Lehnebach1,2 H

Wood specific gravity variations within tree trunk: the case study of Legumes representatives in French Guiana

Romain LEHNEBACH1,2 H. MOREL1, N. AMUSANT3 , S. GRIFFON1, J.F. BARCZI1, J. BEAUCHÊNE3 and E. NICOLINI1 1JRU AMAP : botAny and bioinforMatics of Plant Architecture, Montpellier, France 2University of Montpellier II, Montpellier, France 3JRU EcoFoG : Wood sciences laboratory, Kourou, French Guiana

Wood sciences underpinning tropical forest ecology & management, Tervuren, Belgium, May 2015 Introduction:

probability to sample species with coloured ? heartwood

‘bottom’ ‘top’ wood wood ‐ Longitudinal trend

Introduction WSUFEM, Tervuren, Belgium, May 2015 Objectives

1. How WSG varies radially and longgyitudinally within tree and between s pecies?

2. Wha t are the tren ds o f WSG ra dia l var ia tions an d w ha t are the poss ible m is in terpre ta tions of these trends due to the effect of heartwood?

3. Does these variations and patterns, are related to the successional status of the species ?

Objectives WSUFEM, Tervuren, Belgium, May 2015 Experimental site:

Paracou field station

Suriname

Guyana

Brasil 100 km (AMAPA)

400 m

30 years of monitoring Access to growth rate, mortality rate ..

Study site WSUFEM, Tervuren, Belgium, May 2015 Data collection: Field sampling and WSG measurements

top-crown height

mid-crown height

under crown

tangential cut every 0.5cm

4 m Basi c WSG = Oven dry weight Volume

10 < DBH < 15 cm 0.5 m 1 to 3 iidiidlndividual/species

Field sampling & Measurements WSUFEM, Tervuren, Belgium, May 2015 Statistical analysis - Radial and Longitudinal gradient assessed by:

Under crown µ(WSG near pith 0.5m) new/oldradial = µ(WSG below bark 0.5m) > 1  WSG increases on the radius < 1  WSG decreases on the radius

µ(WSG near pith under crown) old/oldheight = µ(WSG near pith 0.5m) old/oldheight > 1  WSG increases with height < 1  WSG decreases with height

050.5m new/oldradial

- Radial trends assessed by linear mixed model (lme) :

linear: WSG ~ β0 + β1*radial distance ,

quadratic: WSG ~ β0 + β1*radial distance + β3*radial distance²

Statistical analysis WSUFEM, Tervuren, Belgium, May 2015 Collected species: phylogeny & overall WSG Swartzia panacoco Bocoa prouacensis Enterolobium schomburgkii Inga stipularis Zygia tetragona Abarema jupunba PkiitidParkia nitida Parkia velutina Parkia pendula Tachigali guianensis Recordoxylon speciosum Vouacapoua americana Dicorynia guianensis Eperua falcata Eperua grandiflora Peltogyne venosa

0.2 0.4 0.6 0.8 1.0 Overall WSG Faboideae Mimosoideae Caesalpinioideae Collected Species WSUFEM, Tervuren, Belgium, May 2015 Defining successional status

Using Paracou database : computation of variables related to successional status

- RGR95: (95th quantile of the growth rate distribution)

- Mr: morta lity rate:

-DBHmax

-4 -3 -2 -1 0 1 2 PC1 Axis (71.7%)

PC1 scores uses as synthetic variable describing successional status

Assessment of ecological behaviour WSUFEM, Tervuren, Belgium, May 2015 Taxon new/oldradial old/oldheight Results: Radial & longitudinal gradients Parkia nitida 1,977 *** 2,212 * Parkia nitida 2,073 *** 1,959 *** ‐ *** / * Parkia nitida 2,549 *** 1,455 * PN Parkia pendula 1,357 *** 1,445 *** Parkia velutina 1,537 *** 1,198 ns 2.0 Parkia velutina 1,320 *** 1,212 *** PN*** / *** Tachigali guianensis 1,090 * 1,243 * Parkia pendula 1,276 *** 1,215 ** Parkia pendula 1,307 ** 1,115 ns crease Tachigali guianensis 1,226 *** 1,118 ns nn Abarema jupunba 1,204 *** 0,838 * Dicorynia guianensis 1,042 * 0,770 *** Enterolobium schomburgkii 1,300 *** 1,146 *** 1.5 *** / * dinal i *** / *** Inga stiplularis 1,229 *** 0,976 ns PP PN Dicorynia guianensis 1,038 ns 0,795 ** uu Dicorynia guianensis 1,151 *** 0,790 *** Eperua falcata 0,913 ns 0,788 *** * / * TG Eperua falcata 0,836 *** 0,821 ** *** / ***** / *** PP PV *** / ns Longit PV SG Eperua falcalta 0,968 ns 0,852 ** *** / *** Eperua grandiflora 0,832 *** 0,781 *** ES WW *** / ns ** / ns Inga stipularis 1,358 *** 1,052 ns TG PP

*** / ns Eperua grandiflora 0,851 *** 0,796 *** IS Recordoxylon speciosum 1,105 *** 0,810 ***

1.0 *** / * Mean *** / ns ZT *** / * Vouacapoua americana 0,967 ** 0,919 ** ZT IS al

Swartzia panacoco 1,230 *** 0,904 *** *** / ** ** / ** *** / * nn SP

ee SP VA *** / *** Recordoxylon speciosum 1,181 *** 0,840 ** SP *** / *** RS Eperua grandiflora 0,941 0,858 ** / *** ** *** EG ns / ** EF *** ***/ ** / * RSAJ Recordoxylon speciosum 1,079 *** 0,881 *** *** / ** * / *** EF BP*** / *** *** / *** *** / ***BP*** / *** RS *** / ***PVe ns / ** Swartzia panacoco 1,209 *** 0,921 * EGPVe *** / ***ns / *** DG *** / *** *** / ***BP ***EF / *** DG EG PVe * / ***

Swartzia panacoco 1,168 *** 0,924 ** ngitudi DG decreas

Peltogyne sp. 0,886 *** 0,781 *** oo

Peltogyne sp. 0,906 *** 0,803 *** L 1.0 1.5 2.0 2.5 Peltogyne sp. 0,856 *** 0,800 *** Bocoa prouacensis 0,913 * 0,819 *** Bocoa prouacensis 0,884 *** 0,790 *** Radial Radial increase Bocoa prouacensis 0,908 *** 0,808 *** decrease Zygia tetragona 0,896 *** 0,968 * + Zygia tetragona 0,8853 *** 0,984 * Radial & Longitudinal gradients WSUFEM, Tervuren, Belgium, May 2015 Taxon new/oldradial old/oldheight Results: Radial & longitudinal gradients Parkia nitida 1,977 *** 2,212 * Parkia nitida 2,073 *** 1,959 *** ‐ *** / * Parkia nitida 2,549 *** 1,455 * PN Parkia pendula 1,357 *** 1,445 *** Parkia velutina 1,537 *** 1,198 ns 2.0 *** / *** Parkia velutina 1,320 *** 1,212 *** PN Tachigali guianensis 1,090 * 1,243 * Parkia pendula 1,276 *** 1,215 ** crease

Parkia pendula 1,307 ** 1,115 ns nn Tachigali guianensis 1,226 *** 1,118 ns Abarema jupunba 1,204 *** 0,838 * Dicorynia guianensis 1,042 * 0,770 *** 1.5 *** / * Enterolobium schomburgk 1,300 *** 1,146 *** dinal i *** / *** PP PN

Inga stiplularis 1,229 *** 0,976 ns uu Dicorynia guianensis 1,038 ns 0,795 ** Dicorynia guianensis 1,151 *** 0,790 ***

* / * Eperua falcata 0,913 ns 0,788 *** TG *** / ***** / *** PP PV *** / ns Eperua falcata 0,836 *** 0,821 ** Longit PV SG

Eperua falcalta 0,968 ns 0,852 ** *** / *** ES WW *** / ns ** / ns Eperua grandiflora 0,832 *** 0,781 *** TG PP Inga stipularis 1,358 *** 1,052 ns *** / ns IS Eperua grandiflora 0,851 *** 0,796 ***

1.0 *** / * Mean Recordoxylon speciosum 1,105 *** 0,810 *** *** / ns ZT *** / * ZT IS

Vouacapoua americana 0,967 ** 0,919 ** al *** / ** ** / ** *** / * nn SP ee VA SP Swartzia panacoco 1,230 *** 0,904 *** *** / *** SP *** / *** Recordoxylon speciosum 1,181 *** 0,840 ** RS ** / *** ns / ** EG EF Eperua grandiflora 0,941 0,858 *** ***/ ** / * ** *** RSAJ *** / ** * / *** EF BP*** / *** *** / *** Recordoxylon speciosum 1,079 *** 0,881 *** *** / ***BP*** / *** RS *** / ***PVe ns / ** EGPVe *** / ***ns / *** DG *** / *** *** / ***BP ***EF / *** DG Swartzia panacoco 1,209 *** 0,921 * EG PVe * / ***

ngitudi DG Swartzia panacoco 1,168 *** 0,924 ** decreas oo

Peltogyne sp. 0,886 *** 0,781 *** L 1.0 1.5 2.0 2.5 Peltogyne sp. 0,906 *** 0,803 *** Peltogyne sp. 0,856 *** 0,800 *** Bocoa prouacensis 0,913 * 0,819 *** Radial Radial increase Bocoa prouacensis 0,884 *** 0,790 *** Bocoa prouacensis 0,908 *** 0,808 *** decrease Zygia tetragona 0,896 *** 0,968 * + Zygia tetragona 0,8853 *** 0,984 * Radial & Longitudinal gradients WSUFEM, Tervuren, Belgium, May 2015 Taxon new/oldradial old/oldheight Results: Radial & longitudinal gradients Parkia nitida 1,977 *** 2,212 * Parkia nitida 2,073 *** 1,959 *** ‐ *** / * Parkia nitida 2,549 *** 1,455 * PN Parkia pendula 1,357 *** 1,445 *** Parkia velutina 1,537 *** 1,198 ns 2.0 *** / *** Parkia velutina 1,320 *** 1,212 *** PN Tachigali guianensis 1,090 * 1,243 * Parkia pendula 1,276 *** 1,215 ** crease

Inga stiplularis 1,229 *** 0,976 ns uu Dicorynia guianensis 1,038 ns 0,795 ** Dicorynia guianensis 1,151 *** 0,790 ***

Eperua falcata 0,913 ns 0,788 *** * / * TG *** / ***** / *** PP PV *** / ns Eperua falcata 0,836 *** 0,821 ** Longit PV SG

Eperua falcalta 0,968 ns 0,852 ** *** / *** ES WW *** / ns ** / ns Eperua grandiflora 0,832 *** 0,781 *** TG PP Inga stipularis 1,358 *** 1,052 ns *** / ns Eperua grandiflora 0,851 *** 0,796 *** IS

Recordoxylon speciosum 1,105 *** 0,810 *** 1.0 *** / * Mean *** / ns ZT *** / * ZT IS

Vouacapoua americana 0,967 ** 0,919 ** al *** / ** ** / ** *** / * nn SP

ee SP StiSwartzia panacoco 1,230 *** 0,904 *** VA *** / *** SP *** / *** Recordoxylon speciosum 1,181 *** 0,840 ** RS ** / *** EG ns / ** Eperua grandiflora 0,941 ** 0,858 *** EF *** ***/ ** / * RSAJ *** / ** * / *** Recordoxylon speciosum 1,079 *** 0,881 *** EF BP*** / *** *** / *** *** / ***BP*** / *** RS *** / ***PVe ns / ** EGPVe *** / ***ns / *** DG *** / *** *** / ***BP ***EF / *** DG Swartzia panacoco 1,209 *** 0,921 * EG PVe * / ***

ngitudi DG Swartzia panacoco 1,168 *** 0,924 ** decreas oo Peltogyne sp. 0,886 *** 0,781 *** L 1.0 1.5 2.0 2.5 Peltogyne sp. 0,906 *** 0,803 *** Peltogyne sp. 0,856 *** 0,800 *** Bocoa prouacensis 0,913 * 0,819 *** Radial Radial increase Bocoa prouacensis 0,884 *** 0,790 *** Bocoa prouacensis 0,908 *** 0,808 *** decrease Zygia tetragona 0,896 *** 0,968 * + Zygia tetragona 0,8853 *** 0,984 * Radial & Longitudinal gradients WSUFEM, Tervuren, Belgium, May 2015 Taxon new/oldradial old/oldheight Results: Radial & longitudinal gradients Parkia nitida 1,977 *** 2,212 * Parkia nitida 2,073 *** 1,959 *** ‐ *** / * Parkia nitida 2,549 *** 1,455 * PN Parkia pendula 1,357 *** 1,445 *** Parkia velutina 1,537 *** 1,198 ns 2.0 *** / *** Parkia velutina 1,320 *** 1,212 *** PN Tachigali guianensis 1,090 * 1,243 * Parkia pendula 1,276 *** 1,215 ** crease

Inga stiplularis 1,229 *** 0,976 ns uu Dicorynia guianensis 1,038 ns 0,795 ** Dicorynia guianensis 1,151 *** 0,790 ***

Eperua falcata 0,913 ns 0,788 *** * / * TG *** / ***** / *** PP PV *** / ns Eperua falcata 0,836 *** 0,821 ** Longit PV SG

Eperua falcalta 0,968 ns 0,852 ** *** / *** ES WW *** / ns ** / ns Eperua grandiflora 0,832 *** 0,781 *** TG PP Inga stipularis 1,358 *** 1,052 ns *** / ns Eperua grandiflora 0,851 *** 0,796 *** IS

Recordoxylon speciosum 1,105 *** 0,810 *** 1.0 *** / * Mean *** / ns ZT *** / * ZT IS

Vouacapoua americana 0,967 ** 0,919 ** al *** / ** ** / ** *** / * nn SP

*** / * PN 2.0 PN*** / *** crease nn 1.5 *** / * dinal i *** / *** PP PN uu

* / * TG *** / ***** / *** PP PV *** / ns Longit PV

*** / *** ES *** / ns ** / ns TG PP

*** / ns IS

1.0 *** / * *** / ns ZT *** / * ZT IS al

*** / ** ** / ** *** / * nn SP

ee SP VA *** / *** SP *** / *** RS ** / *** EG ns / ** EF *** ***/ ** / * RSAJ *** / ** * / *** EF BP*** / *** *** / *** *** / ***BP*** / *** RS *** / ***PVe ns / ** EGPVe *** / ***ns / *** DG *** / *** *** / ***BP ***EF / *** DG EG PVe * / ***

ngitudi DG decreas oo

L 1.0 1.5 2.0 2.5

-4 -3 -2 -1 0 1 Radial Radial increase PCA Scores Early decrease succesional ------> Late succesional

Radial & Longitudinal trends WSUFEM, Tervuren, Belgium, May 2015 Radial trends Radial WSG

0.2 0.4 0.6 0.8 1.0 heartwood/sapwood sapwood td ht h ear 123456 t woo Distance from pith (cm) d Results: Radialtrends(1) Parkia nitida ovxcriierincrease curvilinear Convex WSUFEM, Tervuren, Belgium, May 2015

Radial trends Radial WSG

0.2 0.4 0.6 0.8 1.0 heartwood/sapwood sapwood td ht h ear 123456 t woo Distance from pith (cm) d ioyi guianensis Dicorynia Results: Radialtrends(1) Parkia nitida ovxcriierincrease curvilinear Convex WSUFEM, Tervuren, Belgium, May 2015 ierincrease Linear

Radial trends Radial WSG

0.2 0.4 0.6 0.8 1.0 RdlR ecor heartwood/sapwood sapwood td ht h ear 123456 t woo d oxy Distance from pith (cm) d l nspec on ioyi guianensis Dicorynia i osum Results: Radialtrends(1) Parkia nitida ovxcriierincrease curvilinear Convex increase Concave curvilinear WSUFEM, Tervuren, Belgium, May 2015 ierincrease Linear

Results: Radial trends (2) Eperua grandiflora

sapwood heartwood/sapwood Heartwood Specific gravity corrected by 20% Extractives content

0 heartwood 1. 0.9 SG WS 0.8 0.7 0.6 12345601234560123456 Distance from pith (cm) Distance from pith (cm) Distance from pith (cm)

Radial trends WSUFEM, Tervuren, Belgium, May 2015 Links between radial trends and successional status

b 0 1.

b 88 0. SG WW

0.6 ab

0.4 a 0.2

-4 -3 -2 -1 0 1 246 PCA Scores Distance from pith Early succesional ------> Late succesional

Discussion & Conclusion WSUFEM, Tervuren, Belgium, May 2015 Links between radial trends and successional status

Adapted from

Woodcock & Shier, 2002 .2 1.0 1 0.8 0.4 0.6 0.2 0.0 corrected WSG

WSUFEM, Tervuren, Belgium, May 2015 Links between radial trends and successional status

Adapted from

Woodcock & Shier, 2002 .2

Very late

1.0 1 successional (Sciaphilic)

0.8 Late successional Early succesional 0.4 0.6 Pionneers 0.2

Inner OtOuter wood wood 0.0 corrected Successional Radial trend WSG status

Discussion & Conclusion WSUFEM, Tervuren, Belgium, May 2015 Links between radial trends and successional status

Adapted from

Woodcock & Shier, 2002 .2

Very late

1.0 1 successional (Sciaphilic)

0.8 Late successional Early succesional 0.4 0.6 Pionneers 0.2

Inner OtOuter BttBottom Top wood wood wood wood 0.0 corrected Successional Radial trend Longitudinal WSG status trend

WSUFEM, Tervuren, Belgium, May 2015 Conclusion 1. Radial and longitudinally variations ?

- Increase or constant WSG along the radius but NO radial decrease ! - Decreasing WSG from bottom to top is a general case, excepted for species with low WSG. - All studied species tend to the same range of WSG values with height (~ 0.6-0.9) ! Supported by a higher WSG under bark within trunk

2. Radial trends of WSG // the effect of heartwood?

- Po ten tia l decreasi ng tren ds are due to hear twoo d e ffec t in our sampl es.

3. Does these variations and patterns, are related to the successional status of the species ?

- Extreme radial increase & slight increase with height = pioneers species. - Radial increase & longitudinal decrease = late successional - Sciaphilic species differs by their constant WSG across the radius

Discussion & Conclusion WSUFEM, Tervuren, Belgium, May 2015 Xplo Software to visualize and compute biomass estimates ..

Parkia nitida Swartzia panacoco Biomass estimate by Xplo 16.8 Kg 73.23Kg Biomass estimate by WSG under 25.9 Kg 85.1 Kg bark at 1m30

Over-estimation 35% Over-estimation 14% WSUFEM, Tervuren, Belgium, May 2015