -0.00055 *** July water deficit (mm) -0.049 *** Nitrogen index (Ellenberg) - a k 0.86 ** 1.01 *** 1.17 *** 0.89 *** ph 2 - -0.098 *** Limestone bedrock (boolean) -6.82 *** Siliceous bedrock (Boolean) 3.36 *** Light access Geo. residual effects Light index (Ellenberg) - -3.06 *** -0.82 * 9.17 ** Light index (Ellenberg) 2 - -1.02 *** Density index - 14.94 *** 4.90 *** 18.26 *** 2.96 ** Density index 2 - -7.06 *** -6.97 * Geographical Unit (Oceanic North-West) (boolean) -2.91 ** Geographical Unit (Massif Central mountains) (boolean) -1.85 ** Geographical Unit (Alps mountains) (boolean) -3.74 ** Age for ½ asymptotic value b 51.56 *** 37.91 *** 24.62 *** 36.21 *** 32.86 *** Shape factor c 1.14 *** 1.87 *** 1.86 *** 2.04 *** 1.96 ***

12 Analysis method using NFI data GENERAL PRINCIPLE Step 1: Selection of a pure stand dataset in NFI data Selection of a mixed stand dataset in NFI data Step 2: Development of specific dominant height growth model in pure stand Hdom = f(age, climatic & edaphic variables) Step 3: Model applied in mixed stand dataset Expected dominant height (control) Calculation of mixture effects Mixture effect = Hdom observed - Hdom expected Hdom expected Analyses of the mixture effects

13 Mean mixture effects on target species MIXED STANDS EXAMPLE FOR OAK AND SPRUCE Effects can be negative, null or positive

14 Mean mixture effects by species associations MIXED STANDS Effects can be negative, null or positive Species association Species 1 Species 2 Number of plots Effect of species 2 on species 1 Effect of species 1 on species 2 Sessile oak Common beech 195 +2.0% - 5.4% Sessile oak Scots pine 26 +1.3% +6.3% Common beech Silver fir 63 + 4.8% +3.2% Common beech Norway spruce 35 + 7.7% - 7.8% Common beech Scots pine 27 +2.8% + 16.7% Scots pine Norway spruce 14 + 11.6% -6.5% Silver fir Norway spruce 87 + 3.3% - 3.9% Significant positive value Significant negative value

15 Mixture effect depends on height difference between species Species on the top: Negative effect Species on the bottom: Positive effect Oak pine case: Close H dom dynamics no mixture effect Beech Spruce case: Differentiated H dom dynamics mixture effect Each species reach the other one

16 Generalization of the result ALL SPECIES ASSOCIATIONS TOGETHER When the associated species is smaller, the mixture effect is negative When the associated species is taller, the mixture effect is positive

17 Comparison with results on basal area MAUDE TOÏGO S PHD : 5 COUPLES OF SPECIES Beech 129 Sessile Oak 334 Mixture 469 Scots pine 100 Sessile oak 334 Mixture 107 Beech 138 Spruce 257 Mixture 117 Beech 138 Silver fir 172 Mixture 246 Spruce 257 Silver fir 172 Mixture 292

18 Mean mixture effect MIXTURE EFFECT ON BASAL AREA Species level Diversity effect (For a 50% - 50% mixture) +60% +40% +20% 0 beech fir oak pine beech spruce fir spruce beech oak

19 Comparison of height and basal area mixture effects CAUTION: Only qualitative comparison Volume ~ α x G x H Mixture effect is not a compensation between the compartments but a real effect on volume

20 Conclusion Mixture effect on dominant height depends on height differences between species Leveling process: The taller species limits its height growth The smaller species increases its height growth Lower magnitude compared to effects on basal area From -7.8% to +16.7% for dominant height Basal area effects from -5.8% to 50.8% (Toïgo et al. 2015), and usually in the same direction Real overyielding, and not a compensation between radial and height productivity