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Mensah, S., Glèlè Kakaï, R., Seifert, T., 2016. Patterns of biomass allocation be-
tween foliage and woody structure: the effects of tree size and specific functional
traits. Ann. For. Res. 59(1): 49-60.
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Ann. For. Res. 59(1): 49-60, 2016 Research article
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Mensah et al. Patterns of biomass allocation between foliage and woody structure...
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Ann. For. Res. 59(1): 49-60, 2016 Research article
Results
Figure 1 Distribution of leaf to wood mass ratio Within study species, the biomass allocated to
among studied species; CA - C. africana, foliage per unit of wood mass (LWR) ranged
CK - C. kraussii, CS - C. sylvaticus, SG from 0.0038 to 0.0225 for C. africana, 0.0071
- S. gerrardii, TD - T. dregeana, XM - X. to 0.0704 for C. kraussii, 0.0200 to 0.0916
monospora. Species with the same letter
for C. sylvaticus, 0.0073 to 0.1443 for S. ger-
are not significantly different at p = 0.05
(Student-Newman-Keuls test). rardii, 0.0113 to 0.0485 for T. dregeana and
0.0053 to 0.0503 for X. monospora (Figure
1). There were significant differences between
of LWR due to tree size by using the residu-
species (F = 13.4; P < 0.001). On average, C.
als of LWR-DBH model as response variable,
sylvaticus and T. dregeana showed the highest
and each specific trait as explanatory variable.
values of LWR, followed by C. kraussii and S.
Finally, we grouped the species and compared
gerrardii, while C. africana and X. monospora
the trend lines of LWR-dbh scaling relation-
had the same and lowest values (Figure 1).
ship between groups of species. We used the
Tree diameter and species identity showed
specific functional traits as a grouping factor,
significant effects (P < 0.001) on the biomass
thus distinguishing (1) between species with
allocation patterns, with 77.96 % of the total
larger and smaller leaf area, and (2) between
variance being explained (Table 2). Tree diam-
species with higher and lower wood density.
eter alone explained 44.44 % of the variance of
After grouping, we found that the range for
LWR for all species (P < 0.001). The effect of
tree diameter was greater in the group of high
tree size was shown by a significant decrease
wood density species (0.7- 94.5 cm). There-
in LWR with increasing tree diameter (Figure
fore, we excluded the largest trees from the set
2a), regardless of the species. Results from
of study species to have approximatively the
GLM also showed species’ significant effects
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Ann. For. Res. 59(1): 49-60, 2016 Research article
on the leaf to wood mass ratio (P < 0.001, Ta- lower wood density species (slope = -4.10-4,
ble 2), and a large variability of species-spe- Figure 3b). Accordingly, species with higher
cific slopes. For a given tree diameter, C. wood density had slightly lower LWR than
sylvaticus, T. dregeana, C. kraussii and S. ger- species with lower wood density. In contrast, a
rardii showed slopes of 1.27±0.14, 1.13±0.14, more remarkable differentiation was noted be-
0.81±0.14 and 0.53±0.14 respectively, higher tween species with larger leaf area and species
than the one in C. africana, which was consid- with smaller leaf area (Figures 3b): those with
ered as the baseline (Table 2). X. monospora larger leaf area had greater slope and intercept,
was ranked last, and had a slope that did not and therefore greater biomass allocated to foli-
differ significantly from the one in the baseline age per unit of wood biomass.
(P = 0.320).
Table 2 Output of GLMs describing the effects of tree diameter, species identity, wood density and leaf
area on leaf to wood mass ratio
Pseudo R
Estimate SE t P (>|t|) Deviance P (>Chi)
square (%)
Intercept -3.092 0.164 -18.896 <0.001
Log (DBH) -0.525 0.041 -12.769 <0.001 24.79 <0.001
Species 18.70 <0.001
C. kraussii 0.813 0.139 5.866 <0.001
77.96
C. sylvaticus 1.273 0.143 8.893 <0.001
S. gerrardii 0.533 0.141 3.777 <0.001
T. dregeana 1.126 0.143 7.866 <0.001
X. monospora 0.141 0.141 1.001 0.320
Intercept 1.387 0.403 3.444 <0.001
12.61
Wood density -2.672 0.767 -3.482 <0.001 4.11 <0.001
Intercept -0.464 0.093 -4.972 <0.001
30.17
Leaf area 0.011 0.002 6.131 <0.001 9.96 <0.001
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Mensah et al. Patterns of biomass allocation between foliage and woody structure...
Figure 2 Individual effects of (a) tree size (DBH), (b) leaf area and (c) wood density on leaf
to wood mass ratio. The regression coefficients (estimates) and values of pseudo
R-square are given in Table 2.
et al. 2002). Such a reduced production of foli- and heterospecific neighbours. In the mean-
age biomass is in part a result of the declining time, additional resources are invested for root
production of branch foliage when branches growth and for below ground mechanical safe-
get older (King 1997). This is intrinsically ty (Poorter et al. 2015). Therefore, increasing
linked with increasing amount of heartwood tree size would result in different mechanical
in stems and branches with increasing age and architectures that enable plants to differen-
in line with the findings that sapwood area is tially access the resources in the below and
highly correlated with total foliage biomass aboveground compartments (Fourcaud et al.
according to the pipe model theory (Shinoza- 2008). While our results support the hypothe-
ki et al. 1964a,b, Marchand 1984, Morataya et sis of size-related allocation patterns, the var-
al. 1999). The higher LWR in younger trees iance explained by tree diameter leaves much
indicates that more carbohydrate resources part of variation to be attributed to species-spe-
would have been allocated to foliage to under- cific differences (Weiner 2004, McCarthy et al.
take photosynthetic activities and allow rapid 2007, Poorter et al. 2015), and / or environ-
vertical growth. In natural environments (e.g. mental effects (McCarthy & Enquist 2007, Re-
natural forests), it is remarkable that resource ich et al. 2014).
partitioning among plant organs at early devel- The significant effect of species identity on
opment stages is part of plant’s strategy built the biomass partitioning patterns stresses the
as a response of competition for light. But as plasticity of different species in acquiring re-
plant size increases, more resources are allo- sources and adjusting biomass allocation. In-
cated for stem growth (height and diameter) deed, different species are expected to obtain
to enable plants to compete with conspecific resources in various ways because of the in-
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Ann. For. Res. 59(1): 49-60, 2016 Research article
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Mensah et al. Patterns of biomass allocation between foliage and woody structure...
tree species and enables them to fill forest can- to consider if we are to catch the total varia-
opy gaps quickly. On the contrary, in higher bility in the biomass allocation patterns. While
wood density species, the conductive tissues recent studies put forward the importance of
are most expensive to construct (Suzuki 1999), above and belowground resources available,
thus slowing tree growth. Furthermore, it was accounting for the role of plant functional
found that LWR declined with increasing tree traits that could capture the leaf and wood eco-
size for both higher and lower wood density nomics spectra, would provide deeper insights
species, consistently with what is expected into the mechanisms behind resource utilisa-
from the size dependency hypothesis. How- tion and biomass allocation.
ever, the fact that LWR declines more steeply
in higher wood density species than in lower
wood density species indicates that the latter Acknowledgements
allocates more biomass to foliage and less
biomass to stem and branches. This outcome This research was partly supported by the
suggests that additional resources have prob- SHARE Intra-ACP project and the National
ably been allocated to foliage to maximize Research Foundation of South Africa through
photosynthetic activities in low wood density the project Catchman Letaba in the RTF fund-
species. ing scheme of DST. Special thanks go to the
All being considered, it is worth mentioning colleagues Otto Pienaar and Andrew Perkins
that co-existing species show quite different who provided help during the data collection.
patterns of aboveground allocation and dif- We also acknowledge the anonymous referees
ferent correlations with structural traits. This for their significant contribution.
may serve as proof that competition in the
Mistbelt forests is interacting with tree struc-
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