For Peer Review
Diversity increases ca
rbon storage and tree productivity in
Spanish forests
Journal:
Global Ecology and Biogeography
Manuscript ID:
GEB-2012-0248.R3
Manuscript Type:
Research Papers
Date Submitted by the Author:
n/a
Complete List of Authors:
Ruiz-Benito, Paloma; INIA,CIFOR, Dpt. Forest Ecosystems and Resources;
Universidad de Alcalá, Departamento de Ciencias de la Vida
Gómez-Aparicio, Lorena; CSIC, Instituto de Recursos Naturales y
Agrobiología de Sevilla (IRNAS)
Paquette, Alain; UQAM, CEF
Messier, Christian; UQAM, CEF
Kattge, Jens; Max-Planck, Institute for Biogeochemistry
de Zavala, Miguel; Universidad de Alcalá, Departamento de Ciencias de la
Vida
Keywords:
climate, competition, ecosystem functions and services, forest
management, functional diversity, functional identity, maximum likelihood
techniques, national forest inventory
Global Ecology and Biogeography
For Peer Review
Ruiz-Benito et al. 1
Diversity increases carbon storage and tree productivity in Spanish
1
forests
2
Paloma Ruiz-Benito
1,2,3*
, Lorena Gómez-Aparicio
4
, Alain Paquette
3
, Christian 3
Messier
3
, Jens Kattge
5
, Miguel A. Zavala
2
4
1
CIFOR-INIA. Ctra. de la Coruña, Km. 7,5. 28040. Madrid, Spain. 5
2
Forest Ecology and Restoration Group, Department of Life Sciences, Science Building, 6
University of Alcala, Campus Universitario, 28871 Alcalá de Henares, Madrid, Spain. 7
3
CEF, Universit du Quebec à Montréal au Canda. 8
4
Instituto de Recursos Naturales y Agrobiología de Sevilla (IRNAS), CSIC, PO BOX 9
1052. 41080. Sevilla. Spain 10
5
Max Planck Institute for Biogeochemistry, 07745 Jena, Germany. 11
*
Correspondence: Paloma Ruiz Benito. Forest Ecology and Restoration Group, 12
Department of Life Sciences, Science Building, University of Alcala, Campus 13
Universitario, 28871 Alcalá de Henares, Madrid, Spain. E-mail: 14
palomaruizbenito@gmail.com 15
Article type: Research Paper. 16
Running title: Diversity effects on forest carbon storage and productivity.
17
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ABSTRACT
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Aim Biodiversity loss could reduce primary productivity and carbon storage provided 19
by forests; however the underpinning mechanisms of biodiversity effects on multiple 20
ecosystem functions are not completely understood. Spanish forests are of particular 21
interest because of the broad variation in environmental conditions and management 22
history. We tested for the existence of a relationship among diversity effects and both 23
carbon storage and tree productivity, and examined the relative importance of 24
complementarity and selection mechanisms in a wide variety of forests, from cold 25
deciduous Atlantic to xeric Mediterranean evergreen forests. 26
Location Continental Spain. 27
Methods We used c. 54,000 plots of the Spanish Forest Inventory and maximum 28
likelihood techniques to quantify how climate, stand structure and diversity shape 29
carbon storage and tree productivity. Diversity effects included both complementarity 30
and selection mechanisms, measured respectively through functional diversity and 31
functional identity measures. 32
Results Diversity had a significant effect on both carbon storage and tree productivity, 33
even when controlling for climatic and stand structural confounding factors. A 34
consistent positive effect of functional diversity on carbon storage and tree productivity 35
was observed in all seven forest types studied. This relationship was not linear, and the 36
largest changes in carbon storage and tree productivity were observed at low functional 37
diversity levels. However, the importance of complementarity effects was not consistent 38
with the productivity of different forest types. Selection effects were particularly 39
important in deciduous and Mediterranean pine forests, but had very little effect on 40
mountain pines. 41
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Main conclusions We found a generally positive effect of diversity on carbon storage 42
and tree productivity, supported by both complementarity and selection mechanisms. 43
Thus, both functionally diverse forests and functionally important species should be 44
maintained to adequately preserve and promote key ecosystem functions such as carbon 45
storage and tree productivity. 46
Keywords: climate, competition, community-weighted means of trait value, continental 47
Spain, diversity, ecosystem functions and services, forest management, maximum 48
likelihood techniques, national forest inventory. 49
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INTRODUCTION
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Under global change the increase of human-mediated modifications in ecosystems could 51
lead to important biodiversity losses (Cardinale et al., 2012). Reductions of biodiversity 52
may alter the quality and quantity of ecosystem functions and services provided by 53
terrestrial ecosystems (e.g. Isbell et al., 2011). Thus, biodiversity-ecosystem functioning 54
(BEF) relationships are an important topic in ecology and have been subject of 55
considerable debate during the last decades (e.g. Loreau et al., 2001; Hooper et al., 56
2012). Most studies of BEF relationships have used species richness as a measure of 57
diversity (e.g. Maestre et al., 2012). However, it has recently been shown that functional 58
diversity better connects the underlying mechanisms of biodiversity effects to 59
ecosystem functioning (e.g. Hooper et al., 2005). Trait based approaches are a 60
promising avenue to disentangle the underlying mechanisms of the diversity effects on 61
productivity (see Mokany et al., 2008; Paquette & Messier, 2011; Roscher et al., 2012). 62
Two main non-exclusive mechanisms of the positive effects of diversity on 63
ecosystem functioning have been proposed: the complementarity and the selection 64
effects (Grime, 1998; Loreau & Hector, 2001). The complementarity effect increases an 65
ecosystem function through facilitation and niche partitioning, because functionally 66
diverse species assemblages would enhance resource use efficiency and nutrient 67
retention (Loreau, 2000; Morin et al., 2011). Some authors have suggested that 68
complementarity effects could be particularly important in low productive or harsh 69
environments, where species interactions are less affected by competitive exclusion 70
(Warren et al., 2009; Paquette & Messier, 2011). Yet, other authors have observed that 71
complementarity effects are similar across different forest biomes (Zhang et al., 2012). 72
The selection effect (i.e. selection of particular species or functional traits) proposes that 73
high species richness increases the probability of including the most productive species 74
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