Forest Ecology and Management 534 (2023) 120882

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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

The impact of planting density on forest monospecific plantations:

An overview
Mônica Moreno Gabira a, *, Miguel Montoro Girona b, c, Annie DesRochers b, Dagma Kratz a,
Richardson Barbosa Gomes da Silva d, Manoela Mendes Duarte a, Natalia Saudade de Aguiar a,
Ivar Wendling e
a
Department of Forest Sciences, Federal University of Paraná (UFPR), av. Prefeito Lothário Meissner, 623, Jardim Botânico, Curitiba, 80210 170, PR, Brazil
b
Groupe de Recherche en Écologie de la MRC Abitibi, Institut de recherche sur les forêts (IRF), Université du Québec en Abitibi-Témiscamingue (UQAT), 341, rue
Principale Nord, Amos, J9T 2L8, QC, Canada
c
Department of Wildlife, Fish and Environmental Studies, Swedish University of Agricultural Sciences, Umeå, SE-901 83, Sweden
d
Department of Forest Science, Soils and Environment, School of Agriculture, São Paulo State University (UNESP), Botucatu, 18610-034, SP, Brazil
e
Embrapa Florestas, Estrada da Ribeira, 101, Parque Monte Castelo, Colombo, 83411 000, PR, Brazil

A R T I C L E I N F O A B S T R A C T

Keywords: Knowledge of the morphological and physiological responses of trees to planting density is important for
Growth and yield adjusting plantation establishment practices to attain the desired size and characteristics of wood products. In
Forestry this review, we provide an overview of how planting density is managed as a silvicultural practice and sum­
Forest stands
marize recent advances, trends, and gaps to guide future research in this field. We applied a PRISMA method­
Silviculture
ology to select papers published between 1981 and 2020 that tested how monospecies planting density affects
Stand density
Tree spacing the morphological and physiological characteristics of planted trees; we observed an increasing trend in the
number of relevant publications over this period. These studies focused mainly on species used for timber
production (Eucalyptus and Pinus), and research was concentrated in the United States, China, and Brazil. In
general, the 255 selected studies identified that planting density has a major impact on stem diameter and
volume, resulting in greater stand productivity of high-density stands, especially for trees having a smaller in­
dividual volume. Studies evaluating the physiological response of trees to planting density also showed that
species ecology, solar radiation, water requirements, and site-specific characteristics have a greater influence on
plant growth than planting density. Considering the physiological aspects in high-density plantations, water
relations based on transpiration and water use efficiency are the main aspect guiding plants growth rate. Despite
major advances in intensive silviculture around the world, there remains a knowledge gap for tree species grown
for their leaves, fruits, or seeds and a lack of data related to the physiological response of trees to planting
density. The results of this literature survey can improve forest management decisions in regard to forest stand
use and develop novel study avenues for silviculture.

1. Introduction
Forest-based industry is a crucial sector of socioeconomic develop­
ment around the world and provides several raw products to fiber,
timber, energy, and non-timber forest-product industries. Forest plan­
tations for industrial purposes cover about 131 million ha, correspond­
ing to 2 % of the world’s total forest cover (FAO, 2020). Asia contains
the largest area of plantation forests (79,332,000 ha of planted forests),
and the highest ratio of introduced species in planted forests is in South
America, where 97 % of the planted areas contain non-native species.
The ecological, economical, and social importance of planted forests has
been discussed extensively, and plantations are considered important for
community development, particularly in the context of climate change
and post COVID-19 pandemic (Celentano et al., 2022; Olmos, 2022). In
general, plantation forests are established with a design aimed at
providing the optimal conditions for tree growth and maximum pro­
ductivity and for managing timber quality with a focus on the desired
final products.
Silviculture in forests and plantations has a main goal of managing
competition for light, nutrients, and water among neighboring trees.

* Corresponding author.

https://doi.org/10.1016/j.foreco.2023.120882
Received 8 August 2022; Received in revised form 15 February 2023; Accepted 17 February 2023
Available online 22 February 2023
0378-1127/© 2023 Elsevier B.V. All rights reserved.
M.M. Gabira et al.
Competition is defined as the demand by individuals for a common
required resource that may exceed the immediate supply; thus,
competition reduces tree productivity (Begon and Townsend, 2020).
Light is often considered a source of competition between trees because
it is the only environmental factor that cannot be easily managed in
even-aged and un-thinned forest monocultures (Chase et al., 2016;
Nelson et al., 2016); this competition leads to physiological and
morphological changes—also called shade-avoidance responses—which
include hyponasty, stem and petiole elongation, apical dominance, leaf
positioning, and life-cycle adjustments (Bongers et al., 2018). Huber
et al. (2021) explored the physiological, biochemical, and genetic as­
pects of shade-avoidance responses caused by light signaling; they
demonstrated that competition for light may impact the canopy struc­
ture, species composition, and size of individuals in plant communities.
Competition affects plant physiology directly, mediating individual
growth by regulating resource acquisition (Benomar et al., 2012). In
drier regions, light may not be as limiting as water for productivity,
which limits plant growth and affects resource- use efficiency and
biomass allocation (Gonçalves et al., 2004; Stape et al., 2004). Space is
also a limiting factor for tree growth because it influences resource
consumption per area; however, space is not considered a resource for
which plants compete because it is probably never limiting in terrestrial
ecosystems, as opposed to water and other essential resources (Wilson
et al., 2007). Planted forests are sites of relatively homogeneous
competition because trees are usually even-aged, regularly spaced in­
dividuals of the same species and deal with a relatively uniform resource
availability at the stand scale; however, changes in environmental re­
sources availability at a stand scale are known to influence productivity,
as observed by Resende et al. (2018). Thus, understanding species re­
sponses to spatiotemporal variations in resources is essential to provide
theoretical guidance to stand management in different regions.
Despite this focus on competition, silviculturists must consider that
trees are not always competing with each other; facilitation relation­
ships also occur in forest stands, and silvicultural practices aiming to
reduce tree density do not always positively affect growth (Quer et al.,
2020). In general, facilitation relationships create a buffer that mitigates
variations in soil humidity, atmospheric temperature, and light inter­
ception and favors plant establishment and growth (Bravo-Oviedo et al.,
2018). An example of facilitation is to mix species in forest stands as a
way to increase its growth, as explored by Forrester (2015); although the
author observed that not all trees increased their growth in comparison
to monospecific stands. Natural root grafting has also been reported as a
means of creating facilitation relationships among individual jack pine
(Pinus banksiana Lamb.) to improve resource use by the trees (Tarroux
and Desrochers, 2011). When these stands are thinned, however, the
root system of the removed trees increases the carbon demand on the
residual trees, decreasing the latter’s growth relative to non-root-grafted
trees, despite benefiting from more resources (Tarroux et al., 2010). This
example illustrates how tree growth can depend not only on competition
but also facilitation (Begon and Townsend, 2020); therefore, facilitation
must be better understood in forest stands to promote tree growth and
increase the efficacy of resource use.
In the design and planning of plantations, tree spacing is a major
factor because of its impact on profitability—close spacing requires
more seedlings—and productivity because closer spacings can improve
tree form and timber quality for some industries, e.g., the veneer in­
dustry (Charlton et al., 2020; Glencross et al., 2012; Naji et al., 2016;
Tonini et al., 2018). Tree growth tends to reach a maximum at a certain
density and then declines with additional trees; however, this trend also
depends on stand age, site quality, and environmental conditions (Will
et al., 2001). As a key to understanding density-related patterns, Weiner
and Freckleton (2010) reviewed the empirical and mathematical basis of
constant final yield. They concluded that constant final yield was the
most robust pattern in ecology and deemed it as essential for under­
standing and predicting plant community growth. Intertree competition
is generally minimal for resources at very low tree densities, leading to a
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Forest Ecology and Management 534 (2023) 120882
regular increase in biomass accumulation up to a certain point where
biomass accumulation decreases because natural resources become
limiting for growth.
Given the importance of forest plantations worldwide and the po­
tential of planting density to be used as management tool, several studies
have been developed to clarify species responses to planting density at
individual and stand levels in forest plantations. However, to our
knowledge, the results of these studies have never been combined and
analyzed through a systematic review, which offers many advantages
over other research synthesis techniques, as its results can be evaluated,
investigated, and updated more accurately. In this review, we discuss
how planting density affects plantation forest growth, yield, and phys­
iological responses in temperate and tropical regions. We focus on
monospecific planted forests because they are some of the most inten­
sively managed stands, contain generally only one or two species, and
are characterized by even-aged regularly spaced trees. We provide an
overview of how planting density is managed as a silvicultural practice
and summarize recent advances, trends, and gaps to guide future
research in this field.
2. Material and methods
2.1. Criteria for selecting planting density − growth studies
We searched the existing literature for studies published between
1981 and 2020 focused on the morphological and physiological re­
sponses of plants to planting density in forest stands. We selected this 40-
year period to highlight the most up-to-date advances in the field of
planting density available in online databases. The literature review was
conducted according to the PRISMA methodology (Moher et al., 2009;
Fig. 1).
We used two indexing electronic databases—Web of Science and
Scopus—to identify the available literature and encompass a wide va­
riety of high-quality journals. Title, abstract, and keywords were
searched for the words “forest,” “planting,” “plant,” “tree,” “plant,”
“plantation,” “stem,” “stand,” “spacing,” “stock,” “stocking,” and “den­
sity,” using Boolean operators, e.g., OR, AND. Our search was limited to
research papers published between 1981 and 2020 in English-language
scientific journals, and we excluded review articles, event papers,
technical reports, and short communications. Our database search
returned 11,602 papers in Web of Science and 8097 papers in Scopus. A
total of 10,820 papers remained after removing duplicates.
In the screening process, we analyzed and excluded papers that did
not meet specific criteria. We excluded papers that (a) focused on
agricultural crop species, such as cotton, wheat, maize, and soy; (b) were
based on field trials established in drylands or wetlands; (c) focused on
the development of GIS and remote-sensing technologies; and (d) were
based on modelling or mathematical models, not on field trials. We
included papers focused on either temperate or tropical species (or
both), given that the most relevant planting density studies occur in both
zones. After the application of these criteria, we retained 188 papers.
The remaining papers were then evaluated in terms of the clarity of the
supplied information and the application of stand density within the
field trials. We included in our study only papers that (a) were based on
data collected from planted forests and that provided clear, specific in­
formation about spacing and evaluation methods; (b) used tree density
as a factor affecting growth in planted stands, thus excluding studies
having density as a dependent variable in natural forest stands; (c)
evaluated planted tree species response, not the response of natural
understory regeneration, shrubs, or herbaceous species; and (d) evalu­
ated tree morphological and physiological responses to planting density,
thereby excluding papers investigating wood quality or environmental
aspects, such as soil properties and understory species occurrence. After
this second group of criteria, we retained 255 papers for this review.
M.M. Gabira et al.
Fig. 1. Workflow description of the PRISMA searching methodology, .
adapted from Moher et al. (2009)
2.2. Data analysis
We used descriptive statistics to analyze general trends related to
publication frequency, study regions, study species, and spacing size. All
selected papers were evaluated, and we extracted and analyzed relevant
information. We evaluated relationships between growth variables and
planting density. For this analysis, we considered the morphological and
physiological characteristics of studied trees detailed in selected papers.
These characteristics included total height, diameter at breast height,
individual and stand basal area, individual and stand timber volume,
leaf biomass, leaf area index, crown height and diameter, intercepted
radiation, carbon assimilation, transpiration, respiration, water-use ef­
ficiency, leaf water potential, and chlorophyll content. We extracted all
relevant information related to these variables, and for each paper we
classified our results as a neutral (*), positive (+), or negative (− )
response to spacing. From this classification, we produced tables pre­
senting the studied species and the associated response of morphological
Fig. 2. Annual number of studies focused on planting density and its effect on tree growth published between 1981 and 2020.
3
Forest Ecology and Management 534 (2023) 120882
or physiological growth characteristics to tree spacing.
3. Results and discussion
3.1. Evolution of planting density–growth studies between 2001 and 2020
We observed an increasing trend in the number of publications
focused on planting density and growth over the 1981–2020 period
(Fig. 2). Publication abundance fluctuated between zero and 33 papers
per year, with the highest peaks observed in 2019 and 2020, whereas we
found only one planting density study in 1985,1986,1989, 1991, and
2006, and we did not locate studies published in 1981, 1982, 1983,
1987, and 1995. It is important to mention that planting density in forest
stands was already a focus of studies before 1981, but most papers are
not available online. A greater focus on increasing timber quality (i.e.,
straight shape and decay-resistant wood) and reducing rotation times
drove the last 20-year increase in planting density studies, with financial
M.M. Gabira et al.
costs and environmental change as a consequence of planting density
also being central elements in these papers (Bentancor et al., 2019;
Charlton et al., 2020; Duan et al., 2019; Smith et al., 2016). Increasing
planting density provides an easy means of increasing wood production,
especially in regions where areas available for forest plantations are
scarce because of competing crops or resources availability. These
concerns have been raised in the last decades by several international
institutions and worldwide agreements aimed at reducing deforestation
and mitigating climate change, e.g., the FAO and the Paris Agreement.
Globally, studies of planting density were widespread, although
many countries had only a single study on their territory (Fig. 3). The
subject of planting density was most frequent in trials located in the
United States (US – 60), China (31), and Brazil (31). These countries
have a strong tradition of plantation forestry, although the species and
products generated from these forests serve various industries—mainly
timber, pulp and paper, and biomass for energy production. The US and
China lie in temperate climates, with studies mainly involving planted
forests of coniferous species (Groot and Cortini, 2016; Harrington et al.,
2009; Will et al., 2005, 2001; Zhao et al., 2020, 2011), whereas Brazil
lies in the tropics and contains planted forests of various broad-leaved
species (Narducci et al., 2016; Páscoa et al., 2019; Schwerz et al.,
2020a, 2020b; Silva et al., 2012; Tonini et al., 2018). We thus observed a
greater number of studies on Pinus spp. in the US and China, and more
studies on Eucalyptus spp. in Brazil. Other countries in which the forest
industry is an important economic sector did not present high numbers
of planting density studies because the studies focused on natural forests
did not have regular spacing between plants, explored thinning exper­
iments, and evaluated ecological variables other than growth. These
latter studies were removed from this review during the screening
process.
Species from the genera Pinus and Eucalyptus were the most studied
in planting density trials, representing 28 % and 19 % of the analyzed
studies, respectively (Fig. 4). Among the Pinus taxa, P. taeda L. was the
most used in the trials, being found in 40 studies. The most-widely
studied Eucalyptus species was E. grandis W. Hill ex Maiden, used in 18
studies. These abovementioned taxa are the most important species for
plantations in several countries, and they are used by timber, pulp and
paper, and energy industries.
Nonetheless, the lack of information on tree species cultivated for
products other than wood was obvious from our literature search; we
only found a handful of species evaluated for leaves and/or fruits:
Fig. 3. Location and abundance of published studies between 2001 and 2020 that assessed planting density effects on growth.
4
Forest Ecology and Management 534 (2023) 120882
Camellia sinensis L. (Kigalu, 2007a, 2007b), Hevea brasiliensis (Willd. ex
A. Juss.) Müll. Arg. (Naji et al., 2014), Fraxinus angustifolia Vahl
(Özbayram and Çiçek, 2020), Gliricidia sepium (Jacq.) Kunth ex Walp.
(Silva et al., 2012), Jatropha curcas L. (Andrianirina et al., 2019), and
Moringa oleifera Lam. (Mendieta-Araica et al., 2013). Thus, we cannot
identify how planting density affects many species, despite the major
effort undertaken in the studies of selected economically important taxa;
the results of these species-specific studies cannot always be extrapo­
lated to other taxa.
We observed that planting densities differed between coniferous and
broad-leaved species (Fig. 5). Planting density varied among species and
ranged between 25 stems⋅ha− 1 in a Quercus robur L. Nelder Wheel trial
(Uhl et al., 2015) to 167,000 stems⋅ha− 1 for Moringa oleifera Lam.
(Mendieta-Araica et al., 2013). Differences in planting density related
mainly to the planned end products from these trials. Pinus spp., rep­
resenting the commonly studied coniferous species in the reviewed
studies, are used mainly for timber production, especially for the
sawmill industry. This industry requires larger-diameter timber, which
cannot be produced at high planting densities. Conversely, Eucalyptus
spp. and Salix spp., among other broad-leaved species, are used for pulp
and paper or energy industries; smaller diameter boles can be used for
this purpose and be grown under conditions of shorter rotation periods
and higher planting densities. Studies with Camellia sinensis L. also use
higher densities because the main products of these plantations are the
leaves used in the tea industry (Kigalu, 2007a, 2007b).
3.2. How does planting density affect tree growth?
Studies of tree density in planted forest stands mainly address
biomass production and timber quality, which reflected the most
commonly measured morphological variables: diameter at breast height
(DBH, 74 % of studies), height (H, 64 %), and stand volume (SV, 48 %).
Table 1 presents the growth response of trees species to increasing
planting density. In general, there is a trade-off between tighter spacing
and biomass production per area, and initial spacing strongly influences
plant growth at various times during stand development. Higher
planting densities reduce stem and/or crown diameter and, in most
studies, there is a linear relationship between these growth variables
(Benomar et al., 2012; Dassot et al., 2015; Han et al., 2020; Harrington
et al., 2009; Páscoa et al., 2019; Silva et al., 2012). Height growth tends
to increase with planting density; however, this trend is not consistent
M.M. Gabira et al.
Fig. 4. Frequency of genus studied in planting density–growth studies published between 2001 and 2020.
Fig. 5. Average density of genera studied in planting density–growth trials published between 2001 and 2020.
and not present in many studies. Height is not always a planting den­
sity–dependent variable, and soil fertility has also been deemed the most
critical factor determining height growth (Henskens et al., 2001;
Woodruff et al., 2002); therefore, competition for soil nutrients could be
the main cause for reduced height growth for trees in high-density
stands. Kholdaenko et al. (2022), for example, demonstrated that a
high planting density in Picea obovate Ledeb. plantations reduced height
growth considerably in suppressed trees because of an earlier onset of
competition for nutrients.
In the initial years after planting, the interaction between trees is
limited and competition may not be observed until a tree reaches the
above- and belowground space of its neighbor. Studies analyzed in our
review evaluated trees from 1 (Schwerz et al., 2019; Silva et al., 2019) to
90 (Węgiel and Polowy, 2020) years after planting, and results related to
the morphological characteristics were highly heterogeneous. For
Acacia mearnsii De Wild., Schwerz et al. (2019) observed a greater
biomass allocation in branches and leaves until seven years after
planting – an increase of 29 % in the narrowest spacing, compared to the
widest spacing. Seven years after planting, they observed a change in the
pattern of biomass partitioning, with a relative reduction in leaf biomass
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Forest Ecology and Management 534 (2023) 120882
and an increase in total stand timber biomass, regardless of planting
density. In a Eucalyptus plantation, Silva et al. (2019) observed differ­
ences in DBH and volume per tree between planting densities as early as
24 months after planting – an increase of 22 % in DBH and 90 % in
volume per tree in the widest spacing compared to the other planting
spacings. The authors attributed the greater DBH and volume per tree to
the higher light interception by tree crowns within low-density planta­
tions, which led to a higher production of photoassimilates. Studies that
examined biomass allocation along gradients of light, water, and
nutrient availability also found support for the hypothesis that plants
tend to allocate biomass to organs that acquire the most limiting re­
sources (Ma et al., 2021; McCarthy and Enquist, 2007; Rodríguez-García
and Bravo, 2013); therefore, plants competing for resources in stands
tend to change allocation patterns according to stand age and planting
density. The greater allocation to the growth of branches and leaves in
the first years form the canopy, important for light interception and
determining forest productivity (Barbier et al., 2010). Changes in tree
growth after canopy closure are often related to competition with
neighboring trees, which is more intense in high-density plantations
(Will et al., 2001).
M.M. Gabira et al.
Table 1
Summary of studies published between 2001 and 2020 that measured morphological characteristics related to tree growth in response to increasing planting density.
Species
Acacia auriculiformis A.
Cunn. ex Benth.
Acacia farnesiana Linnaeus
Acacia koa A. Gray
Acacia mangium Willd.
Acacia mearnsii De Wild.
Acacia nilotica subsp.
cupressiformis (J.L.
Stewart) Ali & Faruqi
Acer velutinum Boiss.
Alnus rubra Bong.
Ateleia glazioviana Baill.
Betula alba L.
Betula alnoides Buch.-Ham.
ex D. Don
Betula pendula Roth.
Betula pubescens Ehrh.
Camellia sinensis L.
Cariniana legalis (Mart.)
Kuntze
Cassia siamea Lam.
Cordia africana Lam.
Cordia alliodora (Ruiz &
Pav.) Oken
Cryptomeria japonica D. Don
Cunninghamia lanceolata
(Lamb.) Hook
Endospermum medullosum LS
Smith
Eremanthus erythropappus
(DC.) MacLeish
Eucalyptus benthamii Maiden
& Cambage
Eucalyptus camaldulensis
Dehnh.
Eucalyptus cloeziana F.
Muell.
Eucalyptus dunnii Maiden
Eucalyptus globulus Labill.
Eucalyptus grandis W. Hill ex
Maiden
E. grandis W. Hill ex Maiden
× E. camaldulensis Dehnh.
E. grandis W. Hill ex Maiden
× E. urophylla S.T. Blake
Eucalyptus macarthurii H.
Deane & Maiden
Eucalyptus marginata Donn
ex Sm.
Eucalyptus nitens (H.Deane &
Maiden) Maiden
Eucalyptus paniculata Sm.
Eucalyptus pilularis DC.
Eucalyptus saligna (Sm.)
Eucalyptus sideroxylon
Woolls
Eucalyptus smithii F.Muell. ex
R.T.Baker
Forest Ecology and Management 534 (2023) 120882
S H DBH CH CD IV SV IBA SBA LB LAI Reference
− + − (Feng and Xue, 2019)
* * * + * (Goel and Behl, 2004)
+ (Meinzer et al., 1996)
*/− */+ − + (Nirsatmanto, 2016; Tonini et al., 2018)
+ + + (Eloy et al., 2018; Schwerz et al., 2019)
− − − * − (Goel and Behl, 2004)
− (Naji et al., 2016)
*/+ +/− − + + (DeBell and Harrington, 2002; Negrave et al.,
2007)
* − +/− + + + (Eloy et al., 2018; Schwerz et al., 2020a, 2020b)
− + (Fernández-Núñez et al., 2010)
− * * + * * (Wang et al., 2016, 2018, 2019, 2020)
* */− − + − * + (Ilomäki et al., 2003; Johansson, 2007)
* * − − */+ + (Johansson, 1989; 2007)
+ (Kigalu, 2007b)
* − − + (Oliveira et al., 2018)
* * * + (Goel and Behl, 2004)
* − − (Mehari and Abte, 2006)
* − (Hummel, 2000)
+ (Lam and Guan, 2020)
− */− − − + + + + (Bai et al., 2020; Farooq et al., 2020; Liu et al.,
2019; Zhang et al., 2017, 2020b)
+ * − */+ + (Glencross et al., 2012; Smith et al., 2016)
*/− − − (Páscoa et al., 2019; Scolforo et al., 2016)
* * − − + (Benin et al., 2014; Binkley et al., 2017; Resquin
et al., 2018,2019, 2020 )
+ */− − */− − + + (Akhtar et al., 2008; Bernardo et al., 1998;
Horner et al., 2009; Nagar et al., 2015; Paula
et al., 2013; Thoranisorn et al., 1990)
* − * (Alcorn et al., 2007)
* */− − − */+ + (Binkley et al., 2017; Bentancor et al., 2019;
Boreham and Pallett, 2009; Crous et al., 2013,
2019; Pallett and Sale, 2004; Resquin et al., 2018,
2019, 2020; Stape and Binkley, 2010)
− * − − +/− − + + (Forrester et al., 2013; Henskens et al., 2001;
Khanna, 1997; Luo et al., 2018; Miranda et al.,
2015)
*/+ */− */− − +/− * + + + (Binkley et al., 2017; Boreham and Pallett, 2009;
Crous et al., 2013, 2019; du Plessis and Kotze,
2011; Eastham et al., 1990; Eloy et al., 2016,
2018; Ferraz Filho et al., 2018; Forrester and
Smith, 2012; Kearney et al., 2007; Kirongo et al.,
2012; Pallett and Sale, 2004; Resquin et al., 2018,
2019, 2020; Schwerz et al., 2020a, 2020b)
* − − − + + (Binkley et al., 2017; Crous et al., 2013;
Hakamada et al., 2017,2020a, 2020b )
− */− */− + − + + (Binkley et al., 2017; Hakamada et al., 2017,
2020a, 2020b; Medeiros et al., 2020; Ramalho
et al., 2019; Silva et al., 2019; Tavares et al.,
2020)
+ (Boreham and Pallett, 2009; Crous et al., 2019;
Pallett and Sale, 2004)
* − + + (Grigg et al., 2008)
* − + − + (Boreham and Pallett, 2009; Pallett and Sale,
2004)
* − − − (Crous et al., 2013)
*/+/− */− */+/− − * (Alcorn et al., 2007; Cassidy et al., 2013; Forrester
and Smith, 2012; Kearney et al., 2007; West and
Smith, 2019, 2020)
+ * * (Arias-Aguilar et al., 2020; Binkley et al., 2017)
* * * * (Crous et al., 2013)
+ (Boreham and Pallett, 2009; Crous et al., 2019;
Pallett and Sale, 2004)
(continued on next page)
6
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882

Table 1 (continued )
Species S H DBH CH CD IV SV IBA SBA LB LAI Reference

Eucalyptus tereticornis Smith + * * − + (Arias-Aguilar et al., 2020; Rai and Srinivasan,

1990)
Eucalyptus urophylla S.T. * − − + + (Bernardo et al., 1998; Binkley et al., 2017;
Blake Hakamada et al., 2017,2020a)
E. urophylla S.T. Blake × − + (Binkley et al., 2017)
Eucalyptus globulus Labill.
E. urophylla S.T. Blake × − + (Binkley et al., 2017)
Eucalyptus tereticornis
Smith
Fagus orientalis Lipsky − * + (Guner et al., 2010)
Fagus sylvatica L. − * */− * − (Dassot et al., 2015; Gemmel et al., 1996)
Fraxinus angustifolia Vahl */− * + * (Çiçek et al., 2010; Özbayram and Çiçek, 2020)
Gliricidia sepium (Jacq.) */− − − * * (Ramos-Trejo et al., 2020; Silva et al., 2012,
Kunth ex Walp. 2014)
Gmelina arborea Roxb. − * * − + + (Tenorio et al., 2019)
Grevillea robusta A.Cunn. ex * − − + (Kalinganire, 1996)
R.Br.
Hevea brasiliensis (Willd. ex */− − + (Lokmal et al., 2008; Naji et al., 2014)
A. Juss.) Müll. Arg.
Jatropha curcas L. * * (Andrianirina et al., 2019; Lima et al., 2016)
Juglans nigra L. − + − (Bohanek and Groninger, 2003)
Leucaena leucocephala */− */− − */− − + (Liu et al., 2018; Pachas et al., 2018; Prasad et al.,
(Lam.) de Wit 2011)
Liquidambar styraciflua L. − (Zutter et al., 1998)
Mimosa caesalpiniifolia − − − * * (Silva et al., 2012, 2014)
Benth.
Mimosa scabrella Benth. * − +/− + + + (Eloy et al., 2018; Schwerz et al., 2020a,2020b)
Moringa oleifera Lam. */+ + + (Basra et al., 2015; Kumar et al., 2017; Mendieta-
Araica et al., 2013; Zheng et al., 2016)
Morus alba L. * (Ramos-Trejo et al., 2020)
Mytilaria laosensis Lecomte * − (Wang et al., 2020)
Peltophorum dubium + − − − (Matos et al., 2015)
(Spreng.) Taub.
Picea abies (L) */− */− */− + − + + + (Coordes, 2013; Gizachew et al., 2012; Hager and
Kazda, 1985; Nilsson, 1994; Štefančík, 2020)
Picea glauca (Moench) Voss. */− */+ − − − − + + (McClain et al., 1994; Metsaranta, 2019)
Picea mariana (Mill.) Britton, */+/− */− − − − − + + (Groot and Cortini, 2016; Metsaranta, 2019;
Sterns & Poggenb. Newton, 2015)
Pinus banksiana Lamb. */− − + − − (Hébert et al., 2016; McClain et al., 1994;
Newton, 2015, Tong and Zhang, 2005; Wells,
1994)
Pinus contorta Douglas ex +/− − − + */+ (Liziniewicz et al., 2012; Long and Smith, 1988)
Loudon
Pinus elliottii Engelm. − * − + * + + + + (Barron-Gafford et al., 2003; Burkes et al., 2003;
Gadow and Kotze, 2014; Lee et al., 1997; Will
et al., 2001; Zhao and Kane, 2012; Zhao et al.,
2019)
Pinus massoniana Lamb. * − + − * (Yan et al., 2019)
Pinus patula Schiede ex +/− */+/− − * (Charlton et al., 2020; Erasmus et al., 2018, 2020;
Schltdl. & Cham. Gadow and Kotze, 2014)
Pinus pinaster Ait. * (Mazza et al., 2015)
Pinus pinea L. */+ */− */- (Loewe-Muñoz et al., 2019, 2020)
Pinus ponderosa Douglas ex – */– */− − * – (Gyenge et al., 2010; Finley and Zhang, 2019;
C. Lawson Peracca and O’Hara, 2008; J. Zhang et al., 2019,
2020a)
Pinus radiata D. Don − * − + */+ * (Fernández-Núñez et al., 2010; Lasserre et al.,
2005; Lin et al., 2013; Snowdon et al., 1998;
Waghorn et al., 2007; Waghorn and Watt, 2013)
Pinus resinosa Torr. − * − + + (Metsaranta, 2019; Penner et al., 2001)
Pinus sibirica Du Tour − (Milyutina et al., 2013)
Pinus sylvestris L. */+ */− */− − */+ * * (Egbäck et al., 2012; Kerr, 2003; Milyutina et al.,
2013; Nan et al., 2020; Nilsson and Albresktson,
1994; Węgiel and Polowy, 2020; Węgiel and
Polowy, 2020)
Pinus taeda L. +/− */+/− */− +/− − − */+ +/− +/− */+ + (Albaugh et al., 2020; Akers et al., 2013; Amateis
and Buckhart (2013); Antón-Fernández et al.,
2011, 2012; Aspinwall et al., 2011; Baldwin et al.,
2000; Barron-Gafford et al., 2003; Bose et al.,
2018; Buford, 1991; Burkes et al., 2003; Cardoso
et al., 2013; Clark et al., 2008; Dean and Baldwin,
1996; Egbäck et al., 2015; Ferreira et al., 2020;
Harms et al., 2000; Lee et al., 1997; Quicke et al.,
1999; Radtke and Harold, 1999; Roth et al.,
2007a, 2007b; Russell et al., 2009; Shi and Cao,
1997; Subedi et al., 2012; Torreano et al., 2012;
Will et al., 2001, 2005, 2010; Yáñez et al., 2015;
(continued on next page)

7
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882

Table 1 (continued )
Species S H DBH CH CD IV SV IBA SBA LB LAI Reference

Zhao and Kane, 2012; Zhao et al., 2011; 2012;

Zhao et al., 2019, 2020; Zutter et al., 1998)
Platanus occidentalis L. * + − * + + (Domec et al., 2017)
Populus balsamifera L. × + * − + − − (Benomar et al., 2012)
P. trichocarpa Torr. & A.
Gray ex Hook
P. balsamifera L. × */+ * − + − − (Benomar et al., 2012; Remaury et al., 2019)
P. maximowiczii A. Henry
Populus × canadenses */− * − − (Di Matteo et al., 2012; Pannacci et al., 2009)
Moench
Populus × canadenses − − * (Truax et al., 2018)
Moench × P. maximowiczii
A. Henry
Populus deltoides W. Bartram * */− − − */+ (Fang et al., 2007; Rončević et al., 2013; Zhang
ex Marshall et al., 2020c)
P. deltoides W. Bartram ex * * − (Pannacci et al., 2009)
Marshall × P. ×
canadenses Moench
P. deltoides W. Bartram ex * − − * (Tun et al., 2018)
Marshall × P. ×
euramericana (Dode)
Guinier
P. deltoides W. Bartram ex */+ − (Toillon et al., 2013;Yáñez et al., 2019 )
Marshall × P. nigra L.
P. nigra L. × P. × generosa A. */− * − − (Di Matteo et al., 2012; Pannacci et al., 2009)
Henry
P. simonii Carr. × P. nigra L. − − − + (Ahmed et al., 2020)
Populus tomentosa Carr. * */− − * − − + (Han et al., 2020; Huijuan et al., 2020)
Populus trichocarpa Torr. & * − (Yáñez et al., 2019)
A.Gray ex Hook.
P. trichocarpa Torr. & A.Gray */− */− − + (Auclair and Bouvarel, 1992; DeBell and
ex Hook. × P. deltoides W. Harrington, 2002; Yáñez et al., 2019)
Bartram ex Marshall
P. trichocarpa Torr. & A.Gray * − (Yáñez et al., 2019)
ex Hook. ×
P. maximowiczii A. Henry
P. trichocarpa Torr. & A.Gray * − (Yáñez et al., 2019)
ex Hook. × P. nigra L. × P.
Populus × euramericana * − − */+ (Fang et al., 2007; Ning et al., 2019; Xue et al.,
(Dode) Guinier 2012; Zhang et al., 2020c )
Pseudotsuga menziesii (Mirb.) +/− */+/− +/− + − − */+ */+ (Cole and Newton, 1986; Harrington et al., 2009;
Franco Isaac-Renton et al., 2020; Marziliano et al., 2015,
2017; Peracca and O’Hara, 2008; Shainsky et al.,
1992; Scott et al., 1998; Woodruff et al., 2002)
Quercus acutissima Carruth. * − − + + (Cheng et al., 2020)
Quercus mongolica Fisch. ex − * − − (Imada et al., 1997)
Ledeb.
Quercus petraea (Matt.) */+ */− − * * (Andrzejczyk et al., 2015; Colin et al., 2012;
Liebl. Farque et al., 2001; Uhl et al. 2015)
Quercus robur L. − */+ − + − * (Gemmel et al., 1996; Uhl et al, 2015)
Quercus rugosa Nee. + − + (Uhl et al, 2015)
Robinia pseudoacacia L. * * − + (Rédei, 2009)
Salix spp. */+ (Larsen et al., 2019)
Salix alba L. − * * (Stolarski et al. 2017,2019a, 2019b ; Załuski
et al., 2018)
Salix dasyclados Wimm. * + (Baksiene and Titova, 2018)
Salix schwerinii E.L.Wolf − * + (Cao et al., 2012)
Salix schwerinii E. Wolf × * − + (Baksiene and Titova, 2018; Castaño-Díaz et al.,
Salix viminalis L. 2018)
Salix triandra L. × Salix * − + (Castaño-Díaz et al., 2018)
viminalis L.
Salix viminalis cv. Jorunn */− * */− − + (Bergkvist and Ledin, 1998; Bullard et al., 2002;
Stolarski et al., 2017, 2019a, 2019b; Załuski et al.,
2018)
Salix viminalis L. × (Salix * − + (Castaño-Díaz et al., 2018)
schwerinii E. Wolf × Salix
viminalis L.)
Sassafras tzumu (Hemsl.) − + (Sun et al., 2018)
Hemsl.
Sclerobium paniculatum − + − (Narducci et al., 2016)
Vogel
Sequoiadendron giganteum * − − + − − * + (O’Hara and York, 2014; Peracca and O’Hara,
(Lindl.) J. Buchholz 2008; York et al., 2013)
Tectona grandis L.f. + − + − − + * (Ola-Adams, 1993; Pachas et al., 2019)
Terminalia arjuna (Roxb. ex + + * + + + (Srivastava et al., 1999)
DC.) Wight & Arn.
(continued on next page)

8
M.M. Gabira et al.
Table 1 (continued )
Species S H DBH CH CD
Terminalia superba Engl. & − − − +
Diels
Thuja plicata Donn ex D.Don − (DeBell and Gartner, 1997)
Tsuga heterophylla [Raf.] * +
Sarg.
Ulmus pumila Walter * +
Zelkova serrata (Thunb.) − − − (Tanabe et al., 2019)
Makino
S, survival; H, total height; DBH, diameter at breast height; CH, crown height; CD, crown diameter; IV, individual volume; SV, stand volume; IBA, individual basal area;
SBA, stand basal area; LB, leaf biomass; LAI, leaf area index; *, neutral influence of increasing planting density; +, positive influence of increasing planting density; − ,
negative influence of increasing planting density.
Several shade-avoidance responses related to the production and
transport of auxin and cytokinin phytohormones drive tree growth and
development in high-density stands. Auxins are known to drive plant
growth and development—through the elongation of hypocotyl and
petiole—and upward leaf movements, enhancing access to light (Galvão
and Fankhauser, 2015). The biosynthesis and transport of auxins are
related to photoreceptor signaling (Küpers et al., 2020), and in high-
density stands, auxins stimulate height growth rather than stem lateral
growth and branch development in the lower portion of trees, which
define plant shape. Auxin metabolism and its influence on plant growth
and development was extensively reviewed by Casanova-Sáez and Voβ
(2019). Cytokinin influence on plant growth and acclimation to the light
gradient in high-density stands relate to differences in transpiration
rates in shaded and sun-exposed leaves. The reduced influx of cytokinins
in shaded leaves, resulting from a lower transpiration rate, decreases
protein synthesis and amino acid export, in turn inducing a reallocation
of resources to sun-exposed leaves, which become stronger sinks
because of higher transpiration rates (Pons et al., 2001). Because of
changes induced by these phytohormones, specific leaf area (SLA) is
reduced in trees subject to high-density spacing, although the leaf area
index (LAI) can increase in these circumstances (Parker, 2020).
In general, leaf expansion is reduced in shaded leaves and stimulated
in sunbathed ones, which directly influences the LAI and SLA (Pons
et al., 2001). As a measure of canopy closure, LAI is indicative of stand
productivity and biomass accumulation, and it can be an alternative
measure of growing space and stocking control in plantations (O’Hara
and York, 2014). However, few studies have considered this index
among measured growth variables. Only 24 studies in our survey (9 %,
Tab.1) considered LAI as a response variable to planting density, and all
of these studies revealed a neutral or positive relationship with density
(Akers et al., 2013; Bullard et al., 2002; Dean and Baldwin, 1996; Farooq
et al., 2020; Farque et al., 2001; Grigg et al., 2008; Hager and Kazda,
1985; Hakamada et al., 2020a; Henskens et al., 2001; Johansson, 1989,
2007; Meinzer et al., 1996; O’Hara and York, 2014; de Paula et al., 2013;
Pinkard and Neilsen, 2003; Shi and Cao, 1997; Schwerz et al., 2019,
2020b; Tenorio et al., 2019; Tun et al., 2018; Wang et al., 2019; Will
et al., 2001, 2005; Xue et al., 2012; Yan et al., 2019; Zhao et al., 2012).
LAI is often used as an indicator of canopy closure and light interception
in agricultural crops (Sharma-Natu and Ghildiyal, 2005). Nonetheless,
in forest plantations we must consider that plant (tree) density is
markedly different from that in agricultural plantations, and thus light
interception differs between forest and field settings. Hakamada et al.
(2020a) indicated that increased LAI (90 % higher than in wider spac­
ings) in clonal Eucalyptus high-density plantations was directly related to
light interception and consequently, to more efficient biomass accu­
mulation. The authors also observed that genotype-related characteris­
tics influenced the clonal response to planting density, which is a key
factor to be considered in breeding programs. Akers et al. (2013) ob­
tained comparable results for Pinus taeda L. plantations under various
planting densities. The authors attributed the increase in LAI (26 %
higher in high-density planting) and light interception to a greater
light-use efficacy in high-density stands relative to low-density stands.
Forest Ecology and Management 534 (2023) 120882
IV SV IBA SBA LB LAI Reference
* (Okojie et al., 1988; Ola-Adams, 1993)
+ (Negrave et al., 2007)
− + (Pérez et al., 2014)
Nonetheless, an increase in LAI or SLA is not always accompanied by
greater photosynthesis efficacy, as increased leaf expansion can produce
thicker leaves having a lower stomatal density and, consequently,
reduced gas exchange potential (Mcelwain et al., 2016; Thyroff et al.,
2019).
Competition for resources cannot always be managed by the addition
of water and/or nutrients or the application of other silvicultural
treatments, and the effects of resource management are more likely to be
observed for leaf, fruit, and seed yield than for wood (Kigalu, 2007a,
2007b; Mendieta-Araica et al., 2013; Narducci et al., 2016). For other
species, fruit and seed yields are not influenced by density, independent
of other resource management interventions, as observed for Pinus pinea
L. (Loewe-Muñoz et al., 2020). This limited influence of density in­
dicates a potential to manage spacing arrangements in interaction with
other silvicultural treatments to increase yield even when other re­
sources become scarce. Establishing high-density plantations for these
non-timber taxa in sites having a high availability of water and nutrients
is a strategy for increasing productivity by reducing competition be­
tween trees without increasing plantation costs (Harrington et al., 2009;
Sun et al., 2018; Toillon et al., 2013; Zhao et al., 2020).
The increase in total plantation biomass with concomitant reductions
in individual tree biomass is an important aspect to consider for forest
producers. There is a potential of producing wood for energy in high-
density stands; however, for more valuable wood products, such as
lumber and veneer, these stands should be submitted to thinning man­
agement to produce larger boles. At high initial densities, thinning may
be the best option for producers to ensure highly valuable wood prod­
ucts at the end of rotation by regulating timber form and volume
(Charlton et al., 2020). Smith et al. (2016) highlighted the need to
evaluate the economic aspects and potential use of thinned stems, and
Glencross et al. (2012) also suggested coupling high planting densities
with thinning treatments to obtain various products at different times to
ensure both a high volume for the stand and individual trees. Adapted
management strategies aim to ensure both wood and energy products in
forest plantations; these interventions include an increased tree density,
additional thinning, and a reduced rotation length to heighten financial
gains (Scott and Tiarks, 2008). Nonetheless, in plantations characterized
by wider spacings or that are subjected to thinning, wind can penetrate
more easily, increasing mechanical stresses on residual trees and
increasing tree vulnerability to windthrow (Girona et al., 2019; Scott
and Mitchell, 2005).
Another important alternative to increase biomass production in
forest stands is species mixture, which create distinct interactions be­
tween individuals and provide growth advantages (Bello et al., 2019). As
an example, Forrester et al. (2004) observed that biomass accumulation
in E. globulus increased when in admixture with A. mearnsii. In contrast,
Bouillet et al. (2013) observed that changes in competitive or facilitation
processes in monospecific stands or in mixed-stands of Eucalyptus spp.
and A. mangium depended on ecological conditions, indicating the
importance of evaluating each species responses and other environ­
mental aspects such as water and nutrient availability. In our review, we
did not consider papers evaluating plant growth in mixed stands
9
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882

considering that they have similarities and differences, resulting re­ Table 2
sponses that could not be compared to studies evaluating monospecific Summary of studies (2001–2020) focused on the physiological response of a tree
stands; however, we recognise and encourage the development of a re­ species to increased planting density.
view evaluating the influence of mixed stands on tree growth. Species IR A E R WUE LWP Cl Reference
Although we consider the constant final yield as an index of the Acacia koa A. + + (Meinzer
maximum planting density in a stand, most studies evaluated in this Gray et al., 1996)
review did not attain a planting density that would be considered Acacia mangium + (Cienciala
limiting for tree growth. A possibility is that most researchers focused on Willd et al., 2000)
Acacia mearnsii De * (Schwerz
planting densities that achieved an increase in stand biomass, all the
Wild. et al., 2019)
while considering the production of high-quality wood products. Alnus rubra Bong. − (Cole and
Nonetheless, the risks of drought-induced problems for tree survival and Newton,
growth may increase above densities of 1,000 trees⋅ha− 1 (Hakamada 2019)
et al., 2020b), and lower stocking plantations could be a strategy to Ateleia glazioviana * (Schwerz
Baill. et al., 2020b)
improve the balance between sustaining wood production and main­ Camellia sinensis L. − + (Kigalu,
taining ecosystem services in forest landscapes. In some studies, both 2007a)
individual and stand volumes did not present a linear response to Eucalyptus * (Henskens
increasing stand density, indicating that increased planting densities did globulus Labill. et al., 2001)
Eucalyptus grandis * (Eastham
not always heighten biomass productivity (Gadow and Kotze, 2014; − +
W. Hill ex et al., 1990;
Guner et al., 2010; Marziliano et al., 2015; Özbayram and Çiçek, 2020; Maiden Schwerz
Tavares et al., 2020; Węgiel and Polowy, 2020). It is important to note et al., 2020b)
that most plantations are not fertilized during the entire rotation; E. grandis W. Hill + * − (Hakamada
therefore, a lower planting density would also reduce soil nutrient ex Maiden × et al., 2017,
E. camaldulensis 2020a,
extraction (Medeiros et al., 2020).
Dehnh. 2020b)
E. grandis W. Hill + * − (Hakamada
3.3. How does planting density affect tree physiological responses? ex Maiden × et al., 2017,
E. urophylla S.T. 2020a,
Blake 2020b)
Physiological evaluations in response to tree density are less com­
Eucalyptus + * − (Hakamada
mon relative to dendrological measurements, likely because they are urophylla S.T. et al., 2017,
more complicated to undertake in the field. Furthermore, these physi­ Blake 2020a,
ological studies particularly focused on the influence of light and water 2020b)
on trees (Table 2). Fagus orientalis L. * (Guner et al.,
2010)
The first and most obvious influence of closer plant spacing is Fagus sylvatica L. + (Gemmel
reduced light interception, which directly influences the physiological et al., 1996)
functions of a plant. In a study of Pinus plantations of varying density, Leucaena * + (Pachas
Will et al. (2001) observed that tree growth correlated with intercepted leucocephala et al., 2018)
(Lam.) de Wit
radiation; as stand density increased, the amount of radiation inter­
Mimosa scabrella * (Schwerz
cepted per unit of leaf area decreased because of self-shading. The au­ Benth. et al., 2020b)
thors observed that differences in net photosynthesis and respiration Moringa oleifera * * * (Mabapa
rates were minimal and did not correlate with growth at the various Lam. et al., 2018)
stand densities, possibly because the resources necessary for the plants’ Pinus elliottii + * + (Will et al.,
Engelm. 2001)
physiological activities were never limiting in any treatment. Toillon Pinus sibirica Du + (Milyutina
et al. (2013) observed that elevated planting densities increased height Tour et al., 2013)
growth (14 % higher in high planting density, in one study site) rather Pinus sylvestris L. + (Milyutina
than larger stem diameters (10–31 % lower in high planting densities) et al., 2013)
Pinus taeda L. */+ * (Akers et al.,
probably because of an accentuated competition for light in poplar + −
2013;
(Populus spp.) plantations, although the authors did not evaluate inter­ Ingwers
cepted radiation. Trees planted closer together experience a lower red: et al., 2018;
far-red light ratio, indicating a competition for light and a decrease in Will et al.,
the total light intensity within the stand (Huber et al., 2021); however, 2001, 2005)
Populus (Benomar
only Benomar et al. (2011) observed a decrease in C assimilation related
+ −
balsamifera L. × et al., 2011)
to high planting density among the studies in our review, a reduction of P. trichocarpa
23 % in C assimilation compared to plants in wider spacings. Torr. & A. Gray
Direct measurements of transpiration rate gradients in forest can­ ex Hook
Populus deltoides (Toillon
opies are quite limited, although there is some evidence that this −
W. Bartram ex et al., 2013)
physiological parameter varies with light and growth rates, and species Marshall ×
(Pons et al., 2001). Transpiration in Eucalyptus plantations and respi­ P. nigra L.
ration in Pinus plantations have been shown to increase with a higher Populus + − (Benomar
planting density, a pattern related to the water-use efficiency of each maximowiczii A. et al., 2011)
Henry ×
genotype and to the microclimate created by the proximity of in­ P. balsamifera L.
dividuals (Hakamada et al., 2017, 2020a, 2020b; Will et al., 2001); Pseudotsuga + (Cole and
however, it does not mean that plants in this condition will be more menziesii Newton,
susceptible to droughts because other water-related process also influ­ (Mirb.) Franco 1986)
Quercus robur L. (Gemmel
ence water availability, as interspecies interactions, understory cover, +
et al., 1996)
and soil type. Moore et al. (2011) demonstrated that the greatest vari­ Tsuga heterophylla − (Cole and
ability in both productivity and transpiration in Pseudotsuga menziesii [Raf.] Sarg. Newton,
and Alnus rubra plantations (1100 trees ha− 1) was mainly determined by 2019)

10
M.M. Gabira et al.
IR, intercepted radiation; A, carbon assimilation; E, transpiration; R, respiration;
WUE, water-use efficiency; LWP, leaf water potential; Cl, chlorophyll content; *,
neutral influence of increasing planting density; +, positive influence of
increasing planting density; − , negative influence of increasing planting density.
site conditions. Camellia sinensis L. demonstrated increased water-use
efficiency in high-density stands as transpiration rates decreased, and
the response of Eucalyptus clones varied according to genotype (Haka­
mada et al., 2020a, 2020b; Kigalu, 2007a). Forrester (2015) explained
that there is a complementarity effect of transpiration and water use
efficiency in forest plantations that influences plant growth.
Increasing planting density may also have implications for ecosystem
water balance, as high-density stands require greater water resources at
the landscape scale (Forrester, 2015). In this scenario, the increase in
drought events due to climate changes has induced mortality in forest
plantations worldwide (Klein and Hartmann, 2018). Looney and Zhang
(2022) commented that climate change–driven wildfires have increased
in frequency and extent; researchers have advocated manipulating the
spacing in plantings during reforestation as a means of enhancing
structural complexity to improve stand resistance and resilience against
fires. According to Campo et al. (2022), thinning can be an effective
means of helping the remaining trees to cope with climate change im­
pacts, e.g., more frequent droughts and wildfires, as thinning ensures a
more hydrated forest system. In the face of worsening climate-change
scenarios, Ungar et al. (2013) indicated that the Pinus halepensis Mill.
(Aleppo pine) forest in Israel’s semiarid northern Negev (300 trees ha− 1)
should be thinned to stands of 250 or 190 trees ha− 1 in order to remain
sustainable under annual rainfall regimes of 200 or 150 mm, respec­
tively. For Hevea brasiliensis monocultures, Hardanto et al. (2017)
concluded that stand density (300 to 668 trees ha− 1) played a prominent
role as a potential controlling factor of stand water use. Thus, thinning
measures appear as a potentially viable management option for influ­
encing water uptake patterns and stand transpiration rates in favor of
cultivated rubber trees.
Furthermore, some authors suggest that mixing species or reducing
planting density would increase stand resilience to drought, although
results demonstrate that it depends on what species are used, how the
stand is managed, what is the growth rate, and availability of other re­
sources (Forrester et al., 2004; Forrester, 2015; Tsamir et al., 2019).
Under severe soil water deficit conditions, Grossiord et al. (2014)
highlighted an interesting differential response pattern in terms of
transpiration and water use efficiency between Quercus cerris and
Quercus petraea when they are growing in pure plots or when they are
competing for water resources in mixed species conditions. Q. cerris was
negatively impacted in its water use when growing in mixed conditions
with Q. petraea (689 trees ha− 1), an oak species that is known to be able
to extract water from deep soil layers.
Differences in characteristics such as drought resistance, the number
of stomata, and leaf type can affect physiological responses in high-
density stands (Hakamada et al., 2020b), however, according to
Resquin et al. (2019) and Will et al. (2005), the physiological response of
trees in high-density stands is mainly mediated by competition for re­
sources, indicating the importance of resource availability and under­
standing the species-specific resource requirements for monospecific
forest plantations. Understanding this physiological dynamic is funda­
mental for planning appropriate management scenarios and for
obtaining high-value products (Forrester, 2015). Several studies selected
in this review demonstrate that site characteristics have a major effect
on tree growth and physiological response to planting density. This
identified importance of site characteristics and studies detailing tree
physiological response to planting density indicate the critical need for
physiological evaluations in field trials to develop specific management
programs tailored to individual species (Akers et al., 2013; Guner et al.,
2010; Toillon et al., 2013; Will et al., 2005, 2001).
11
Forest Ecology and Management 534 (2023) 120882
4. Final considerations
Tree density and spacing are two key elements in plantation design;
and understanding their impact on plant growth and development is
vital. We found an increase in the number of studies focused on plant
morphological and physiological responses to planting density in forest
plantations over the last 40 years. Most studies occurred in countries
having well-established markets for timber, pulp and paper, and energy,
and a high number of these studies focused on Pinus and Eucalyptus
species. The most common variables evaluated in field trials were DBH
and timber volume, reflecting the dominant focus of these studies on
timber production. We, therefore, highlight the need to develop more
research on the production of other forest products, such as leaves,
fruits, and seeds, and we underscore the need to consider the use of
physiological evaluations in field trials to provide more accurate re­
sponses of plants to resource competition, tree thinning, and mainly site-
specific characteristics.
Increasing planting density in forest plantations is important in re­
gions where forest plantations compete with other high-value crops or
where space is a limiting factor for the development of the forest in­
dustry. High-density forest stands can increase the value and yield for a
variety of products; however, we still need to improve our under­
standing of trees grown for products other than timber. Finally, when
planning for planting density within forest plantations, water, nutrient,
and carbon-balance effects on tree growth and physiology must be
assessed and understood. The results of this survey of existing literature
can serve to improve forest management decisions in regard to forest
stand use and develop novel study avenues for silviculture.
CRediT authorship contribution statement
Mônica Moreno Gabira: Conceptualization, Methodology, Formal
analysis, Investigation, Data curation, Writing – original draft, Writing –
review & editing, Visualization. Miguel Montoro Girona: Methodol­
ogy, Resources, Writing – review & editing, Supervision. Annie Des­
Rochers: Methodology, Resources, Writing – review & editing,
Supervision. Dagma Kratz: Writing – original draft, Writing – review &
editing. Richardson Barbosa Gomes da Silva: Formal analysis,
Investigation, Data curation, Writing – original draft, Writing – review &
editing. Manoela Mendes Duarte: Methodology, Investigation, Data
curation, Writing – original draft. Natalia Saudade de Aguiar: Meth­
odology, Investigation, Data curation, Writing – original draft. Ivar
Wendling: Conceptualization, Resources, Writing – review & editing,
Supervision, Project administration.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgements
This project was funded by the Ph.D. scholarship of the Coordination
for the Improvement of Higher Education Personnel of Brazil (Capes
–financing code 001) obtained by Gabira MM, the Ministry of Interna­
tional Relations and La Francophonie of Québec (MRiF – Call for Pro­
jects Québec-Brésil 2019-2021 and 2021-2023) obtained by Gabira MM,
Girona MM, and DesRochers A for a research internship scholarship at
GREMA in the Québec University at Abitibi-Temiscamingue (UQAT),
and the NSERC–Alliance – Silviculture grant UQAT-UQAC ALLRP
557166–20 obtained by Girona MM.
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882

References Bohanek, J.R., Groninger, J.W., 2003. Impacts of intensive management on black walnut
(Juglans nigra L.) growth and bole quality at mid-rotation. For. Sci. 49, 522–529.
Bongers, F.J., Pierik, R., Anten, N.P.R., Evers, J.B., 2018. Subtle variation in shade
Ahmed, A.K.M., Fu, Z., Ding, C., Jiang, L., Han, X., Yang, A., Ma, Y., Zhao, X., 2020.
avoidance responses may have profound consequences for plant competitiveness.
Growth and wood properties of a 38-year-old Populus simonii × P. nigra plantation
Ann. Bot. (Lond.) 121, 863–873.
established with different densities in semi-arid areas of northeastern China. J. For.
Boreham, G.R., Pallett, R.N., 2009. The influence of tree improvement and cultural
Res. 31, 497–506.
practices on the productivity of Eucalyptus plantations in temperate South Africa.
Akers, M.K., Kane, M., Zhao, D., Teskey, R.O., Daniels, R.F., 2013. Effects of planting
South. For. 71, 85–93.
density and cultural intensity on stand and crown attributes of mid-rotation loblolly
Bose, A.K., Nelson, A.S., Kane, M., Rigling, A., 2018. Density reduction in loblolly pine
pine plantations. For. Ecol. Manage. 310, 468–475.
(Pinus taeda L.) stands to increase tree C assimilation: an approach with the dual δ13C
Akhtar, J., Saqib, Z.A., Qureshi, R.H., Haq, M.A., Iqbal, M.S., Marcar, N.E., 2008. The
and δ18O isotope signatures in needles. Ann. For. Sci. 75, 8.
effect of spacing on the growth of Eucalyptus camaldulensis on salt-affected soils of
Bouillet, J.-P., Laclau, J.-P., Gonçalves, J.L.D.M., Voigtlaender, M., Gava, J.L., Leite, F.P.,
the Punjab. Pakistan. Can. J. For. Res. 38, 2434–2444.
Hakamada, R., Mareschal, L., Mabiala, A., Tardy, F., Levillain, J., Deleporte, P.,
Albaugh, T.J., Maier, C.A., Campoe, O.C., Yáñez, M.A., Carbaugh, E.D., Carter, D.R.,
Epron, D., Nouvellon, Y., 2013. Eucalyptus and Acacia tree growth over entire
Cook, R.L., Rubilar, R.A., Fox, T.R., 2020. Crown architecture, crown leaf area
rotation in single-and mixed-species plantations across five sites in Brazil and Congo.
distribution, and individual tree growth efficiency vary across site, genetic entry,
For. Ecol. Manag. 301, 89–101.
and planting density. Trees (Berl.) 34, 73–88.
Bravo-Oviedo, A., Pretzch, H., del Río, M., 2018. Dynamics, silviculture and management
Alcorn, P.J., Pyttel, P., Bauhus, J., Smith, R.G.B., Thomas, D., James, R., Nicotra, A.,
of mixed forests. In: Tomé, M., Seifert, T., Kurttila, M. (Eds.), Managing Forest
2007. Effects of initial planting density on branch development in 4-year-old
Ecosystems. Springer International Publishing, Cham, Switzerland, p. 420.
plantation grown Eucalyptus pilularis and Eucalyptus cloeziana trees. For. Ecol.
Buford, M.A., 1991. Performance of four yield models for predicting stand dynamics of a
Manage. 252 (1), 41–51.
30-year-old loblolly pine (Pinus taeda L.) spacing study. For. Ecol. Manage. 46,
Amateis, R.L., Burkhart, H.E., 2013. Relating quantity, quality, and value of lumber to
23–38.
planting density for loblolly pine plantations. South. J. Appl. For. 37, 97–101.
Bullard, M.J., Mustill, S.J., Carver, P., Nixon, P.M.I., 2002. Yield improvements through
Andrianirina, Z.T., Martin, M., Dongmeza, E., Senger, E., 2019. Effects of genotype,
modification of planting density and harvest frequency in short rotation coppice
direct sowing and plant spacing on field performance of Jatropha curcas L. Agronomy
Salix spp. - 2. Resource capture and use in two morphologically diverse varieties.
(Basel) 9, 465.
Biomass Bioenergy 22 (1), 27–39.
Andrzejczyk, T., Liziniewicz, M., Drozdowski, S., 2015. Effect of spacing on growth and
Burkes, E.C., Will, R.E., Barron-Gafford, G.A., Teskey, R.O., Shiver, B., 2003. Biomass
quality parameters in sessile oak (Quercus petraea) stands in central Poland: results 7
partitioning and growth efficiency of intensively managed Pinus taeda and Pinus
years after planting. Scand. J. For. Res. 30, 710–718.
elliottii stands of different planting densities. For. Sci. 49, 225–234.
Antón-Fernández, C., Burkhart, H.E., Strub, M., Amateis, R.L., 2011. Effects of initial
Cao, Y., Lehto, T., Piirainen, S., Kukkonen, J.V.K., Pelkonen, P., 2012. Effects of planting
spacing on height development of loblolly pine. For. Sci. 57, 201–211.
orientation and density on the soil solution chemistry and growth of willow cuttings.
Antón-Fernández, C., Burkhart, H.E., Amateis, R.L., 2012. Modeling the effects of initial
Biomass and Bioenergy 46, 165–173.
spacing on stand basal area development of loblolly pine. For. Sci. 58, 95–105.
Cardoso, D.J., Lacerda, A.E.B., Rosot, M.A.D., Garrastazú, M.C., Lima, R.T., 2013.
Arias-Aguilar, D., Valverde, J.C., Campos, R., 2020. Effect of planting density and tree
Influence of spacing regimes on the development of loblolly pine (Pinus taeda L.) in
species selection on forest bioenergy systems: tree growth, nutrient storage and
Southern Brazil. For. Ecol. Manage. 310, 761–769.
wood chemical properties. Greenh. Gases Sci. Technol. 10, 1165–1175.
Casanova-Sáez, R., Voß, U., 2019. Auxin metabolism controls developmental decisions in
Aspinwall, M.J., King, J.S., Mckeand, S.E., Bullock, B.P., 2011. Genetic effects on stand-
land plants. Trends Plant Sci. 24, 741–754.
level uniformity and above-and belowground dry mass production in juvenile
Cassidy, M., Palmer, G., Smith, R.G.B., 2013. The effect of wide initial spacing on wood
loblolly pine. For. Ecol. Manage. 262, 609–619.
properties in plantation grown Eucalyptus pilularis. New For. 44, 919–936.
Auclair, D., Bouvarel, L., 1992. Influence of spacing and short rotations on Populus
Castaño-Díaz, M., Barrio-Anta, M., Afif-Khouri, E., Cámara-Obregón, A., 2018. Willow
trichocarpa × deltoides coppice. Can. J. For. Res. 22, 541–548.
short rotation coppice trial in a former mining area in Northern Spain: effects of
Bai, Y.F., Shen, Y.Y., Jin, Y.D., Hong, Y., Liu, Y.Y., Li, Y.Q., Liu, R., Zhang, Z.W., Jiang, C.
clone, fertilization and planting density on yield after five years. Forests 9, 154.
Q., Wang, Y.J., 2020. Selective thinning and initial planting density management
Celentano, D., Moraes, M., Ferreira, J., Nahur, A., Coutinho, B., Rousseau, G.X.,
promote biomass and carbon storage in a chronosequence of evergreen conifer
Martins, M.B., Vasconcelos, L.G.T.R., Rodrigues, F., Freire, R., Pinto, A.,
plantations in Southeast China. Glob. Ecol. Conserv. 24, e01216.
Kanashiro, M., 2022. Forest restoration to promote a fair post COVID-19 recovery in
Baksiene, E., Titova, J., 2018. Effects of cultivation technologies on woody biomass yield
the Brazilian Amazon. Land Use Policy 116, 106076.
of various willow (Salix spp.) cultivars. Zemdirbyste-Agriculture 105, 339–348.
Charlton, R.A., Naghizadeh, Z., Ham, C., Wessels, C.B., 2020. A value chain comparison
Baldwin, V.C., Peterson, K.D., Clark, A., Ferguson, R.B., Strub, M.R., Bower, D.R., 2000.
of Pinus patula sawlog management regimes based on different initial planting
The effects of spacing and thinning on stand and tree characteristics of 38-year-old
densities and effect on wood quality. For. Policy Econ. 111, 102067.
Loblolly Pine. For. Ecol. Manag. 137, 91–102.
Chase, C.W., Kimsey, M.J., Shaw, T.M., Coleman, M.D., 2016. The response of light,
Barbier, N., Couteron, P., Proisy, C., Malhi, Y., Gastellu-Etchegorry, J.P., 2010. The
water, and nutrient availability to pre-commercial thinning in dry inland Douglas-fir
variation of apparent crown size and canopy heterogeneity across lowland
forests. For. Ecol. Manage. 363, 98–109.
Amazonian forests. Glob. Ecol. Biogeogr. 19, 72–84.
Cheng, X., Yuan, H., Xing, W., Wang, Y., Yu, M., 2020. Effects of provenance and initial
Barron-Gafford, G.A., Will, R.E., Burkes, E.C., Shiver, B., Teskey, R.O., 2003. Nutrient
planting density on growth and wood properties in young sawtooth oak (Quercus
concentrations and contents, and their relation to stem growth, of intensively
acutissima) plantations. Eur. J. For. Res. 139, 1067–1078.
managed Pinus taeda and Pinus elliottii stands of different planting densities. For. Sci.
Çiçek, E., Yilmaz, F., Tilki, F., Cicek, N., 2010. Effects of spacing and post-planting
49, 291–300.
treatments on survival and growth of Fraxinus angustifolia seedlings. J. Environ. Biol.
Basra, S.M.A., Nouman, W., Hafeez-ur-Rehman, U.M., Nazli, Z.H., 2015. Biomass
31, 515–519.
production and nutritional composition of Moringa oleifera under different cutting
Cienciala, E., Kučera, J., Malmer, A., 2000. Tree sap flow and stand transpiration of two
frequencies and planting spacings. Int. J. Agric. Biol. 17, 1055–1060.
Acacia mangium plantations in Sabah. Borneo. J. Hydrology 236, 109–120.
Begon. M., Townsend CR 2020 Ecology : from individuals to ecosystems. John Wiley &
Clark, A., Lewis, I., Schimleck, J.L., Daniels, R.F., 2008. Effect of initial planting spacing
Sons, Inc, Hoboken, NJ, p 864.
on wood properties of unthinned loblolly pine at age 21. For. Prod. J. 58, 78–83.
Bello, J., Vallet, P., Perot, T., Balandier, P., Seigner, V., Perret, S., Couteau, C.,
Cole, E.C., Newton, M., 1986. Nutrient, moisture, and light relations en 5-year-old
Korboulewsky, N., 2019. How do mixing tree species and stand density affect
Douglas-fir plantations under variable competition. Can. J. For. Res. 16, 727–732.
seasonal radial growth during drought events? For. Ecol. Manag. 432, 436–445.
Cole, E.C., Newton, M., 2019. Plant water stress and soil depletion in variable-density,
Benin, C.C., Wionzek, F.B., Watzlawick, L.F., 2014. Initial assessments on the plantation
red alder/western hemlock coastal Oregon plantations. For. Sci. 66, 304–313.
of Eucalyptus benthamii Maiden et Cambage deployed in different spacing. Brazilian
Colin, F., Ningre, F., Fortin, M., Huet, S., 2012. Quantification of Quercus petraea Liebl.
J. Appl. Technol. Agric. Sci. 7, 55–61.
forking based on a 23-year-long longitudinal survey. For. Ecol. Manage. 282,
Benomar, L., Desrochers, A., Larocque, G.R., 2011. Changes in specific leaf area and
133–141.
photosynthetic nitrogen-use efficiency associated with physiological acclimation of
Coordes, R., 2013. Influence of planting density and rotation age on the profitability of
two hybrid poplar clones to intraclonal competition. Can. J. For. Res. 41,
timber production for Norway spruce in Central Europe. Eur. J. For. Res. 132,
1465–1476.
297–311.
Benomar, L., DesRochers, A., Larocque, G.R., 2012. The effects of spacing on growth,
Crous, J., Burger, L., Sale, G., 2013. Growth response at age 10 years of five Eucalyptus
morphology and biomass production and allocation in two hybrid poplar clones
genotypes planted at three densities on a drought-prone site in KwaZulu-Natal, South
growing in the boreal region of Canada. Trees (Berl.) 26, 939–949.
Africa. South. For. 75, 189–198.
Bentancor, L., Hernández, J., del Pino, A., Califra, Á., Resquín, F., González-Barrios, P.,
Crous, J., Sale, G., Naidoo, T., 2019. The influence of species, tree improvement and
2019. Evaluation of the biomass production, energy yield and nutrient removal of
cultural practices on rotation-end fibre production of Eucalyptus pulpwood
Eucalyptus dunnii Maiden grown in short rotation coppice under two initial planting
plantations in South Africa. South. For. 81, 307–317.
densities and harvest systems. Biomass Bioenergy 122, 165–174.
da Páscoa, K.J.V., Scolforo, J.R.S., Ferraz Filho, A.C., Altoé, T.F., Gomide, L.R., 2019.
Bergkvist, P., Ledin, S., 1998. Stem biomass yields at different planting designs and
Growth of Eremanthus erythropappus (DC.) MacLeish in different planting spacings.
spacings in willow coppice systems. Biomass and Bioenergy 14, 149–156.
Floresta Ambient. 26, e20170834.
Bernardo, A.L., Reis, M.G.F., Reis, G.G., Harrison, R.B., Firme, D.J., 1998. Effect of
Dassot, M., Constant, T., Ningre, F., Fournier, M., 2015. Impact of stand density on tree
spacing on growth and biomass distribution in Eucalyptus camaldulensis, E. pellita and
morphology and growth stresses in young beech (Fagus sylvatica L.) stands. Trees
E. urophylla plantations in southeastern Brazil. For. Ecol. Manag. 104, 1–13.
(Berl.) 29, 583–591.
Binkley, D., Campoe, O.C., Alvares, C., Carneiro, R.L., Cegatta, Í., Stape, J.L., 2017. The
Dean, T.J., Baldwin, V.C., 1996. Growth in loblolly pine plantations as a function of stand
interactions of climate, spacing and genetics on clonal Eucalyptus plantations across
density and canopy properties. For. Ecol. Manage. 82, 49–58.
Brazil and Uruguay. For. Ecol. Manage. 405, 271–283.

12
M.M. Gabira et al.
DeBell, J.D., Gartner, B.L., 1997. Stem characteristics on the lower log of 35-year-old
western redcedar grown at several spacings. West. J. Appl. For. 12, 9–14.
DeBell, D.S., Harrington, C.A., 2002. Density and rectangularity of planting influence 20-
year growth and development of red alder. Can. J. For. Res. 32, 1244–1253.
del Campo, A.D., Otsuki, K., Serengil, Y., Blanco, J.A., Yousefpour, R., Wei, X., 2022.
A global synthesis on the effects of thinning on hydrological processes: implications
for forest management. For. Ecol. Manage. 519, 120324.
Di Matteo, G., Sperandio, G., Verani, S., 2012. Field performance of poplar for bioenergy
in southern Europe after two coppicing rotations: effects of clone and planting
density. IForest 5, 224–229.
Domec, J.C., Ashley, E., Fischer, M., Noormets, A., Boone, J., Williamson, J.C., King, J.S.,
2017. Productivity, biomass partitioning, and energy yield of low-input short-
rotation american sycamore (Platanus occidentalis L.) grown on marginal land:effects
of planting density and simulated drought. Bioenergy Res. 10, 903–914.
du Plessis, M., Kotze, H., 2011. Growth and yield models for Eucalyptus grandis grown in
Swaziland. South. For. 73, 81–89.
Duan, A., Lei, J., Hu, X., Zhang, J., Du, H., Zhang, X., Guo, W., Sun, J., 2019. Effects of
planting density on soil bulk density, pH and nutrients of unthinned Chinese fir
mature stands in south subtropical region of China. Forests 10, 351.
Eastham, J., Rose, C.W., Charles-Edwards, D.A., Cameron, D.M., Rance, S.J., 1990.
Planting density effects on water use efficiency of trees and pasture in an
agroforestry experiment. New Zealand J. For. Sci. 20, 39–53.
Egbäck, S., Liziniewicz, M., Högberg, K.A., Ekö, P.M., Nilsson, U., 2012. Influence of
progeny and initial stand density on growth and quality traits of 21 year old half-sib
Scots pine (Pinus sylvestris L.). For. Ecol. Manage. 286, 1–7.
Egbäck, S., Bullock, B.P., Isik, F., McKeand, S.E., 2015. Height-diameter relationships for
different genetic planting stock of loblolly pine at age 6. For. Sci. 61, 424–428.
Eloy, E., da Silva, D.A., Schmidt, D., Trevisan, R., Caron, B.O., Elli, E.F., 2016. Effect of
planting age and spacing on energy properties of Eucalyptus grandis W. Hill ex
Maiden. Rev. Árvore 40, 749–758.
Eloy, E., da Silva, D.A., Caron, B.O., Elli, E.F., Schwerz, F., 2018. Effect of age and
spacing on biomass production in forest plantations. Rev. Árvore 42, e420214.
Erasmus, J., Wessels, C.B., 2020. The effect of stand density management on Pinus patula
lumber properties. Eur. J. For. Res. 139, 247–257.
Erasmus, J., Kunneke, A., Drew, D.M., Wessels, C.B., 2018. The effect of planting spacing
on Pinus patula stem straightness, microfibril angle and wood density. Forestry 91,
247–258.
Fang, S., Xue, J., Tang, L., 2007. Biomass production and carbon sequestration potential
in poplar plantations with different management patterns. J. Environ. Manage. 85,
672–679.
Food and Agriculture Organization of the United Nations (FAO) (2020a) Global forest
resources assessment 2020: Main report. Food and Agriculture Organization of the
United Nations, Rome, Italy, p 186.
Farooq, T.H., Yan, W., Chen, X., Shakoor, A., Rashid, M.H.U., Gilani, M.M., He, Z.,
Wu, P., 2020. Dynamics of canopy development of Cunninghamia lanceolata mid-age
plantation in relation to foliar nitrogen and soil quality influenced by stand density.
Glob. Ecol. Conserv. 24, e01209.
Farque, L., Sinoquet, H., Colin, F., 2001. Canopy structure and light interception in
Quercus petraea seedlings in relation to light regime and plant density. Tree Physiol.
21, 1257–1267.
Feng, H., Xue, L., 2019. Competition-density effect of tree organs in Acacia auriculiformis
stands. J. For. Res. 30, 891–898.
Fernández-Núñez, E., Rigueiro-Rodríguez, A., Mosquera-Losada, M.R., 2010. Carbon
allocation dynamics one decade after afforestation with Pinus radiata D. Don and
Betula alba L. under two stand densities in NW Spain. Ecol. Eng. 36, 876–890.
Ferraz Filho, A.C., Mola-Yudego, B., González-Olabarria, J.R., Scolforo, J.R.S., 2018.
Thinning regimes and initial spacing for Eucalyptus plantations in Brazil. An. Acad.
Bras. Cienc. 90, 255–265.
Ferreira, G.W.D., Rau, B.M., Aubrey, D.P., 2020. Herbicide, fertilization, and planting
density effects on intensively managed loblolly pine early stand development. For.
Ecol. Manage. 472.
Finley, K., Zhang, J., 2019. Climate effect on ponderosa pine radial growth varies with
tree density and shrub removal. Forests 10, 477.
Forrester, D.I., 2015. Transpiration and water-use efficiency in mixed-species forests
versus monocultures: effects of tree size, stand density and season. Tree Physiol. 35,
289–304.
Forrester, D.I., Smith, R.G.B., 2012. Faster growth of Eucalyptus grandis and Eucalyptus
pilularis in mixed-species stands than monocultures. For. Ecol. Manage. 286, 81–86.
Forrester, D.I., Bauhus, J., Khanna, P.K., 2004. Growth dynamics in a mixed-species
plantation of Eucalyptus globulus and Acacia mearnsii. For. Ecol. Manage. 193, 81–95.
Forrester, D.I., Wiedemann, J.C., Forrester, R.I., Baker, T.G., 2013. Effects of planting
density and site quality on mean tree size and total stand growth of Eucalyptus
globulus plantations. Can. J. For. Res. 43, 846–851.
Gadow, K.V., Kotze, H., 2014. Tree survival and maximum density of planted forests –
Observations from South African spacing studies. For. Ecosyst. 1, 21.
Galvão, V.C., Fankhauser, C., 2015. Sensing the light environment in plants:
photoreceptors and early signaling steps. Curr. Opin. Neurobiol. 34, 46–53.
Gemmel, P., Nilsson, U., Welander, T., 1996. Development of oak and beech seedlings
planted under varying shelterwood densities and with different site preparation
methods in southern Sweden. New For. 12, 141–161.
Girona, M.M., Morin, H., Lussier, J.M., Ruel, J.C., 2019. Post-cutting mortality following
experimental silvicultural treatments in unmanaged boreal forest stands. Front. For.
Glob. Change 2, 4.
Gizachew, B., Brunner, A., Øyen, B.H., 2012. Stand responses to initial spacing in
Norway spruce plantations in Norway. Scand. J. For. Res. 27, 637–648.
13
Forest Ecology and Management 534 (2023) 120882
Glencross, K., Nichols, J.D., Grant, J.C., Sethy, M., Smith, R.G.B., 2012. Spacing affects
stem form, early growth and branching in young whitewood (Endospermum
medullosum) plantations in Vanuatu. Int. For. Rev. 14, 442–451.
Goel, V.L., Behl, H.M., 2004. Productivity assessment of three leguminous species under
high-density plantations on degraded soil sites. Biomass and Bioenergy 27, 403–409.
Gonçalves, J.L.D.M., Stape, J.L., Laclau, J.-P., Smethurst, P., Gava, J.L., 2004.
Silvicultural effects on the productivity and wood quality of eucalypt plantations.
For. Ecol. Manage. 193, 45–61.
Grigg, A.H., Macfarlane, C., Evangelista, C., Eamus, D., Adams, M.A., 2008. Does initial
spacing influence crown and hydraulic architecture of Eucalyptus marginata? Tree
Physiol. 28, 753–760.
Groot, A., Cortini, F., 2016. Effects of initial planting density on tree and stand
development of planted black spruce up to age 30. For. Chron. 92, 200–209.
Grossiord, C., Gessler, A., Granier, A., Pollastrini, M., Bussotti, F., Bonal, D., 2014.
Interspecific competition influences the response of oak transpiration to increasing
drought stress in a mixed Mediterranean forest. For. Ecol. Manage. 318, 54–61.
Guner, S., Yagci, V., Tilki, F., Celik, N., 2010. The effects of initial planting density on
above-and below-ground biomass in a 25-year-old Fagus orientalis Lipsky plantation
in Hopa. Turkey. Sci. Res. Essays 5, 1856–1860.
Gyenge, J.E., Fernández, M.E., Schlichter, T.M., 2010. Effect of stand density and
pruning on growth of ponderosa pines in NW Patagonia. Argentina. Agrofor. Syst.
78, 233–241.
Hager, H., Kazda, M., 1985. The influence of stand density and canopy position on sulfur
content in needles of Norway spruce (Picea abies L. Karst.). Water, Air, and Soil
Pollution 25, 321–329.
Hakamada RE, Hubbard RM, Stape JL, Lima W de P, Moreira GG, Ferraz SF de B 2020b
Stocking effects on seasonal tree transpiration and ecosystem water balance in a fast-
growing Eucalyptus plantation in Brazil. For. Ecol. Manage. 466:118149.
Hakamada, R., Hubbard, R.M., Ferraz, S., Stape, J.L., Lemos, C., 2017. Biomass
production and potential water stress increase with planting density in four highly
productive clonal Eucalyptus genotypes. South. For. 79, 251–257.
Hakamada, R.E., Hubbard, R.M., Moreira, G.G., Stape, J.L., Campoe, O., Ferraz, S.F.de B.,
2020a. Influence of stand density on growth and water use efficiency in Eucalyptus
clones. For. Ecol. Manage. 466, 118125.
Han, Q., Zhang, Y., Geng, X., Du, K., Yang, J., Kang, X.A., Kang, X., 2020. Response of
tree growth, crown, and branch development to planting density in four Populus
tomentosa clones. Aust. J. For. Sci. 3, 179–198.
Hardanto, A., Röll, A., Hendrayanto, H.D., 2017. Tree soil water uptake and transpiration
in mono-cultural and jungle rubber stands of Sumatra. For. Ecol. Manage. 397,
67–77.
Harms, W.R., Whitesell, C.D., DeBell, D.S., 2000. Growth and development of loblolly
pine in a spacing trial planted in Hawaii. For. Ecol. Manage. 126, 13–24.
Harrington, T.B., Harrington, C.A., DeBell, D.S., 2009. Effects of planting spacing and site
quality on 25-year growth and mortality relationships of Douglas-fir (Pseudotsuga
menziesii var. menziesii). For. Ecol. Manage. 258, 18–25.
Hébert, F., Krause, C., Plourde, P.Y., Achim, A., Prégent, G., Ménétrier, J., 2016. Effect of
tree spacing on tree level volume growth, morphology, and wood properties in a 25-
year-old Pinus banksiana plantation in the boreal forest of Quebec. Forests 7, 276.
Henskens, F.L., Battaglia, M., Cherry, M.L., Beadle, C.L., 2001. Physiological basis of
spacing effects on tree growth and form in Eucalyptus globulus. Trees (Berl.) 15,
365–377.
Horner, G.J., Baker, P.J., Mac Nally, R., Cunningham, S.C., Thomson, J.R., Hamilton, F.,
2009. Mortality of developing floodplain forests subjected to a drying climate and
water extraction. Glob. Change Biol. 15, 2176–2186.
Huber, M., Nieuwendijk, N.M., Pantazopoulou, C.K., Pierik, R., 2021. Light signalling
shapes plant-plant interactions in dense canopies. Plant Cell Environ. 44,
1014–1029.
Huijuan, B., Jialei, Z., Runzhe, Z., Lishui, N., Lianjun, S., 2020. Effects of planting density
and clone type on growth and wood volume of Populus tomentosa stands. J. For. Res.
25 (6), 444–449.
Hummel, S., 2000. Height, diameter and crown dimensions of Cordia alliodora associated
with tree density. For. Ecol. Manage. 127, 31–40.
Ilomäki, S., Nikinmaa, E., Mäkelä, A., 2003. Crown rise due to competition drives
biomass allocation in silver birch. Can. J. For. Res. 33, 2395–2404.
Imada, M., Kunisaki, T., Mizoue, N., Teraoka, Y., 1997. Optimum planting density for
Japanese oak (Quercus mongolica var. grosseserrata) based on spacing experiment
with systematic design. J. For. Res. 2, 89–93.
Ingwers, M.W., Zhao, D., Kane, M.B., Markewitz, D., Teskey, R.O., 2018. Relationships
among growth, δ13C, foliar nitrogen concentration, foliar nitrogen content and
intercepted radiation at different cultural intensities, planting densities and site
indices reveal the importance of water use efficiency in mid-rotation lobloll. For.
Ecol. Manage. 422, 233–240.
Isaac-Renton, M., Stoehr, M., Bealle Statland, C., Woods, J., 2020. Tree breeding and
silviculture: Douglas-fir volume gains with minimal wood quality loss under variable
planting densities. For. Ecol. Manage. 465, 118094.
Johansson, T., 1989. Irradiance within canopies of young trees of european aspen
(Populus tremula) and european birch (Betula pubescens) in stands of different
spacings. For. Ecol. Manage. 28, 217–236.
Johansson, T., 2007. Biomass production and allometric above- and below-ground
relations for young birch stands planted at four spacings on abandoned farmland.
Forestry 80, 41–52.
Kalinganire, A., 1996. Performance of Grevillea robusta in plantations and on farms under
varying environmental conditions in Rwanda. For. Ecol. Manage. 80, 279–285.
Kearney, D., James, R., Montagu, K., Smith, R.G.B., 2007. The effect of initial planting
density on branching characteristics of Eucalyptus pilularis and E. grandis. Aust. For.
70, 262–268.
M.M. Gabira et al.
Kerr, G., 2003. Effects of spacing on the early growth of planted Fraxinus excelsior L. Can.
J. For. Res. 33, 1196–1207.
Khanna, P.K., 1997. Comparison of growth and nutrition of young monocultures and
mixed stands of Eucalyptus globulus and Acacia mearnsii. For. Ecol. Manage. 94,
105–113.
Kholdaenko, Y.A., Belokopytova, L.V., Zhirnova, D.F., Upadhyay, K.K., Tripathi, S.K.,
Koshurnikova, N.N., Sobachkin, R.S., Babushkina, E.A., Vaganov, E.A., 2022. Stand
density effects on tree growth and climatic response in Picea obovata Ledeb.
plantations. For. Ecol. Manage. 519, 120349.
Kigalu, J.M., 2007a. Effects of planting density on the productivity and water use of tea
(Camellia sinensis L.) clones. I. measurement of water use in young tea using sap flow
meters with a stem heat balance method. Agric. Water Manage. 90, 224–232.
Kigalu, J.M., 2007b. Effects of planting density and drought on the productivity of tea
clones (Camellia sinensis L.): Yield responses. Phys. Chem. Earth Parts ABC 32,
1098–1106.
Kirongo, B.B., Mbelase, A.S., Senelwa, K., Hitimana, J., Etiegni, L., 2012. Spacing and
genotype on height and diameter growth of four Eucalyptus under short rotation.
J. Trop. For. Manag. 18, 1–9.
Klein, T., Hartmann, H., 2018. Climate change drives tree mortality. Science 362, 758.
Kumar, R., Lata, K., Khadda, B.S., Jadav, J.K., Rai, A.K., Khajuria, S., 2017. Influence of
planting density on vegetative morphomatrix, yield and economics of drumstick
(Moringa oleifera) cv. PKM 1 under rainfed condition of semi-arid ecosystem. Indian
J. Agric. Sci. 87, 866–869.
Küpers, J.J., Oskam, L., Pierik, R., 2020. Photoreceptors regulate plant developmental
plasticity through auxin. Plants 9, 940.
Lam, T.Y., Guan, B.T., 2020. Modeling stand basal area growth of Cryptomeria japonica D.
Don under different planting densities in Taiwan. J. For. Res. 25, 174–182.
Larsen, S.U., Jørgensen, U., Lærke, P.E., 2019. Harvest interval and row spacing of SRC
willow influence yield and nutrient content. Biomass and Bioenergy 126, 181–189.
Lasserre, J.P., Mason, E.G., Watt, M.S., 2005. The effects of genotype and spacing on
Pinus radiata [D. Don] corewood stiffness in an 11-year old experiment. For. Ecol.
Manage. 205, 375–383.
Lee, Y.-J., Lenhart, J.D., Temple, A., 1997. Influence of planting density on diameter and
height in East Texas pine plantations. South. J. Appl. For. 22, 241–244.
Lima RLS, Campos ARF, Azevedo CAV de, Calado JAW, Silva S de S, Nascimento R do
2016 Effects of planting density on vegetative growth and production components of
jatropha (Physic nut L). Aust. J. Crop Sci. 10:632–636.
Lin, Y., Yang, H., Ivković, M., Gapare, W.J., Colin Matheson, A., Wu, H.X., 2013. Effect of
genotype by spacing interaction on radiata pine genetic parameters for height and
diameter growth. For. Ecol. Manage. 304, 204–211.
Liu, F., Gao, C., Chen, M., Li, K., 2018. Above- and below-ground biomass relationships
of Leucaena leucocephala (Lam.) de Wit in different plant stands. PLoS One 13,
e0207059.
Liu, L., Li, Y., Zhang, J., Sun, H., 2019. Impact of initial planting density on the optimal
economic rotation of Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) in an
experimental forest plantation. Forests 10, 713.
Liziniewicz, M., Ekö, P.M., Agestam, E., 2012. Effect of spacing on 23-year-old lodgepole
pine (Pinus contorta Dougl. var. latifolia) in southern Sweden. Scand. J. For. Res. 27,
361–371.
Loewe-Muñoz, V., Balzarini, M., Del Río, R., Delard, C., 2019. Effects of stone pine (Pinus
pinea L.) plantation spacing on initial growth and conelet entry into production. New
For. 50, 489–503.
Loewe-Muñoz, V., Del Río, R., Balzarini, M., 2020. Stone pine (Pinus pinea) growth and
cone yield as a function of planting density in Chile. Bosque 41, 381–386.
Lokmal, N., Zaki, A.M., Fazwa, M.A.F., Suhaimi, W.C., Azmy, Y., Zakaria, I., Tan, H.,
Khoo, S.K., Wan-Akil, T.A., 2008. Growth of several rubber clones for timber
production. J. Trop. For. Sci. 20, 175–180.
Long, J.N., Smith, F.W., 1988. Leaf area - sapwood area relations of lodgepole pine as
influenced by stand density and site index. Can. J. For. Res. 18, 247–250.
Looney, C.E., Zhang, J., 2022. Site quality and intensive early stand management
practices affect growth dominance, structural complexity, and tree growth in
ponderosa pine plantations. For. Ecol. Manage. 519, 120318.
Luo, J., He, M., Qi, S., Wu, J., Gu, X.S., 2018. Effect of planting density and harvest
protocol on field-scale phytoremediation efficiency by Eucalyptus globulus. Environ.
Sci. Pollut. Res. 25, 11343–11350.
Ma, S., Wang, X., Miao, W., Wang, X., Sun, H., Guo, Z., 2021. Relative influence of
environmental, stand factors and functional traits on allocation of forest productivity
during the restoration of subtropical forests in central China. For. Ecol. Manage. 482,
118814.
Mabapa, M.P., Ayisi, K.K., Mariga, I.K., 2018. Seasonal effect on Moringa oleifera gaseous
exchange and water use efficiency under diverse planting densities. J. Appl. Bot.
Food. Qual. 91, 219–225.
Marziliano, P.A., Coletta, V., Menguzzato, G., Nicolaci, A., Pellicone, G., Veltri, A., 2015.
Effects of planting density on the distribution of biomass in a douglas-fir plantation
in southern Italy. iForest – Biogeosci. For. 8, 368–376.
Marziliano, P.A., Menguzzato, G., Scuderi, A., Scalise, C., Coletta, V., 2017. Biomass
conversion and expansion factors in Douglas-fir stands of different planting density:
variation according to individual growth and prediction equations. For. Syst. 26,
e003.
Matos FA, Daniel O, Serra AP, Heid DM, Nascimento JM do, Nogueira IMB, Ensinas SC,
Altomar PH, Conrad V do A, Potrich DC, Martinez MA 2015 Effects of intra-specific
tree competition on dendrometric parameters of Peltophorum dubium. Aust. J. Crop
Sci. 9:1003–1009.
Mazza, G., Cutini, A., Manetti, M.C., 2015. Influence of tree density on climate-growth
relationships in a Pinus pinaster Ait. forest in the northern mountains of Sardinia
(Italy). IForest 8, 456–463.
14
Forest Ecology and Management 534 (2023) 120882
McCarthy, M.C., Enquist, B.J., 2007. Consistency between an allometric approach and
optimal partitioning theory in global patterns of plant biomass allocation. Funct.
Ecol. 21, 713–720.
McClain, K.M., Morris, D.M., Hills, S.C., Buse, L.J., 1994. The effects of initial spacing on
growth and crown development for planted northern conifers: 37-year results. For.
Chron. 70, 174–182.
McElwain, J.C., Yiotis, C., Lawson, T., 2016. Using modern plant trait relationships
between observed and theoretical maximum stomatal conductance and vein density
to examine patterns of plant macroevolution. New Phytol. 209, 94–103.
Medeiros, P.L., Silva, G.G.C., Oliveira, E.M.M., Ribeiro, C.O., Silva, J.M.S., Pimenta, A.S.,
2020. Efficiency of nutrient use for biomass production of a Eucalyptus clone as a
function of planting density in short-rotation cropping. Aust. For. 83, 66–74.
Mehari, A., Habte, B., 2006. Influence of initial spacing on growth and branching
characteristics of Cordia africana trees established on Eritrean highland. New For. 31,
185–193.
Meinzer, F.C., Fownes, J.H., Harrington, R.A., 1996. Growth indices and stomatal control
of transpiration in Acacia koa stands planted at different densities. Tree Phys. 16,
607–615.
Mendieta-Araica, B., Spörndly, E., Reyes-Sánchez, N., Salmerón-Miranda, F., Halling, M.,
2013. Biomass production and chemical composition of Moringa oleifera under
different planting densities and levels of nitrogen fertilization. Agrofor. Syst. 87,
81–92.
Metsaranta, J.M., 2019. Long-term tree-ring derived carbon dynamics of an experimental
plantation in relation to species and density in Northwestern Ontario. Canada. For.
Ecol. Manage. 441, 229–241.
Milyutina, I.L., Sudachkova, N.E., Romanova, L.I., 2013. Response of the antioxidant
system of light-demanding and shade-bearing pine species to phytocenotic stress.
Contemp. Probl. Ecol. 6, 149–155.
Miranda, I., Gominho, J., Pereira, H., 2015. Heartwood, sapwood and bark variation in
coppiced Eucalyptus globulus trees in 2nd rotation and comparison with the single-
stem 1st rotation. Silva Fenn. 49, 1141.
Moher D, Liberati A, Tetzlaff J, Altman DG, Group P 2009 Preferred reporting items for
systematic reviews and meta-analyses: The PRISMA statement. PLoS Med. 6:
e1000097.
Moore, G.W., Bond, B.J., Jones, J.A., 2011. A comparison of annual transpiration and
productivity in monoculture and mixed-species Douglas-fir and red alder stands. For.
Ecol. Manage. 262, 2263–2270.
Nagar, B., Rawat, S., Rathiesh, P., Sekar, I., 2015. Impact of initial spacing on growth and
yield of Eucalyptus camaldulensis in arid region of India. World Appl. Sci. J. 33,
1362–1368.
Naji, H.R., Bakar, E.S., Soltani, M., Ebadi, S.E., Abdul-Hamid, H., Javad, S.K.S., Sahri, M.
H., 2014. Effect of initial planting density and tree features on growth, wood density,
and anatomical properties from a Hevea brasiliensis trial plantation. For. Prod. J. 64,
41–47.
Naji, H.R., Nia, M.F., Kiaei, M., Abdul-Hamid, H., Soltani, M., Faghihi, A., 2016. Effect of
intensive planting density on tree growth, wood density and fiber properties of
maple (Acer velutinum Boiss.). iForest –. Biogeosci. For. 9, 325–329.
Nan, W., Ta, F., Meng, X., Dong, Z., Xiao, N., 2020. Effects of age and density of Pinus
sylvestris var. mongolica on soil moisture in the semiarid Mu Us Dunefield, northern
China. For. Ecol. Manage. 473, 118313.
Narducci, T.S., Yared, J.A.G., Júnior, S.B., 2016. Growth and survival of Sclerolobium
paniculatum Vogel and the relationship between rainfall and the increment in
diameter at different planting spacings. Rev. Árvore 40, 447–454.
Negrave, R.W., Prescott, C.E., Barker, J.E., 2007. Growth and foliar nutrition of juvenile
western hemlock and western redcedar plantations on low- and medium-
productivity sites on northern Vancouver Island: response to fertilization and
planting density. Can. J. For. Res. 37, 2587–2599.
Nelson, A.S., Wagner, R.G., Day, M.E., Fernandez, I.J., Weiskittel, A.R., Saunders, M.R.,
2016. Light absorption and light-use efficiency of juvenile white spruce trees in
natural stands and plantations. For. Ecol. Manage. 376, 158–165.
Newton, P.F., 2015. Occurrence of density-dependent height repression within jack pine
and black spruce populations. Forests 6, 2450–2468.
Nilsson, U., 1994. Development of growth and stand structure in Picea abies stands
planted at different initial densities. Scandinavian J. For. Res. 9, 135–142.
Nilsson, U., Albrektson, A., 1994. Growth and self-thinning in two young Scots pine
stands planted at different initial densities. For. Ecol. Manage. 68, 209–215.
Ning K, Ding C, Huang Q, Zhang W, Yang C, Liang D, Fan R, Su X 2019 Transcriptome
profiling revealed diverse gene expression patterns in poplar (Populus ×
euramericana) under different planting densities. PLoS ONE, 14.
Nirsatmanto, A., 2016. Early growth of improved Acacia mangium at different planting
densities. Jurnal Manajemen Hutan Tropika 22, 105–113.
O’Hara, K.L., York, R.A., 2014. Leaf area development and crown architecture in a giant
sequoia spacing study. For. Sci. 60, 776–783.
Okojie, J.A., Bailey, R.L., Borders, B.E., 1988. Spacing effects in an unthinned 11-year-
old Terminalia superba plantation in the dry lowland rainforest area of Nigeria. For.
Ecol. Manage. 23, 253–260.
Ola-Adams, B.A., 1993. Effects of spacing on biomass distribution and nutrient content of
Tectona grandis Linn. f. (teak) and Terminalia superba Engl. Diels. (afara) in south-
western Nigeria. For. Ecol. Manage. 58, 299–319.
Oliveira, N., del Río, M., Forrester, D.I., Rodríguez-Soalleiro, R., Pérez-Cruzado, C.,
Cañellas, I., Sixto, H., 2018. Mixed short rotation plantations of Populus alba and
Robinia pseudoacacia for biomass yield. For. Ecol. Manage. 410, 48–55.
Olmos, V.M., 2022. Forestry and the forest products sector: Production, income and
employment, and international trade. Forest Policy Econ. 135, 102648.
Özbayram, A.K., Çiçek, E., 2020. Initial planting density experiments of narrow-leaved
ash in Turkey: ten-year results. Sumar. List 144, 269–278.
M.M. Gabira et al.
Pachas, A.N.A., Shelton, H.M., Lambrides, C.J., Dalzell, S.A., Murtagh, G.J., Hardner, C.
M., 2018. Effect of tree density on competition between Leucaena leucocephala and
Chloris gayana using a Nelder Wheel trial II. Belowground interactions. Crop Pasture
Sci. 69, 733–744.
Pachas, A.N.A., Sakanphet, S., Soukkhy, O., Lao, M., Savathvong, S., Newby, J.C.,
Souliyasack, B., Keoboualapha, B., Dieters, M.J., 2019. Initial spacing of teak
(Tectona grandis) in northern Lao PDR: impacts on the growth of teak and companion
crops. For. Ecol. Manage. 435, 77–88.
Pallett, R.N., Sale, G., 2004. The relative contributions of tree improvement and cultural
practice toward productivity gains in Eucalyptus pulpwood stands. For. Ecol.
Manage. 193, 33–43.
Pannacci, E., Bartolini, S., Covarelli, G., 2009. Evaluation of four poplar clones in a short
rotation forestry in central Italy. Ital. J. Agron. 4, 191–198.
Parker, G.G., 2020. Tamm review: Leaf Area Index (LAI) is both a determinant and a
consequence of important processes in vegetation canopies. For. Ecol. Manage. 477,
118496.
Paula, R.R., Reis, G.G., Reis, M.G.F., Oliveira Neto, S.N., Leite, H.G., Melido, R.C.N.,
Lopes, H.N.S., Souza, F.C., 2013. Eucalypt growth in monoculture and silvopastoral
systems with varied tree initial densities and spatial arrangements. Agrofor. Syst. 87,
1295–1307.
Penner, M., Robinson, C., Burgess, D., 2001. Pinus resinosa product potential following
initial spacing and subsequent thinning. For. Chron. 77, 129–139.
Peracca, G.G., O’Hara, K.L., 2008. Effects of growing space on growth for 20-year-old
giant sequoia, ponderosa pine, and Douglas-fir in the Sierra Nevada. West. J. Appl.
For. 23, 156–165.
Pérez, I., Pérez, J., Carrasco, J., Ciria, P., 2014. Siberian elm responses to different
culture conditions under short rotation forestry in Mediterranean areas. Turkish J.
Agric. For. 38, 652–662.
Pinkard, E.A., Neilsen, W.A., 2003. Crown and stand characteristics of Eucalyptus nitens
in response to initial spacing: implications for thinning. For. Ecol. Manage. 172,
215–227.
Pons, T.L., Jordi, W., Kuiper, D., 2001. Acclimation of plants to light gradients in leaf
canopies: evidence for a possible role for cytokinins transported in the transpiration
stream. J. Exp. Bot. 52, 1563–1574.
Prasad, J.V.N.S., Korwar, G.R., Rao, K.V., Mandal, U.K., Rao, G.R., Srinivas, I.,
Venkateswarlu, B., Rao, S.N., Kulkarni, H.D., 2011. Optimum stand density of
Leucaena leucocephala for wood production in Andhra Pradesh, Southern India.
Biomass Bioenergy 35, 227–235.
Quer, E., Baldy, V., DesRochers, A., 2020. Ecological drivers of root grafting in balsam fir
natural stands. For. Ecol. Manage. 475, 118388.
Quicke, H., Glover, G., Glover, R., 1999. Loblolly pine growth response to herbaceous
vegetation control at different planting densities. Can. J. For. Res. 29, 960–967.
Radtke, P.J., Burkhart, H.E., 1999. Basal area growth and crown closure in a loblolly pine
spacing trial. For. Sci. 45, 35–44.
Rai, R.S., Srinivasan, V.M., 1990. High density short rotation studies in Eucalyptus
tereticornis and Casuarina equisetifolia. Int. Crops J. 6, 113–122.
Ramalho, F.M.G., Pimenta, E.M., Goulart, C.P., de Almeida, M.N.F., Vidaurre, G.B.,
Hein, P.R.G., 2019. Effect of stand density on longitudinal variation of wood and
bark growth in fast-growing Eucalyptus plantations. iForest – Biogeosci. For. 12,
527–532.
Ramos-Trejo OS, Canul-Solís JR, Alvarado-Canché A del R, Castillo-Sánchez LE,
Sandoval-Gío JJ, Campos-Navarrete MJ, Piñero-Vázquez AT, Chay-Canul AJ,
Casanova-Lugo F (2020) Growth, forage yield and quality of Morus alba L. and
Gliricidia sepium (Jacq.) Walp. in mixed and pure fodder bank systems in Yucatan,
México. Agrofor. Syst. 94:151–157.
Rédei, K., 2009. Effects of initial spacing on the stand structure and yield of young black
locust (Robinia pseudoacacia L.) stands. Int. J. Hortic. Sci. 15.
Remaury, A., Guittonny, M., Rickson, J., 2019. The effect of tree planting density on the
relative development of weeds and hybrid poplars on revegetated mine slopes
vulnerable to erosion. New For. 50, 555–572.
Resende, R.T., Soares, A.A.V., Forrester, D.I., Marcatti, G.E., dos Santos, A.R.,
Takahashi, E.K., Silva, F.F.e., Grattapaglia, D., Resende, M.D.V., Leite, H.G., 2018.
Environmental uniformity, site quality and tree competition interact to determine
stand productivity of clonal Eucalyptus. For. Ecol. Manage. 410, 76–83.
Resquin F, Navarro-Cerrillo RM, Rachid-Casnati C, Hirigoyen A, Carrasco-Letelier L,
Duque-Lazo J 2018 Allometry, growth and survival of three eucalyptus species
(Eucalyptus benthamii Maiden and Cambage, E. dunnii Maiden and E. grandis Hill ex
Maiden) in high-density plantations in Uruguay. Forests 9:745.
Resquin, F., Navarro-Cerrillo, R.M., Carrasco-Letelier, L., Casnati, C.R., 2019. Influence
of contrasting stocking densities on the dynamics of above-ground biomass and
wood density of Eucalyptus benthamii, Eucalyptus dunnii, and Eucalyptus grandis for
bioenergy in Uruguay. For. Ecol. Manage. 438, 63–74.
Resquin, F., Navarro-Cerrillo, R.M., Carrasco-Letelier, L., Casnati, C.R., Bentancor, L.,
2020. Evaluation of the nutrient content in biomass of Eucalyptus species from short
rotation plantations in Uruguay. Biomass and Bioenergy 134, 105502.
Rodríguez-García, E., Bravo, F., 2013. Plasticity in Pinus pinaster populations of diverse
origins: Comparative seedling responses to light and nitrogen availability. For. Ecol.
Manage. 307, 196–205.
Rončević, S., Andrašev, S., Ivanišević, P., Kovačević, B., Klašnja, B., 2013. Biomass
production and energy potential of some eastern cottonwood (Populus deltoides Bartr.
ex Marsh.) clones in relation to planting spacing. Sumar. List 137, 33–42.
Roth, B.E., Jokela, E.J., Martin, T.A., Huber, D.A., White, T.L., 2007a. Genotype ×
environment interactions in selected loblolly and slash pine plantations in the
Southeastern United States. For. Ecol. Manage. 238, 175–188.
15
Forest Ecology and Management 534 (2023) 120882
Roth, B.E., Li, X., Huber, D.A., Peter, G.F., 2007b. Effects of management intensity,
genetics and planting density on wood stiffness in a plantation of juvenile loblolly
pine in the southeastern USA. For. Ecol. Manage. 246, 155–162.
Russell, M.B., Burkhart, H.E., Amateis, R.L., 2009. Biomass partitioning in a miniature-
scale loblolly pine spacing trial. Can. J. For. Res. 39, 320–329.
Schwerz, F., Eloy, E., Elli, E.F., Caron, B.O., 2019. Reduced planting spacing increase
radiation use efficiency and biomass for energy in black wattle plantations: towards
sustainable production systems. Biomass Bioenergy 120, 229–239.
Schwerz, F., Neto, D.D., Caron, B.O., Nardini, C., Sgarbossa, J., Eloy, E., Behling, A.,
Elli, E.F., Reichardt, K., 2020a. Biomass and potential energy yield of perennial
woody energy crops under reduced planting spacing. Renew. Energy 153,
1238–1250.
Schwerz, F., Neto, D.D., Caron, B.O., Tibolla, L.B., Sgarbossa, J., Eloy, E., Elli, E.F.,
Carvalho, L.G., 2020b. Carbon stocks, partitioning, and wood composition in short-
rotation forestry system under reduced planting spacing. Ann. For. Sci. 77, 67.
Scolforo, J.R.S., Altoé, T.F., Scolforo, H.F., de Mello, J.M., Silva, C.P.C.e., Ferraz Filho, A.
C., 2016. Management strategies of Eremanthus erythropappus (DC.) MacLeish under
different initial spacing. Ciência e Agrotecnologia 40, 298–304.
Scott, W., Meade, R., Leon, R., Hyink, D., Miller, R., 1998. Planting density and tree-size
relations in coast Douglas-fir. Can. J. For. Res. 28 (1), 74–78.
Scott, R.E., Mitchell, S.J., 2005. Empirical modelling of windthrow risk in partially
harvested stands using tree, neighbourhood, and stand attributes. For. Ecol. Manage.
218, 193–209.
Scott, D.A., Tiarks, A., 2008. Dual-cropping loblolly pine for biomass energy and
conventional wood products. South. J. Appl. For. 32, 33–37.
Shainsky, L.J., Newton, M., Radosevich, S.R., 1992. Effects of intra- and inter-specific
competition on root and shoot biomass of young Douglas-fir and red alder. Can. J.
For. Res. 22, 101–110.
Sharma-Natu, P., Ghildiyal, M.C., 2005. Potential targets for improving photosynthesis
and crop yield. Curr. Sci. 88, 1918–1928.
Shi, K., Cao, Q., 1997. Predicted leaf area growth and foliage efficiency of loblolly pine
plantations. For. Ecol. Manage. 95, 109–115.
Silva, W.G., dos Santos, D.M.C.L., Lima, A.P.L., Mattos, F.D., Lima, S.F., de Paula, R.C.M.,
2019. Growth and volumetric production of eucalyptus clones in different planting
spaces. Rev. Agric. Neotrop. 6, 38–47.
Silva PSL, Paiva HN de, Oliveira VR de, Siqueira PL de OF, Soares EB, Monteiro AL,
Tavella LB 2014 Biomass of three species as a response to planting density and
interspecific competition. Rev. Árvore 38:319–329.
Silva, P.S.L., Holanda, A.E.R., de Paiva, H.N., de Oliveira, F.H.T., de Oliveira, O.F., 2012.
Planting density and initial growth of two tree species adapted to the semi-arid
region. Rev. Árvore 36, 951–959.
Smith, R.G.B., Glencross, K., Nichols, J.D., Palmer, G., Viranamangga, R., 2016. Effect of
initial spacing on first thinning product recovery, and financial outcomes in
whitewood (Endospermum medullosum) plantations in Vanuatu. Small-scale For. 15,
45–59.
Snowdon, P., Woollons, R.C., Benson, M.L., 1998. Incorporation of climatic indices into
models of growth of Pinus radiata in a spacing experiment. New For. 16, 101–123.
Srivastava, N., Goel, V.L., Behl, H.M., 1999. Influence of planting density on growth and
biomass productivity of Terminalia arjuna under sodic soil sites. Biomass and
Bioenergy 17, 273–278.
Stape, J.L., Binkley, D., 2010. Insights from full-rotation Nelder spacing trials with
Eucalyptus. South. For. 72, 90–97.
Stape, J.L., Binkley, D., Ryan, M.G., 2004. Eucalyptus production and the supply, use and
efficiency of use of water, light and nitrogen across a geographic gradient in Brazil.
For. Ecol. Manage. 193, 17–31.
Štefančík, I., 2020. Development of the Norway spruce (Picea abies /L./ Karst.) stand
established by various spacings and affected by abiotic harmful factors and ungulate
game. J. For. Sci. 66, 117–131.
Stolarski, M.J., Szczukowski, S., Tworkowski, J., Krzyżaniak, M., Załuski, D., 2017.
Willow biomass and cuttings’ production potential over ten successive annual
harvests. Biomass and Bioenergy 105, 230–247.
Stolarski, M.J., Szczukowski, S., Tworkowski, J., Krzyżaniak, M., 2019a. Extensive
willow biomass production on marginal land. Polish J. Environ. Stud. 28,
4359–4367.
Stolarski, M.J., Szczukowski, S., Tworkowski, J., Krzyżaniak, M., Załuski, D., 2019b.
Willow production during 12 consecutive years—The effects of harvest rotation,
planting density and cultivar on biomass yield. GCB Bioenergy 11, 635–656.
Subedi, S., Kane, M., Zhao, D., Borders, B., Greene, D., 2012. Cultural intensity and
planting density effects on aboveground biomass of 12-year-old loblolly pine trees in
the Upper Coastal Plain and Piedmont of the southeastern United States. For. Ecol.
Manage. 267, 157–162.
Sun, H., Diao, S., Liu, R., Forrester, D., Soares, A., Saito, D., Dong, R., Jiang, J., 2018.
Relationship between size inequality and stand productivity is modified by self-
thinning, age, site and planting density in Sassafras tzumu plantations in central
China. For. Ecol. Manage. 422, 199–206.
Tanabe, J., Endo, R., Kuroda, S., Ishiguri, F., Narisawa, T., Takashima, Y., 2019. Variance
components and parent–offspring correlations of growth traits vary among the initial
planting spacings in Zelkova serrata. Silvae Genet. 68, 45–50.
Tarroux, E., Desrochers, A., 2011. Effect of natural root grafting on growth response of
jack pine (Pinus banksiana; Pinaceae). Am. J. Bot. 98, 967–974.
Tarroux, E., Desrochers, A., Krause, C., 2010. Effect of natural root grafting on growth
response of jack pine (Pinus banksiana) after commercial thinning. For. Ecol.
Manage. 260, 526–535.
Tavares, A.P.C., Soares, T.S., Cruz, E.S., 2020. Dendrometric characteristics and basic
wood density of eucalyptus clones in different spacing. Rev. Agric. Neotrop. 7,
104–108.
M.M. Gabira et al.
Tenorio, C., Roque, R.M., Valverde, J.C., Arias-Aguilar, D., 2019. Biomass production
and characteristics of short rotation plantations of clones of Gmelina arborea in three
spacings. Silvae Genet. 68 (1), 92–100.
Thoranisorn, S., Sahunalu, P., Yoda, K., 1990. Density effects and sel-thinning in even-
aged pure stands of Eucalyptus camaldulensis Dehn. The Botanical Magazine 103,
283–295.
Thyroff, E.C., Burney, O.T., Mickelbart, M.V., Jacobs, D.F., 2019. Unraveling shade
tolerance and plasticity of semi-evergreen oaks: Insights from maritime forest live
oak restoration. Front. Plant Sci. 10, 1526.
Toillon, J., Fichot, R., Dallé, E., Berthelot, A., Brignolas, F., Marron, N., 2013. Planting
density affects growth and water-use efficiency depending on site in Populus deltoides
× P. nigra. For. Ecol. Manage. 304, 345–354.
Tong, Q.J., Zhang, S.Y., 2005. Impact of initial spacing and precommercial thinning on
jack pine tree growth and stem quality. For. Chron. 81, 418–428.
Tonini, H., Schwengber, D.R., Morales, M.M., Magalhães, C.A.S., de Oliveira, J.M.F.,
2018. Growth, biomass, and energy quality of Acacia mangium timber grown at
different spacings. Pesqui. Agropecu. Bras. 53, 791–799.
Torreano, S.J., Smalley, G.W., Nunley, A.N., Rennie, J.C., 2012. Structure and growth of
40-year-old unthinned loblolly pine planted at three spacings in south-central
Tennessee. South. J. Appl. For. 36, 136–140.
Truax, B., Fortier, J., Gagnon, D., Lambert, F., 2018. Planting density and site effects on
stem dimensions, stand productivity, biomass partitioning, carbon stocks and soil
nutrient supply in hybrid poplar plantations. Forests 9, 293.
Tsamir, M., Gottlieb, S., Preisler, Y., Rotenberg, E., Tatarinov, F., Yakir, D., Tague, C.,
Klein, T., 2019. Stand density effects on carbon and water fluxes in a semi-arid
forest, from leaf to stand-scale. For. Ecol. Manage. 453, 117573.
Tun, T.N., Guo, J., Fang, S., Tian, Y., 2018. Planting spacing affects canopy structure,
biomass production and stem roundness in poplar plantations. Scand. J. For. Res. 33,
464–474.
Uhl, E., Biber, P., Ulbricht, M., Heym, M., Horváth, T., Lakatos, F., Gál, J., Steinacker, L.,
Tonon, G., Ventura, M., Pretzsch, H., 2015. Analysing the effect of stand density and
site conditions on structure and growth of oak species using Nelder trials along an
environmental gradient: experimental design, evaluation methods, and results. For.
Ecosyst. 2, 1–19.
Ungar, E.D., Rotenberg, E., Raz-Yaseef, N., Cohen, S., Yakir, D., Schiller, G., 2013.
Transpiration and annual water balance of Aleppo pine in a semiarid region:
implications for forest management. For. Ecol. Manage. 298, 39–51.
Waghorn, M.J., Mason, E.G., Watt, M.S., 2007. Influence of initial stand density and
genotype on longitudinal variation in modulus of elasticity for 17-year-old Pinus
radiata. For. Ecol. Manage. 252, 67–72.
Waghorn, M.J., Watt, M.S., 2013. Stand variation in Pinus radiata and its relationship
with allometric scaling and critical buckling height. Ann. Bot. 111, 675–680.
Wang, C.-S., Zeng, J., Hein, S., Zhao, Z.-G., Guo, J.-J., Zeng, J., 2016. Crown and branch
attributes of mid-aged Betula alnoides plantations in response to planting density.
Scand. J. For. Res. 32 (8), 679–687.
Wang, C.-S., Tang, C., Hein, S., Guo, J.-J., Zhao, Z.-G., Zeng, J., 2018. Branch
development of five-year-old Betula alnoides plantations in response to planting
density. Forests 9, 42.
Wang, C.-S., Guo, J.-J., Hein, S., Wang, H., Zhao, Z.-G., Zeng, J., 2019. Foliar morphology
and spatial distribution in five-year-old plantations of Betula alnoides. For. Ecol.
Manage. 432, 514–521.
Wang, Z., Yu, N., Li, R., Zou, W., Yang, J., Yin, G., Peng, T., Wang, Z., Yu, N., Li, R.,
Zou, W., Yang, J., Yin, G., Peng, T., 2020. Branch development of eight-year-old
Mytilaria laosensis plantations in response to planting density. Austrian J. For. Sci. 4,
247–266.
Węgiel, A., Polowy, K., 2020. Aboveground carbon content and storage in mature Scots
pine stands of different densities. Forests 11 (2), 240.
Weiner, J., Freckleton, R.P., 2010. Constant final yield. Annu. Rev. Ecol. Evol. Syst. 41,
173–192.
Wells, E.D., 1994. Effects of planting spacing and refertilization on growth and nutrition
of black spruce (Picea mariana (Mill.) B.S.P.) planted on a minerotrophic peatland in
Newfoundland. Canada. Can. J. For. Res. 24, 1302–1311.
West, P.W., Smith, R.G.B., 2019. Inter-tree competitive processes during early growth of
an experimental plantation of Eucalyptus pilularis in sub-tropical Australia. For. Ecol.
Manage. 451, 117450.
West, P.W., Smith, R.G.B., 2020. Effects of tree spacing on branch-size development
during early growth of an experimental plantation of Eucalyptus pilularis in
subtropical Australia. Aust. For. 83, 39–45.
16
Forest Ecology and Management 534 (2023) 120882
Will, R.E., Barron, G.A., Burkes, E.C., Shiver, B., Teskey, R.O., 2001. Relationship
between intercepted radiation, net photosynthesis, respiration, and rate of stem
volume growth of Pinus taeda and Pinus elliottii stands of different densities. For. Ecol.
Manage. 154, 155–163.
Will, R.E., Narahari, N.V., Shiver, B.D., Teskey, R.O., 2005. Effects of planting density on
canopy dynamics and stem growth for intensively managed loblolly pine stands. For.
Ecol. Manage. 205, 29–41.
Will, R.E., Hennessey, T., Lynch, T., Holeman, R., Heinemann, R., 2010. Effects of
planting density and seed source on loblolly pine stands in Southeastern Oklahoma.
For. Sci. 56, 437–443.
Wilson, J.B., Steel, J.B., Steel, S.L.K., 2007. Do plants ever compete for space? Folia
Geobot. 42, 431–436.
Woodruff, D.R., Bond, B.J., Ritchie, G.A., Scott, W., 2002. Effects of stand density on the
growth of young Douglas-fir trees. Can. J. For. Res. 32, 420–427.
Xue, L., Jacobs, D.F., Zeng, S., Yang, Z., Guo, S., Liu, B., 2012. Relationship between
above-ground biomass allocation and stand density index in Populus xeuramericana
stands. Forestry 85, 611–619.
Yan, Z., Zhou, Q., Teng, M., Ji, H., Zhang, J., He, W., Ye, Y., Wang, B., Wang, P., 2019.
High planting density and leaf area index of masson pine forest reduce crown
transmittance of photosynthetically active radiation. Glob. Ecol. Conserv. 20,
e00759.
Yáñez, M.A., Fox, T.R., Seiler, J.R., 2015. Early growth responses of loblolly pine
varieties and families to silvicultural intensity. For. Ecol. Manage. 356, 204–215.
Yáñez, M.A., Zamudio, F., Espinoza, S., Ivković, M., Guerra, F., Espinosa, C., Baettig, R.
M., 2019. Genetic variation and growth stability of hybrid poplars in high-density
short-rotation coppice stands in central Chile. Biomass Bioenergy 120, 84–90.
York RA, O’hara KL, Battles JJ 2013 Density effects on giant sequoia (Sequoiadendron
giganteum) growth through 22 years:Implications for restoration and plantation
management. West. J. Appl. For. 28:30–36.
Załuski, D., Mielniczuk, J., Bronowicka-Mielniczuk, U., Stolarski, M.J., Krzyżaniak, M.,
Szczukowski, S., Tworkowski, J., 2018. Survival analysis of plants grown in long-
term field experiments. Agron. J. 110, 1791–1798.
Zhang, X., Cao, Q.V., Duan, A., Zhang, J., 2017. Modeling tree mortality in relation to
climate, initial planting density and competition in Chinese fir plantations using a
Bayesian logistic multilevel method. Can. J. For. Res. 47, 1278–1285.
Zhang, J., Finley, K.A., Johnson, N.G., Ritchie, M.W., 2019. Lowering stand density
enhances resiliency of ponderosa pine forests to disturbances and climate change.
For. Sci. 65, 496–507.
Zhang, Y., Tian, Y., Ding, S., Lv, Y., Samjhana, W., Fang, S., 2020c. Growth, carbon
storage, and optimal rotation in poplar plantations: a case study on clone and
planting spacing effects. Forests 11, 842.
Zhang, X., Wang, Z., Chhin, S., Wang, H., Duan, A., Zhang, J., 2020b. Relative
contributions of competition, stand structure, age, and climate factors to tree
mortality of Chinese fir plantations:Long-term spacing trials in southern China. For.
Ecol. Manage. 465, 118103.
Zhang, J., Zhang, J., Mattson, K., Finley, K., 2020a. Effect of silviculture on carbon pools
during development of a ponderosa pine plantation. Forests 11, 997.
Zhao, D., Bullock, B.P., Montes, C.R., Wang, M., Greene, D., Sutter, L., 2019. Loblolly
pine outperforms slash pine in the southeastern United States – A long-term
experimental comparison study. For. Ecol. Manage. 450, 117532.
Zhao, D., Bullock, B.P., Montes, C.R., Wang, M., Westfall, J., Coulston, J.W., 2020. Long-
term dynamics of loblolly pine crown structure and aboveground net primary
production as affected by site quality, planting density and cultural intensity. For.
Ecol. Manage. 472, 118259.
Zhao, D., Kane, M., 2012. Differences in growth dynamics of loblolly and slash pine
plantations in the southeastern United States. For. Ecol. Manage. 281, 84–92.
Zhao, D., Kane, M., Borders, B.E., 2011. Growth responses to planting density and
management intensity in loblolly pine plantations in the southeastern USA Lower
Coastal Plain. Ann. For. Sci. 68, 625–635.
Zhao, D., Kane, M., Borders, B., Subedi, S., Akers, M., 2012. Effects of cultural intensity
and planting density on stand-level aboveground biomass production and allocation
for 12-year-old loblolly pine plantations in the upper coastal plain and piedmont of
the southeastern United States. Canadian Journal of Forest Research 42 (1),
111–122.
Zheng, Y., Zhang, Y., Wu, J., 2016. Yield and quality of Moringa oleifera under different
planting densities and cutting heights in southwest China. Ind. Crops Prod. 91,
88–96.
Zutter, B.R., Glover, G.R., Mitchell, R.J., 1998. Influence of plant density and soil organic
matter on the first-year growth of loblolly pine and sweetgum. For. Sci. 44, 397–404.