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    'Staff publications' contains references to publications authored by Wageningen University staff from 1976 onward.

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Author Correction: Climatic controls of decomposition drive the global biogeography of forest-tree symbioses
Steidinger, B.S. ; Crowther, T.W. ; Liang, J. ; Nuland, M.E. Van; Werner, G.D.A. ; Reich, P.B. ; Nabuurs, G.J. ; de-Miguel, S. ; Zhou, M. ; Picard, N. ; Herault, B. ; Zhao, X. ; Zhang, C. ; Routh, D. ; Peay, K.G. ; Abegg, Meinrad ; Adou Yao, C.Y. ; Alberti, Giorgio ; Almeyda Zambrano, Angelica ; Alvarez-Davila, Esteban ; Alvarez-Loayza, Patricia ; Alves, Luciana F. ; Ammer, Christian ; Antón-Fernández, Clara ; Araujo-Murakami, Alejandro ; Arroyo, Luzmila ; Avitabile, Valerio ; Aymard, Gerardo ; Baker, Timothy ; Bałazy, Radomir ; Banki, Olaf ; Barroso, Jorcely ; Bastian, Meredith ; Bastin, Jean Francois ; Birigazzi, Luca ; Birnbaum, Philippe ; Bitariho, Robert ; Boeckx, Pascal ; Bongers, Frans ; Bouriaud, Olivier ; Brancalion, Pedro H.H.S. ; Decuyper, Mathieu ; Hengeveld, Geerten ; Herold, Martin ; Lu, Huicui ; Parren, Marc ; Poorter, Lourens ; Schelhaas, Mart Jan ; Sheil, Douglas ; Zagt, Roderick - \ 2019
Nature 571 (2019)7765. - ISSN 0028-0836

In this Letter, the middle initial of author G. J. Nabuurs was omitted, and he should have been associated with an additional affiliation: ‘Forest Ecology and Forest Management Group, Wageningen University and Research, Wageningen, The Netherlands’ (now added as affiliation 182). In addition, the following two statements have been added to the Supplementary Acknowledgements. (1): ‘We would particularly like to thank The French NFI for the work of the many field teams and engineers, who have made extraordinary efforts to make forest inventory data publicly available.’ (1): ‘Sergio de Miguel benefited from a Serra- Húnter Fellowship provided by the Generalitat of Catalonia.’ Finally, the second sentence of the Methods section should have cited the French NFI, which provided a national forestry database used in our analysis, to read as follows: ‘The GFBi database consists of individual-based data that we compiled from all the regional and national GFBi forest-inventory datasets, including the French NFI (IGN—French National Forest Inventory, raw data, annual campaigns 2005 and following, https://inventaire-forestier.ign.fr/spip.php?rubrique159, site accessed on 01 January 2015)’. All of these errors have been corrected online.

Species mixing effects on forest productivity : A case study at stand-, species- and tree-level in the Netherlands
Lu, Huicui ; Mohren, Godefridus M.J. ; Río, Miren del; Schelhaas, Mart-Jan ; Bouwman, Meike ; Sterck, Frank J. - \ 2018
Forests 9 (2018)11. - ISSN 1999-4907
Basal area increment - Inter- and intra-specific competition - Mixed forest - Overyielding - Periodic annual volume increment

Many monoculture forests have been converted to mixed-species forests in Europe over the last decades. The main reasons for this conversion were probably to increase productivity, including timber production, and enhance other ecosystem services, such as conservation of biodiversity and other nature values. This study was done by synthesizing results from studies carried out in Dutch mixed forests compared with monoculture stands and evaluating them in the perspective of the current theory. Then we explored possible mechanisms of higher productivity in mixed stands, in relation to the combination of species, stand age and soil fertility, and discussed possible consequences of forest management. The study covered five two-species mixtures and their corresponding monoculture stands from using long-term permanent forest plots over multiple decades as well as two inventories (around 2003 and 2013) across the entire Netherlands. These forest plot data were used together with empirical models at total stand level, species level and tree level. Overyielding in Douglas-fir-beech and pine-oak mixtures was maintained over time, probably owing to the intensive thinning and was achieved on the poorer soils. However, this overyielding was not always driven by fast-growing light-demanding species. On individual tree level, intra-specific competition was not necessarily stronger than inter-specific competition and this competitive reduction was less seen at lower soil fertility and dependent on species mixtures. Moreover, size-asymmetric competition for light was more associated with tree basal area growth than size-symmetric competition for soil resources. Overall, this study suggests a substantial potential of species mixing for increasing productivity and implies developing forest management strategies to convert monospecific forests to mixed-species forests that consider the complementarity in resource acquisition of tree species.

Species and soil effects on overyielding of tree species mixtures in the Netherlands
Lu, Huicui ; Condés, Sonia ; Río, Miren del; Goudiaby, Venceslas ; Ouden, Jan den; Mohren, Godefridus M.J. ; Schelhaas, Mart Jan ; Waal, Rein de; Sterck, Frank J. - \ 2018
Forest Ecology and Management 409 (2018). - ISSN 0378-1127 - p. 105 - 118.
Niche complementarity - Productivity - Soil - Species mixing effect - Volume growth

A growing number of studies provides evidence that mixed-species forests often have higher stand productivity than monospecific forests, which is referred to as overyielding. In this study, we explored how the combination of species and soil conditions affect overyielding in terms of periodic annual volume increment (PAIV) in Dutch forests. We studied Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), common beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.), pedunculate oak (Quercus robur L.), and silver birch (Betula pendula Roth) growing in four two species combinations (Douglas-fir–common beech, Scots pine–pedunculate oak, pedunculate oak–common beech, and pedunculate oak–silver birch) from 398 long-term permanent field plots all over the Netherlands. We found that the Douglas-fir–common beech and Scots pine–pedunculate oak mixtures always showed overyielding. This overyielding was largely attributed to the Douglas-fir in the former mixture and to the pedunculate oak in the latter mixture, respectively. In both cases, overyielding was stronger at poor soils than at rich soils. The pedunculate oak–common beech mixtures overyielded at poor soils and underyielded at rich soils, which was attributed to the response of the common beech. Overyielding was not observed for the pedunculate oak–silver birch mixtures, irrespective of soil conditions. The results do not support our hypothesis since overyielding was not always driven by fast-growing light-demanding species. Overyielding was stronger for evergreen–deciduous species combinations, suggesting that differences in leaf phenology are a major driver of overyielding. Secondly, our results imply that overyielding is much stronger at poor soils than at rich soils, which is in line with the prediction of the stress-gradient hypothesis. We conclude that the growth of one species benefits from the admixture species, particularly in evergreen–deciduous species mixtures and that soils affect the extent of overyielding as studied in the Netherlands.

Species mixing effects on forest productivity in the Netherlands
Lu, Huicui - \ 2017
Wageningen University. Promotor(en): G.M.J. Mohren, co-promotor(en): F.J. Sterck. - Wageningen : Wageningen University - ISBN 9789463436397 - 134
mixed forests - temperate zones - deciduous forests - soil fertility - light - yields - netherlands - gemengde bossen - gematigde klimaatzones - loofverliezende bossen - bodemvruchtbaarheid - licht - opbrengsten - nederland

Many monoculture forests (dominated by a single tree species) have been converted to mixed-species forests (dominated by more than one tree species) in Europe over the last decades. The main reason for this conversion was to increase productivity, including timber production, and enhance other ecosystem services, such as conservation of biodiversity and other nature values. In addition, it has been suggested that mixed-species forests are more resistant, resilient and stable to disturbances.

In line with the niche complementarity hypothesis, inter-specific differences in crown architecture, leaf phenology, shade tolerance and root distribution may allow tree species to partition resources in mixed forests. Such mechanisms may lead to a higher productivity of mixed forests versus monoculture forests, a phenomenon often referred to as overyielding. Interestingly, the stress-gradient hypothesis and the resource-ratio hypothesis suggests that such inter-specific interactions vary along a soil fertility gradient, but in different ways. The stress-gradient hypothesis emphasizes that more efficient partitioning increases overyielding at low fertility soils, whereas the resource ratio hypothesis considers that the denser packing of crowns on fertile soils allows for partitioning of light and overyielding on high fertility soils. Several studies have been carried out about species mixing effects on forest productivity, but so far their findings are ambiguous. Probably, this ambiguity comes from the sites that they studied, which differ in species, age, management history, and/or environmental conditions.

This thesis analyses the mixing effect on productivity in relation to the combination of species, stand age and soil fertility, and discusses possible consequences of forest management, for five two-species mixtures in the Netherlands: Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)beech (Fagus sylvatica L.), pine (Pinus sylvestris L.)oak (Quercus robur L.), oakbeech, oakbirch (Betula pendula Roth) and pinebirch. These mixtures and their corresponding monoculture stands were studied using long-term permanent forest plots over multiple decades, but also using two inventories (around 2003 and 2013) across the entire Netherlands. These forest plots data were used together with empirical models at total stand level (chapter 2), species level (chapter 3) and tree level (chapter 4) to evaluate the mixing effect on forest productivity.

In chapter 2, four two-species mixtures and their corresponding monospecific stands were compared for productivity (volume stem wood in m3 ha-1 year-1). It was explored whether mixing species differing in leaf phenology and shade tolerance would lead to overyielding of mixed forest stands, and whether overyielding changes with stand development. In line with the niche complementarity hypothesis, the two evergreen–deciduous species mixtures (Douglas-fir–beech and pine–oak) showed overyielding whereas deciduous–deciduous species mixtures (oakbeech and oakbirch) did not. The overyielding was strongest for the Douglas-fir–beech mixture than the pine–oak mixture, which can be attributed to the greater difference in shade tolerance in the former mixture. Overyielding did not significantly change with stand development. It is argued that the regular thinning maintained the ability of species to partition resources, i.e. the complementary resource use in those mixed stands over all stand ages.

In chapter 3, it was analysed which of the two species in these four mixtures contributed to overyielding, and whether this overyielding changed along a soil fertility gradient. It was discovered that both the fast-growing and the slow-growing species could contribute to overyielding. Yet, it was mainly the fast-growing Douglas-fir that contributed to higher productivity in the Douglas-fir–beech mixtures, and the slow-growing oak that did so in the pine–oak mixtures. For both mixtures, the greatest relative productivity gain was achieved by mixtures on the poorer soils. At first sight, these results seem in line with the stress-gradient hypothesis and not the resource-ratio hypothesis. Yet, it was argued that not only complementary use of soil resources, but also use of light, may contribute to the higher productivity of mixed stands on the poorer soils.

In chapter 4, it was assessed how the growth of individual trees in mixtures was influenced by inter- and intra-specific competition, and whether this competition was mainly size-symmetric for soil resources or size-asymmetric for light on soils differing in fertility. This chapter focussed on three mixtures, i.e. oak–birch, pine–oak and pine–birch, which were available at sufficient numbers in the Dutch national forest inventory data. It was concluded that intra-specific competition was not necessarily stronger than inter-specific competition and this competitive reduction was less seen at lower soil fertility and dependent on species mixtures, which is not in line with the stress-gradient hypothesis. Moreover, size-asymmetric competition for light was more associated with tree basal area growth than size-symmetric competition for soil resources, suggesting that light is the most limiting resource. Competition for light was generally much stronger at high fertility soils, supporting the resource-ratio hypothesis. These results suggest that light is the most limiting resource for tree basal area growth and that reduced competition for light can be explained to some degree by complementarity in light use to increase tree growth in mixed forests.

This thesis thus described the productivity patterns when mixing tree species and explored possible mechanisms of higher productivity in mixed stands compared with monoculture stands in the Netherlands. Complementary use of aboveground and belowground resources probably contributes to the higher productivity in mixed stands, but other factors including pathogens, nutrient cycling and litter decomposition were not addressed but cannot be excluded. Overyielding in Douglas-fir–beech and pine–oak mixtures was maintained over time, probably owing to the intensive thinning in Dutch forests. The results shed new light on the stress-gradient and resource-ratio hypotheses. For mixtures in Dutch forest, the greatest productivity gain in Douglas-fir–beech and pine–oak mixtures was achieved on the poorer soils, and it was argued that this is at least partially driven by complementary use of light, while the role of complementarity in use of soil resources is more obscure. Overall, this thesis suggest a substantial potential of species mixing for increasing productivity, which may run in parallel with enhancing other ecosystem services such as conservation of diversity and other nature values. Yet, more experimental studies on productivity in mixed stands are required to better unravel alternative mechanisms. Such understanding is required to manage the forests effectively in a century of unpreceded human driven changes in environmental conditions.

FEM growth and yield data monoculures - Scots pine (revised edition)
Lu, Huicui ; Jansen, J.J. ; Oosterbaan, A. ; Goudzwaard, L. ; Oldenburger, J.F. ; Mohren, G.M.J. ; Ouden, J. den - \ 2017
growth and yield - even-aged monoculture forest - tree diameter - tree height - crown class - coordinates stem positions - age - top height - dominant height - monitoring - Scots pine - Pinus sylvestris
The current database is part of the FEM growth and yield database, a collection of growth and yield data from even-aged monocultures (douglas fir, common oak, poplar, Japanese Larch, Norway spruce, Scots pine, Corsican pine, Austrian pine, red oak and several other species, with only a few plots, even-aged mixed species forest plots, uneven-aged natural forest, uneven-aged selection forest and roadside plantattions of poplar. The FEM growth and yield data base is currently supervised by Jan den Ouden and Frits Mohren.
Global Forest Biodiversity Initiative (GFBI)- Dataset Global#1
Liang, J. ; Crowther, T.W. ; Picard, N. ; Barrett, Curtis ; Hengeveld, G.M. ; Nabuurs, G.J. ; Schelhaas, M. ; Lu, Huicui ; Zhu, J. - \ 2016
Data underpinning the Research Article entitled Positive Biodiversity–Productivity Relationship Predominant in Global Forests
Positive biodiversity-productivity relationship predominant in global forests
Liang, J. ; Crowther, T.W. ; Picard, N. ; Wiser, S. ; Zhou, M. ; Alberti, G. ; Schulze, E.D. ; Mcguire, A.D. ; Bozzato, F. ; Pretzsch, H. ; Miguel, S. de; Paquette, A. ; Herault, B. ; Scherer-lorenzen, M. ; Barrett, C.B. ; Glick, H.B. ; Hengeveld, G.M. ; Nabuurs, Gert-Jan ; Pfautsch, S. ; Viana, H. ; Vibrans, A.C. ; Ammer, C. ; Schall, P. ; Verbyla, D. ; Tchebakova, N. ; Fischer, M. ; Watson, J.V. ; Chen, Han Y.H. ; Lei, X. ; Schelhaas, M.J. ; Lu, Huicui ; Gianelle, D. ; Parfenova, E.I. ; Salas, C. ; Lee, E. ; Lee, B. ; Kim, H.S. ; Bruelheide, H. ; Coomes, D.A. ; Piotto, D. ; Sunderland, T. ; Schmid, B. ; Gourlet-Fleury, S. ; Sonke, B. ; Tavani, R. ; Zhu, J. ; Brandl, S. ; Vayreda, J. ; Kitahara, F. ; Searle, E.B. ; Neldner, V.J. ; Ngugi, M.R. ; Baraloto, C. ; Frizzera, L. ; Ba Azy, R. ; Oleksyn, J. ; Zawila-Niedzwiecki, T. ; Bouriaud, O. ; Bussotti, F. ; Finer, L. ; Jaroszewicz, B. ; Jucker, T. ; Valladares, F. ; Jagodzinski, A.M. ; Peri, P.L. ; Gonmadje, C. ; Marthy, W. ; Obrien, T. ; Martin, E.H. ; Marshall, A.R. ; Rovero, F. ; Bitariho, R. ; Niklaus, P.A. ; Alvarez-Loayza, P. ; Chamuya, N. ; Valencia, R. ; Mortier, F. ; Wortel, V. ; Engone-Obiang, N.L. ; Ferreira, L.V. ; Odeke, D.E. ; Vasquez, R.M. ; Lewis, S.L. ; Reich, P.B. - \ 2016
Science 354 (2016)6309. - ISSN 0036-8075 - 15 p.
The biodiversity-productivity relationship (BPR) is foundational to our understanding of the global extinction crisis and its impacts on ecosystem functioning. Understanding BPR is critical for the accurate valuation and effective conservation of biodiversity. Using ground-sourced data from 777,126 permanent plots, spanning 44 countries and most terrestrial biomes, we reveal a globally consistent positive concave-down BPR, showing that continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The value of biodiversity in maintaining commercial forest productivity alone—US$166 billion to 490 billion per year according to our estimation—is more than twice what it would cost to implement effective global conservation. This highlights the need for a worldwide reassessment of biodiversity values, forest management strategies, and conservation priorities.
Overyielding of temperate mixed forests occurs in evergreen–deciduous but not in deciduous–deciduous species mixtures over time in the Netherlands
Lu, Huicui ; Mohren, G.M.J. ; Ouden, J. den; Goudiaby, V. ; Sterck, F.J. - \ 2016
Forest Ecology and Management 376 (2016). - ISSN 0378-1127 - p. 321 - 332.
Recent studies show that mixed species forests sometimes have higher stand productivity than monospecific forests, which we refer to as overyielding. Yet, results for temperate forests are ambiguous, possibly because forests differ in local site conditions, thinning history and forest age. In line with the niche complementarity hypothesis, we expect stronger overyielding for forests with species differing in both leaf phenology (evergreen or deciduous) and shade tolerance. We also hypothesize that overyielding will decrease with stand development because of decreasing resource availability. We compared 4 two-species mixtures with their corresponding monospecific stands from long-term field measurements in the Netherlands. The mixtures were Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)–common beech (Fagus sylvatica L.), Scots pine (Pinus sylvestris L.)–common oak (Quercus robur L.), common oak–common beech, and common oak–silver birch (Betula pendula Roth). Overyielding was observed in 2 of the 4 mixtures: Douglas-fir–common beech mixtures had 35.9% and 36.7% higher volume growth relative to Douglas-fir and common beech monocultures, respectively; Scots pine–common oak mixtures had 20.3% and 31.2% higher volume growth relative to Scots pine and common oak monocultures, respectively, on average over time. Furthermore, overyielding was relatively constant for the two mixtures through stand development. This result was robust after accounting for possible effects of site quality and thinning history, where site quality contributed independently to stand productivity and thinning history had no effect. No significant overyielding effects were observed for the two deciduous mixed stands, i.e. common oak–common beech and common oak–silver birch. Mixing tree species in temperate forests resulted in overyielding for evergreen–deciduous species mixtures, but not for deciduous–deciduous species mixtures. This indicates that leaf phenology contributes to overyielding effects. Overyielding was higher in the Douglas-fir–common beech mixtures than the Scots pine–common oak mixtures, which coincides with a stronger contrast in shade tolerance between Douglas-fir and common beech and thus stronger complementarity. Our results support the complementarity hypothesis and imply that such mechanisms are maintained with stand development. It therefore appears that mixing evergreen with deciduous species with contrasting shade tolerance is a valid management strategy for increasing diversity and productivity of temperate forests in the Netherlands.
FEM growth and yield data mixed species forest
Bartelink, H.H. ; Jansen, J.J. ; Goudzwaard, L. ; Lu, Huicui ; Oldenburger, J.F. ; Mohren, G.M.J. ; Ouden, J. den - \ 2016
growth and yield - even aged mixed species forest - tree diameter - tree height - crown class - crown diameter - crown base - coordination stem positions - age - top height - dominant height - dominant diameter - monitoring - ash common beech - common oak - Douglas fir - Japanese larch - Scots pine - silver birch - sycamore - Acer pseudoplatanus - Betula pendula - Fagus sylvatica - Fraxinus excelsior - Larix kaempferi - Quercus robur - Pinus sylvestris - Pseudotsuga mensisii
The current database is part of the FEM growth and yield database, a collection of growth and yield data from even-aged monocultures (douglas fir, common oak, poplar, Japanese Larch, Norway spruce, Scots pine, Corsican pine, Austrian pine, red oak and several other species with only a few plots like grand fir, even-aged mixed species forest plots, uneven-aged natural forest, uneven-aged selection forest and roadside plantations of poplar). The FEM growth and yield data base is currently supervised by Jan den Ouden and Frits Mohren.
FEM growth and yield data - Common oak
Oosterbaan, A. ; Jansen, J.J. ; Goudzwaard, L. ; Lu, Huicui ; Oldenburger, J.F. ; Mohren, G.M.J. ; Ouden, J. den - \ 2016
growth and yield - even-aged monoculture forest - understory - tree height - tree diameter - crown class - coordinates stem positions - age - top height - dominant height - dominant diameter - monitoring - Common oak - Quercus robur
The current database is a part of the FEM growth and yield database, a collecction of growth and yield data from even-aged monocultures (douglas fir, common oak, poplar, Japanese Larch Norway spruce, Scots pine, Corsican pine, Austrian pine, red oak and several other species with only a few plots, even-aged mixed species forest plots, uneven-aged natural forest, uneven-aged selection forest and roadside plantations of poplar. The FEM growth and yield data base is currently supervised by Jan den Ouden and Frits Mohren.
FEM growth and yield data monoculures - Scots pine
Lu, Huicui ; Jansen, J.J. ; Oosterbaan, A. ; Goudzwaard, L. ; Oldenburger, J.F. ; Mohren, G.M.J. ; Ouden, J. den - \ 2016
growth and yield - even-aged monoculture forest - tree diameter - tree height - crown class - coordinates stem positions - age - top height - dominant height - monitoring - Scots pine - Pinus sylvestris
The current database is part of the FEM growth and yield database, a collection of growth and yield data from even-aged monocultures (douglas fir, common oak, poplar, Japanese Larch, Norway spruce, Scots pine, Corsican pine, Austrian pine, red oak and several other species, with only a few plots, even-aged mixed species forest plots, uneven-aged natural forest, uneven-aged selection forest and roadside plantattions of poplar. The FEM growth and yield data base is currently supervised by Jan den Ouden and Frits Mohren.
FEM growth and yield data monocultures - Common beech
Goudzwaard, L. ; Jansen, J.J. ; Oosterbaan, A. ; Oldenburger, J.F. ; Lu, Huicui ; Mohren, G.M.J. ; Ouden, J. den - \ 2016
growth and yield - even-aged monoculture forest - understory - tree diameter - tree height - crown class - coordinates stem positions - age - top height - dominant diameter - monitoring - Common beech - Fagus sylvatica
The current database is part of the FEM growth and yield database, a collection of growth and yield data from even-aged monocultures (douglas fir, common oak, poplar, Japanese Larch, Norway spruce, Scots pine, Corsican pine, Austrian pine, red oak and several other species, with only a few plots, even-aged mixed species forest plots, uneven-aged natural forest, uneven-aged selection forest and roadside plantattions of poplar. The FEM growth and yield data base is currently supervised by Jan den Ouden and Frits Mohren.
FEM growth and yield data monocultures - Silver birch
Oldenburger, J.F. ; Jansen, J.J. ; Oosterbaan, A. ; Lu, Huicui ; Mohren, G.M.J. ; Ouden, J. den - \ 2016
growth and yield - even-aged monoculture forest - tree diameter - tree height - crown class - coordinates stem positions - age - top height - dominant height - dominant diameter - monitoring - silver birch - Betula pendula
The current database is part of the FEM growth and yield database, a collection of growth and yield data from even-aged monocultures (douglas fir, common oak, poplar, Japanese Larch, Norway spruce, Scots pine, Corsican pine, Austrian pine, red oak and several other species, with only a few plots, even-aged mixed species forest plots, uneven-aged natural forest, uneven-aged selection forest and roadside plantattions of poplar. The FEM growth and yield data base is
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