Professional Documents
Culture Documents
Spacing Review
Spacing Review
Spacing Review
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 The ecological, economical, and social importance of planted forests has
been discussed extensively, and plantations are considered important for
Forest-based industry is a crucial sector of socioeconomic develop community development, particularly in the context of climate change
ment around the world and provides several raw products to fiber, and post COVID-19 pandemic (Celentano et al., 2022; Olmos, 2022). In
timber, energy, and non-timber forest-product industries. Forest plan general, plantation forests are established with a design aimed at
tations for industrial purposes cover about 131 million ha, correspond providing the optimal conditions for tree growth and maximum pro
ing to 2 % of the world’s total forest cover (FAO, 2020). Asia contains ductivity and for managing timber quality with a focus on the desired
the largest area of plantation forests (79,332,000 ha of planted forests), final products.
and the highest ratio of introduced species in planted forests is in South Silviculture in forests and plantations has a main goal of managing
America, where 97 % of the planted areas contain non-native species. 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. Forest Ecology and Management 534 (2023) 120882
Competition is defined as the demand by individuals for a common regular increase in biomass accumulation up to a certain point where
required resource that may exceed the immediate supply; thus, biomass accumulation decreases because natural resources become
competition reduces tree productivity (Begon and Townsend, 2020). limiting for growth.
Light is often considered a source of competition between trees because Given the importance of forest plantations worldwide and the po
it is the only environmental factor that cannot be easily managed in tential of planting density to be used as management tool, several studies
even-aged and un-thinned forest monocultures (Chase et al., 2016; have been developed to clarify species responses to planting density at
Nelson et al., 2016); this competition leads to physiological and individual and stand levels in forest plantations. However, to our
morphological changes—also called shade-avoidance responses—which knowledge, the results of these studies have never been combined and
include hyponasty, stem and petiole elongation, apical dominance, leaf analyzed through a systematic review, which offers many advantages
positioning, and life-cycle adjustments (Bongers et al., 2018). Huber over other research synthesis techniques, as its results can be evaluated,
et al. (2021) explored the physiological, biochemical, and genetic as investigated, and updated more accurately. In this review, we discuss
pects of shade-avoidance responses caused by light signaling; they how planting density affects plantation forest growth, yield, and phys
demonstrated that competition for light may impact the canopy struc iological responses in temperate and tropical regions. We focus on
ture, species composition, and size of individuals in plant communities. monospecific planted forests because they are some of the most inten
Competition affects plant physiology directly, mediating individual sively managed stands, contain generally only one or two species, and
growth by regulating resource acquisition (Benomar et al., 2012). In are characterized by even-aged regularly spaced trees. We provide an
drier regions, light may not be as limiting as water for productivity, overview of how planting density is managed as a silvicultural practice
which limits plant growth and affects resource- use efficiency and and summarize recent advances, trends, and gaps to guide future
biomass allocation (Gonçalves et al., 2004; Stape et al., 2004). Space is research in this field.
also a limiting factor for tree growth because it influences resource
consumption per area; however, space is not considered a resource for 2. Material and methods
which plants compete because it is probably never limiting in terrestrial
ecosystems, as opposed to water and other essential resources (Wilson 2.1. Criteria for selecting planting density − growth studies
et al., 2007). Planted forests are sites of relatively homogeneous
competition because trees are usually even-aged, regularly spaced in We searched the existing literature for studies published between
dividuals of the same species and deal with a relatively uniform resource 1981 and 2020 focused on the morphological and physiological re
availability at the stand scale; however, changes in environmental re sponses of plants to planting density in forest stands. We selected this 40-
sources availability at a stand scale are known to influence productivity, year period to highlight the most up-to-date advances in the field of
as observed by Resende et al. (2018). Thus, understanding species re planting density available in online databases. The literature review was
sponses to spatiotemporal variations in resources is essential to provide conducted according to the PRISMA methodology (Moher et al., 2009;
theoretical guidance to stand management in different regions. Fig. 1).
Despite this focus on competition, silviculturists must consider that We used two indexing electronic databases—Web of Science and
trees are not always competing with each other; facilitation relation Scopus—to identify the available literature and encompass a wide va
ships also occur in forest stands, and silvicultural practices aiming to riety of high-quality journals. Title, abstract, and keywords were
reduce tree density do not always positively affect growth (Quer et al., searched for the words “forest,” “planting,” “plant,” “tree,” “plant,”
2020). In general, facilitation relationships create a buffer that mitigates “plantation,” “stem,” “stand,” “spacing,” “stock,” “stocking,” and “den
variations in soil humidity, atmospheric temperature, and light inter sity,” using Boolean operators, e.g., OR, AND. Our search was limited to
ception and favors plant establishment and growth (Bravo-Oviedo et al., research papers published between 1981 and 2020 in English-language
2018). An example of facilitation is to mix species in forest stands as a scientific journals, and we excluded review articles, event papers,
way to increase its growth, as explored by Forrester (2015); although the technical reports, and short communications. Our database search
author observed that not all trees increased their growth in comparison returned 11,602 papers in Web of Science and 8097 papers in Scopus. A
to monospecific stands. Natural root grafting has also been reported as a total of 10,820 papers remained after removing duplicates.
means of creating facilitation relationships among individual jack pine In the screening process, we analyzed and excluded papers that did
(Pinus banksiana Lamb.) to improve resource use by the trees (Tarroux not meet specific criteria. We excluded papers that (a) focused on
and Desrochers, 2011). When these stands are thinned, however, the agricultural crop species, such as cotton, wheat, maize, and soy; (b) were
root system of the removed trees increases the carbon demand on the based on field trials established in drylands or wetlands; (c) focused on
residual trees, decreasing the latter’s growth relative to non-root-grafted the development of GIS and remote-sensing technologies; and (d) were
trees, despite benefiting from more resources (Tarroux et al., 2010). This based on modelling or mathematical models, not on field trials. We
example illustrates how tree growth can depend not only on competition included papers focused on either temperate or tropical species (or
but also facilitation (Begon and Townsend, 2020); therefore, facilitation both), given that the most relevant planting density studies occur in both
must be better understood in forest stands to promote tree growth and zones. After the application of these criteria, we retained 188 papers.
increase the efficacy of resource use. The remaining papers were then evaluated in terms of the clarity of the
In the design and planning of plantations, tree spacing is a major supplied information and the application of stand density within the
factor because of its impact on profitability—close spacing requires field trials. We included in our study only papers that (a) were based on
more seedlings—and productivity because closer spacings can improve data collected from planted forests and that provided clear, specific in
tree form and timber quality for some industries, e.g., the veneer in formation about spacing and evaluation methods; (b) used tree density
dustry (Charlton et al., 2020; Glencross et al., 2012; Naji et al., 2016; as a factor affecting growth in planted stands, thus excluding studies
Tonini et al., 2018). Tree growth tends to reach a maximum at a certain having density as a dependent variable in natural forest stands; (c)
density and then declines with additional trees; however, this trend also evaluated planted tree species response, not the response of natural
depends on stand age, site quality, and environmental conditions (Will understory regeneration, shrubs, or herbaceous species; and (d) evalu
et al., 2001). As a key to understanding density-related patterns, Weiner ated tree morphological and physiological responses to planting density,
and Freckleton (2010) reviewed the empirical and mathematical basis of thereby excluding papers investigating wood quality or environmental
constant final yield. They concluded that constant final yield was the aspects, such as soil properties and understory species occurrence. After
most robust pattern in ecology and deemed it as essential for under this second group of criteria, we retained 255 papers for this review.
standing and predicting plant community growth. Intertree competition
is generally minimal for resources at very low tree densities, leading to a
2
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
We used descriptive statistics to analyze general trends related to 3. Results and discussion
publication frequency, study regions, study species, and spacing size. All
selected papers were evaluated, and we extracted and analyzed relevant 3.1. Evolution of planting density–growth studies between 2001 and 2020
information. We evaluated relationships between growth variables and
planting density. For this analysis, we considered the morphological and We observed an increasing trend in the number of publications
physiological characteristics of studied trees detailed in selected papers. focused on planting density and growth over the 1981–2020 period
These characteristics included total height, diameter at breast height, (Fig. 2). Publication abundance fluctuated between zero and 33 papers
individual and stand basal area, individual and stand timber volume, per year, with the highest peaks observed in 2019 and 2020, whereas we
leaf biomass, leaf area index, crown height and diameter, intercepted found only one planting density study in 1985,1986,1989, 1991, and
radiation, carbon assimilation, transpiration, respiration, water-use ef 2006, and we did not locate studies published in 1981, 1982, 1983,
ficiency, leaf water potential, and chlorophyll content. We extracted all 1987, and 1995. It is important to mention that planting density in forest
relevant information related to these variables, and for each paper we stands was already a focus of studies before 1981, but most papers are
classified our results as a neutral (*), positive (+), or negative (− ) not available online. A greater focus on increasing timber quality (i.e.,
response to spacing. From this classification, we produced tables pre straight shape and decay-resistant wood) and reducing rotation times
senting the studied species and the associated response of morphological drove the last 20-year increase in planting density studies, with financial
Fig. 2. Annual number of studies focused on planting density and its effect on tree growth published between 1981 and 2020.
3
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
costs and environmental change as a consequence of planting density Camellia sinensis L. (Kigalu, 2007a, 2007b), Hevea brasiliensis (Willd. ex
also being central elements in these papers (Bentancor et al., 2019; A. Juss.) Müll. Arg. (Naji et al., 2014), Fraxinus angustifolia Vahl
Charlton et al., 2020; Duan et al., 2019; Smith et al., 2016). Increasing (Özbayram and Çiçek, 2020), Gliricidia sepium (Jacq.) Kunth ex Walp.
planting density provides an easy means of increasing wood production, (Silva et al., 2012), Jatropha curcas L. (Andrianirina et al., 2019), and
especially in regions where areas available for forest plantations are Moringa oleifera Lam. (Mendieta-Araica et al., 2013). Thus, we cannot
scarce because of competing crops or resources availability. These identify how planting density affects many species, despite the major
concerns have been raised in the last decades by several international effort undertaken in the studies of selected economically important taxa;
institutions and worldwide agreements aimed at reducing deforestation the results of these species-specific studies cannot always be extrapo
and mitigating climate change, e.g., the FAO and the Paris Agreement. lated to other taxa.
Globally, studies of planting density were widespread, although We observed that planting densities differed between coniferous and
many countries had only a single study on their territory (Fig. 3). The broad-leaved species (Fig. 5). Planting density varied among species and
subject of planting density was most frequent in trials located in the ranged between 25 stems⋅ha− 1 in a Quercus robur L. Nelder Wheel trial
United States (US – 60), China (31), and Brazil (31). These countries (Uhl et al., 2015) to 167,000 stems⋅ha− 1 for Moringa oleifera Lam.
have a strong tradition of plantation forestry, although the species and (Mendieta-Araica et al., 2013). Differences in planting density related
products generated from these forests serve various industries—mainly mainly to the planned end products from these trials. Pinus spp., rep
timber, pulp and paper, and biomass for energy production. The US and resenting the commonly studied coniferous species in the reviewed
China lie in temperate climates, with studies mainly involving planted studies, are used mainly for timber production, especially for the
forests of coniferous species (Groot and Cortini, 2016; Harrington et al., sawmill industry. This industry requires larger-diameter timber, which
2009; Will et al., 2005, 2001; Zhao et al., 2020, 2011), whereas Brazil cannot be produced at high planting densities. Conversely, Eucalyptus
lies in the tropics and contains planted forests of various broad-leaved spp. and Salix spp., among other broad-leaved species, are used for pulp
species (Narducci et al., 2016; Páscoa et al., 2019; Schwerz et al., and paper or energy industries; smaller diameter boles can be used for
2020a, 2020b; Silva et al., 2012; Tonini et al., 2018). We thus observed a this purpose and be grown under conditions of shorter rotation periods
greater number of studies on Pinus spp. in the US and China, and more and higher planting densities. Studies with Camellia sinensis L. also use
studies on Eucalyptus spp. in Brazil. Other countries in which the forest higher densities because the main products of these plantations are the
industry is an important economic sector did not present high numbers leaves used in the tea industry (Kigalu, 2007a, 2007b).
of planting density studies because the studies focused on natural forests
did not have regular spacing between plants, explored thinning exper 3.2. How does planting density affect tree growth?
iments, and evaluated ecological variables other than growth. These
latter studies were removed from this review during the screening Studies of tree density in planted forest stands mainly address
process. biomass production and timber quality, which reflected the most
Species from the genera Pinus and Eucalyptus were the most studied commonly measured morphological variables: diameter at breast height
in planting density trials, representing 28 % and 19 % of the analyzed (DBH, 74 % of studies), height (H, 64 %), and stand volume (SV, 48 %).
studies, respectively (Fig. 4). Among the Pinus taxa, P. taeda L. was the Table 1 presents the growth response of trees species to increasing
most used in the trials, being found in 40 studies. The most-widely planting density. In general, there is a trade-off between tighter spacing
studied Eucalyptus species was E. grandis W. Hill ex Maiden, used in 18 and biomass production per area, and initial spacing strongly influences
studies. These abovementioned taxa are the most important species for plant growth at various times during stand development. Higher
plantations in several countries, and they are used by timber, pulp and planting densities reduce stem and/or crown diameter and, in most
paper, and energy industries. studies, there is a linear relationship between these growth variables
Nonetheless, the lack of information on tree species cultivated for (Benomar et al., 2012; Dassot et al., 2015; Han et al., 2020; Harrington
products other than wood was obvious from our literature search; we et al., 2009; Páscoa et al., 2019; Silva et al., 2012). Height growth tends
only found a handful of species evaluated for leaves and/or fruits: to increase with planting density; however, this trend is not consistent
Fig. 3. Location and abundance of published studies between 2001 and 2020 that assessed planting density effects on growth.
4
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
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 and an increase in total stand timber biomass, regardless of planting
sity–dependent variable, and soil fertility has also been deemed the most density. In a Eucalyptus plantation, Silva et al. (2019) observed differ
critical factor determining height growth (Henskens et al., 2001; ences in DBH and volume per tree between planting densities as early as
Woodruff et al., 2002); therefore, competition for soil nutrients could be 24 months after planting – an increase of 22 % in DBH and 90 % in
the main cause for reduced height growth for trees in high-density volume per tree in the widest spacing compared to the other planting
stands. Kholdaenko et al. (2022), for example, demonstrated that a spacings. The authors attributed the greater DBH and volume per tree to
high planting density in Picea obovate Ledeb. plantations reduced height the higher light interception by tree crowns within low-density planta
growth considerably in suppressed trees because of an earlier onset of tions, which led to a higher production of photoassimilates. Studies that
competition for nutrients. examined biomass allocation along gradients of light, water, and
In the initial years after planting, the interaction between trees is nutrient availability also found support for the hypothesis that plants
limited and competition may not be observed until a tree reaches the tend to allocate biomass to organs that acquire the most limiting re
above- and belowground space of its neighbor. Studies analyzed in our sources (Ma et al., 2021; McCarthy and Enquist, 2007; Rodríguez-García
review evaluated trees from 1 (Schwerz et al., 2019; Silva et al., 2019) to and Bravo, 2013); therefore, plants competing for resources in stands
90 (Węgiel and Polowy, 2020) years after planting, and results related to tend to change allocation patterns according to stand age and planting
the morphological characteristics were highly heterogeneous. For density. The greater allocation to the growth of branches and leaves in
Acacia mearnsii De Wild., Schwerz et al. (2019) observed a greater the first years form the canopy, important for light interception and
biomass allocation in branches and leaves until seven years after determining forest productivity (Barbier et al., 2010). Changes in tree
planting – an increase of 29 % in the narrowest spacing, compared to the growth after canopy closure are often related to competition with
widest spacing. Seven years after planting, they observed a change in the neighboring trees, which is more intense in high-density plantations
pattern of biomass partitioning, with a relative reduction in leaf biomass (Will et al., 2001).
5
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
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 S H DBH CH CD IV SV IBA SBA LB LAI Reference
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
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
8
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
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 Nonetheless, an increase in LAI or SLA is not always accompanied by
transport of auxin and cytokinin phytohormones drive tree growth and greater photosynthesis efficacy, as increased leaf expansion can produce
development in high-density stands. Auxins are known to drive plant thicker leaves having a lower stomatal density and, consequently,
growth and development—through the elongation of hypocotyl and reduced gas exchange potential (Mcelwain et al., 2016; Thyroff et al.,
petiole—and upward leaf movements, enhancing access to light (Galvão 2019).
and Fankhauser, 2015). The biosynthesis and transport of auxins are Competition for resources cannot always be managed by the addition
related to photoreceptor signaling (Küpers et al., 2020), and in high- of water and/or nutrients or the application of other silvicultural
density stands, auxins stimulate height growth rather than stem lateral treatments, and the effects of resource management are more likely to be
growth and branch development in the lower portion of trees, which observed for leaf, fruit, and seed yield than for wood (Kigalu, 2007a,
define plant shape. Auxin metabolism and its influence on plant growth 2007b; Mendieta-Araica et al., 2013; Narducci et al., 2016). For other
and development was extensively reviewed by Casanova-Sáez and Voβ species, fruit and seed yields are not influenced by density, independent
(2019). Cytokinin influence on plant growth and acclimation to the light of other resource management interventions, as observed for Pinus pinea
gradient in high-density stands relate to differences in transpiration L. (Loewe-Muñoz et al., 2020). This limited influence of density in
rates in shaded and sun-exposed leaves. The reduced influx of cytokinins dicates a potential to manage spacing arrangements in interaction with
in shaded leaves, resulting from a lower transpiration rate, decreases other silvicultural treatments to increase yield even when other re
protein synthesis and amino acid export, in turn inducing a reallocation sources become scarce. Establishing high-density plantations for these
of resources to sun-exposed leaves, which become stronger sinks non-timber taxa in sites having a high availability of water and nutrients
because of higher transpiration rates (Pons et al., 2001). Because of is a strategy for increasing productivity by reducing competition be
changes induced by these phytohormones, specific leaf area (SLA) is tween trees without increasing plantation costs (Harrington et al., 2009;
reduced in trees subject to high-density spacing, although the leaf area Sun et al., 2018; Toillon et al., 2013; Zhao et al., 2020).
index (LAI) can increase in these circumstances (Parker, 2020). The increase in total plantation biomass with concomitant reductions
In general, leaf expansion is reduced in shaded leaves and stimulated in individual tree biomass is an important aspect to consider for forest
in sunbathed ones, which directly influences the LAI and SLA (Pons producers. There is a potential of producing wood for energy in high-
et al., 2001). As a measure of canopy closure, LAI is indicative of stand density stands; however, for more valuable wood products, such as
productivity and biomass accumulation, and it can be an alternative lumber and veneer, these stands should be submitted to thinning man
measure of growing space and stocking control in plantations (O’Hara agement to produce larger boles. At high initial densities, thinning may
and York, 2014). However, few studies have considered this index be the best option for producers to ensure highly valuable wood prod
among measured growth variables. Only 24 studies in our survey (9 %, ucts at the end of rotation by regulating timber form and volume
Tab.1) considered LAI as a response variable to planting density, and all (Charlton et al., 2020). Smith et al. (2016) highlighted the need to
of these studies revealed a neutral or positive relationship with density evaluate the economic aspects and potential use of thinned stems, and
(Akers et al., 2013; Bullard et al., 2002; Dean and Baldwin, 1996; Farooq Glencross et al. (2012) also suggested coupling high planting densities
et al., 2020; Farque et al., 2001; Grigg et al., 2008; Hager and Kazda, with thinning treatments to obtain various products at different times to
1985; Hakamada et al., 2020a; Henskens et al., 2001; Johansson, 1989, ensure both a high volume for the stand and individual trees. Adapted
2007; Meinzer et al., 1996; O’Hara and York, 2014; de Paula et al., 2013; management strategies aim to ensure both wood and energy products in
Pinkard and Neilsen, 2003; Shi and Cao, 1997; Schwerz et al., 2019, forest plantations; these interventions include an increased tree density,
2020b; Tenorio et al., 2019; Tun et al., 2018; Wang et al., 2019; Will additional thinning, and a reduced rotation length to heighten financial
et al., 2001, 2005; Xue et al., 2012; Yan et al., 2019; Zhao et al., 2012). gains (Scott and Tiarks, 2008). Nonetheless, in plantations characterized
LAI is often used as an indicator of canopy closure and light interception by wider spacings or that are subjected to thinning, wind can penetrate
in agricultural crops (Sharma-Natu and Ghildiyal, 2005). Nonetheless, more easily, increasing mechanical stresses on residual trees and
in forest plantations we must consider that plant (tree) density is increasing tree vulnerability to windthrow (Girona et al., 2019; Scott
markedly different from that in agricultural plantations, and thus light and Mitchell, 2005).
interception differs between forest and field settings. Hakamada et al. Another important alternative to increase biomass production in
(2020a) indicated that increased LAI (90 % higher than in wider spac forest stands is species mixture, which create distinct interactions be
ings) in clonal Eucalyptus high-density plantations was directly related to tween individuals and provide growth advantages (Bello et al., 2019). As
light interception and consequently, to more efficient biomass accu an example, Forrester et al. (2004) observed that biomass accumulation
mulation. The authors also observed that genotype-related characteris in E. globulus increased when in admixture with A. mearnsii. In contrast,
tics influenced the clonal response to planting density, which is a key Bouillet et al. (2013) observed that changes in competitive or facilitation
factor to be considered in breeding programs. Akers et al. (2013) ob processes in monospecific stands or in mixed-stands of Eucalyptus spp.
tained comparable results for Pinus taeda L. plantations under various and A. mangium depended on ecological conditions, indicating the
planting densities. The authors attributed the increase in LAI (26 % importance of evaluating each species responses and other environ
higher in high-density planting) and light interception to a greater mental aspects such as water and nutrient availability. In our review, we
light-use efficacy in high-density stands relative to low-density stands. 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. Forest Ecology and Management 534 (2023) 120882
11
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. Forest Ecology and Management 534 (2023) 120882
DeBell, J.D., Gartner, B.L., 1997. Stem characteristics on the lower log of 35-year-old Glencross, K., Nichols, J.D., Grant, J.C., Sethy, M., Smith, R.G.B., 2012. Spacing affects
western redcedar grown at several spacings. West. J. Appl. For. 12, 9–14. stem form, early growth and branching in young whitewood (Endospermum
DeBell, D.S., Harrington, C.A., 2002. Density and rectangularity of planting influence 20- medullosum) plantations in Vanuatu. Int. For. Rev. 14, 442–451.
year growth and development of red alder. Can. J. For. Res. 32, 1244–1253. Goel, V.L., Behl, H.M., 2004. Productivity assessment of three leguminous species under
del Campo, A.D., Otsuki, K., Serengil, Y., Blanco, J.A., Yousefpour, R., Wei, X., 2022. high-density plantations on degraded soil sites. Biomass and Bioenergy 27, 403–409.
A global synthesis on the effects of thinning on hydrological processes: implications Gonçalves, J.L.D.M., Stape, J.L., Laclau, J.-P., Smethurst, P., Gava, J.L., 2004.
for forest management. For. Ecol. Manage. 519, 120324. Silvicultural effects on the productivity and wood quality of eucalypt plantations.
Di Matteo, G., Sperandio, G., Verani, S., 2012. Field performance of poplar for bioenergy For. Ecol. Manage. 193, 45–61.
in southern Europe after two coppicing rotations: effects of clone and planting Grigg, A.H., Macfarlane, C., Evangelista, C., Eamus, D., Adams, M.A., 2008. Does initial
density. IForest 5, 224–229. spacing influence crown and hydraulic architecture of Eucalyptus marginata? Tree
Domec, J.C., Ashley, E., Fischer, M., Noormets, A., Boone, J., Williamson, J.C., King, J.S., Physiol. 28, 753–760.
2017. Productivity, biomass partitioning, and energy yield of low-input short- Groot, A., Cortini, F., 2016. Effects of initial planting density on tree and stand
rotation american sycamore (Platanus occidentalis L.) grown on marginal land:effects development of planted black spruce up to age 30. For. Chron. 92, 200–209.
of planting density and simulated drought. Bioenergy Res. 10, 903–914. Grossiord, C., Gessler, A., Granier, A., Pollastrini, M., Bussotti, F., Bonal, D., 2014.
du Plessis, M., Kotze, H., 2011. Growth and yield models for Eucalyptus grandis grown in Interspecific competition influences the response of oak transpiration to increasing
Swaziland. South. For. 73, 81–89. drought stress in a mixed Mediterranean forest. For. Ecol. Manage. 318, 54–61.
Duan, A., Lei, J., Hu, X., Zhang, J., Du, H., Zhang, X., Guo, W., Sun, J., 2019. Effects of Guner, S., Yagci, V., Tilki, F., Celik, N., 2010. The effects of initial planting density on
planting density on soil bulk density, pH and nutrients of unthinned Chinese fir above-and below-ground biomass in a 25-year-old Fagus orientalis Lipsky plantation
mature stands in south subtropical region of China. Forests 10, 351. in Hopa. Turkey. Sci. Res. Essays 5, 1856–1860.
Eastham, J., Rose, C.W., Charles-Edwards, D.A., Cameron, D.M., Rance, S.J., 1990. Gyenge, J.E., Fernández, M.E., Schlichter, T.M., 2010. Effect of stand density and
Planting density effects on water use efficiency of trees and pasture in an pruning on growth of ponderosa pines in NW Patagonia. Argentina. Agrofor. Syst.
agroforestry experiment. New Zealand J. For. Sci. 20, 39–53. 78, 233–241.
Egbäck, S., Liziniewicz, M., Högberg, K.A., Ekö, P.M., Nilsson, U., 2012. Influence of Hager, H., Kazda, M., 1985. The influence of stand density and canopy position on sulfur
progeny and initial stand density on growth and quality traits of 21 year old half-sib content in needles of Norway spruce (Picea abies L. Karst.). Water, Air, and Soil
Scots pine (Pinus sylvestris L.). For. Ecol. Manage. 286, 1–7. Pollution 25, 321–329.
Egbäck, S., Bullock, B.P., Isik, F., McKeand, S.E., 2015. Height-diameter relationships for Hakamada RE, Hubbard RM, Stape JL, Lima W de P, Moreira GG, Ferraz SF de B 2020b
different genetic planting stock of loblolly pine at age 6. For. Sci. 61, 424–428. Stocking effects on seasonal tree transpiration and ecosystem water balance in a fast-
Eloy, E., da Silva, D.A., Schmidt, D., Trevisan, R., Caron, B.O., Elli, E.F., 2016. Effect of growing Eucalyptus plantation in Brazil. For. Ecol. Manage. 466:118149.
planting age and spacing on energy properties of Eucalyptus grandis W. Hill ex Hakamada, R., Hubbard, R.M., Ferraz, S., Stape, J.L., Lemos, C., 2017. Biomass
Maiden. Rev. Árvore 40, 749–758. production and potential water stress increase with planting density in four highly
Eloy, E., da Silva, D.A., Caron, B.O., Elli, E.F., Schwerz, F., 2018. Effect of age and productive clonal Eucalyptus genotypes. South. For. 79, 251–257.
spacing on biomass production in forest plantations. Rev. Árvore 42, e420214. Hakamada, R.E., Hubbard, R.M., Moreira, G.G., Stape, J.L., Campoe, O., Ferraz, S.F.de B.,
Erasmus, J., Wessels, C.B., 2020. The effect of stand density management on Pinus patula 2020a. Influence of stand density on growth and water use efficiency in Eucalyptus
lumber properties. Eur. J. For. Res. 139, 247–257. clones. For. Ecol. Manage. 466, 118125.
Erasmus, J., Kunneke, A., Drew, D.M., Wessels, C.B., 2018. The effect of planting spacing Han, Q., Zhang, Y., Geng, X., Du, K., Yang, J., Kang, X.A., Kang, X., 2020. Response of
on Pinus patula stem straightness, microfibril angle and wood density. Forestry 91, tree growth, crown, and branch development to planting density in four Populus
247–258. tomentosa clones. Aust. J. For. Sci. 3, 179–198.
Fang, S., Xue, J., Tang, L., 2007. Biomass production and carbon sequestration potential Hardanto, A., Röll, A., Hendrayanto, H.D., 2017. Tree soil water uptake and transpiration
in poplar plantations with different management patterns. J. Environ. Manage. 85, in mono-cultural and jungle rubber stands of Sumatra. For. Ecol. Manage. 397,
672–679. 67–77.
Food and Agriculture Organization of the United Nations (FAO) (2020a) Global forest Harms, W.R., Whitesell, C.D., DeBell, D.S., 2000. Growth and development of loblolly
resources assessment 2020: Main report. Food and Agriculture Organization of the pine in a spacing trial planted in Hawaii. For. Ecol. Manage. 126, 13–24.
United Nations, Rome, Italy, p 186. Harrington, T.B., Harrington, C.A., DeBell, D.S., 2009. Effects of planting spacing and site
Farooq, T.H., Yan, W., Chen, X., Shakoor, A., Rashid, M.H.U., Gilani, M.M., He, Z., quality on 25-year growth and mortality relationships of Douglas-fir (Pseudotsuga
Wu, P., 2020. Dynamics of canopy development of Cunninghamia lanceolata mid-age menziesii var. menziesii). For. Ecol. Manage. 258, 18–25.
plantation in relation to foliar nitrogen and soil quality influenced by stand density. Hébert, F., Krause, C., Plourde, P.Y., Achim, A., Prégent, G., Ménétrier, J., 2016. Effect of
Glob. Ecol. Conserv. 24, e01209. tree spacing on tree level volume growth, morphology, and wood properties in a 25-
Farque, L., Sinoquet, H., Colin, F., 2001. Canopy structure and light interception in year-old Pinus banksiana plantation in the boreal forest of Quebec. Forests 7, 276.
Quercus petraea seedlings in relation to light regime and plant density. Tree Physiol. Henskens, F.L., Battaglia, M., Cherry, M.L., Beadle, C.L., 2001. Physiological basis of
21, 1257–1267. spacing effects on tree growth and form in Eucalyptus globulus. Trees (Berl.) 15,
Feng, H., Xue, L., 2019. Competition-density effect of tree organs in Acacia auriculiformis 365–377.
stands. J. For. Res. 30, 891–898. Horner, G.J., Baker, P.J., Mac Nally, R., Cunningham, S.C., Thomson, J.R., Hamilton, F.,
Fernández-Núñez, E., Rigueiro-Rodríguez, A., Mosquera-Losada, M.R., 2010. Carbon 2009. Mortality of developing floodplain forests subjected to a drying climate and
allocation dynamics one decade after afforestation with Pinus radiata D. Don and water extraction. Glob. Change Biol. 15, 2176–2186.
Betula alba L. under two stand densities in NW Spain. Ecol. Eng. 36, 876–890. Huber, M., Nieuwendijk, N.M., Pantazopoulou, C.K., Pierik, R., 2021. Light signalling
Ferraz Filho, A.C., Mola-Yudego, B., González-Olabarria, J.R., Scolforo, J.R.S., 2018. shapes plant-plant interactions in dense canopies. Plant Cell Environ. 44,
Thinning regimes and initial spacing for Eucalyptus plantations in Brazil. An. Acad. 1014–1029.
Bras. Cienc. 90, 255–265. Huijuan, B., Jialei, Z., Runzhe, Z., Lishui, N., Lianjun, S., 2020. Effects of planting density
Ferreira, G.W.D., Rau, B.M., Aubrey, D.P., 2020. Herbicide, fertilization, and planting and clone type on growth and wood volume of Populus tomentosa stands. J. For. Res.
density effects on intensively managed loblolly pine early stand development. For. 25 (6), 444–449.
Ecol. Manage. 472. Hummel, S., 2000. Height, diameter and crown dimensions of Cordia alliodora associated
Finley, K., Zhang, J., 2019. Climate effect on ponderosa pine radial growth varies with with tree density. For. Ecol. Manage. 127, 31–40.
tree density and shrub removal. Forests 10, 477. Ilomäki, S., Nikinmaa, E., Mäkelä, A., 2003. Crown rise due to competition drives
Forrester, D.I., 2015. Transpiration and water-use efficiency in mixed-species forests biomass allocation in silver birch. Can. J. For. Res. 33, 2395–2404.
versus monocultures: effects of tree size, stand density and season. Tree Physiol. 35, Imada, M., Kunisaki, T., Mizoue, N., Teraoka, Y., 1997. Optimum planting density for
289–304. Japanese oak (Quercus mongolica var. grosseserrata) based on spacing experiment
Forrester, D.I., Smith, R.G.B., 2012. Faster growth of Eucalyptus grandis and Eucalyptus with systematic design. J. For. Res. 2, 89–93.
pilularis in mixed-species stands than monocultures. For. Ecol. Manage. 286, 81–86. Ingwers, M.W., Zhao, D., Kane, M.B., Markewitz, D., Teskey, R.O., 2018. Relationships
Forrester, D.I., Bauhus, J., Khanna, P.K., 2004. Growth dynamics in a mixed-species among growth, δ13C, foliar nitrogen concentration, foliar nitrogen content and
plantation of Eucalyptus globulus and Acacia mearnsii. For. Ecol. Manage. 193, 81–95. intercepted radiation at different cultural intensities, planting densities and site
Forrester, D.I., Wiedemann, J.C., Forrester, R.I., Baker, T.G., 2013. Effects of planting indices reveal the importance of water use efficiency in mid-rotation lobloll. For.
density and site quality on mean tree size and total stand growth of Eucalyptus Ecol. Manage. 422, 233–240.
globulus plantations. Can. J. For. Res. 43, 846–851. Isaac-Renton, M., Stoehr, M., Bealle Statland, C., Woods, J., 2020. Tree breeding and
Gadow, K.V., Kotze, H., 2014. Tree survival and maximum density of planted forests – silviculture: Douglas-fir volume gains with minimal wood quality loss under variable
Observations from South African spacing studies. For. Ecosyst. 1, 21. planting densities. For. Ecol. Manage. 465, 118094.
Galvão, V.C., Fankhauser, C., 2015. Sensing the light environment in plants: Johansson, T., 1989. Irradiance within canopies of young trees of european aspen
photoreceptors and early signaling steps. Curr. Opin. Neurobiol. 34, 46–53. (Populus tremula) and european birch (Betula pubescens) in stands of different
Gemmel, P., Nilsson, U., Welander, T., 1996. Development of oak and beech seedlings spacings. For. Ecol. Manage. 28, 217–236.
planted under varying shelterwood densities and with different site preparation Johansson, T., 2007. Biomass production and allometric above- and below-ground
methods in southern Sweden. New For. 12, 141–161. relations for young birch stands planted at four spacings on abandoned farmland.
Girona, M.M., Morin, H., Lussier, J.M., Ruel, J.C., 2019. Post-cutting mortality following Forestry 80, 41–52.
experimental silvicultural treatments in unmanaged boreal forest stands. Front. For. Kalinganire, A., 1996. Performance of Grevillea robusta in plantations and on farms under
Glob. Change 2, 4. varying environmental conditions in Rwanda. For. Ecol. Manage. 80, 279–285.
Gizachew, B., Brunner, A., Øyen, B.H., 2012. Stand responses to initial spacing in Kearney, D., James, R., Montagu, K., Smith, R.G.B., 2007. The effect of initial planting
Norway spruce plantations in Norway. Scand. J. For. Res. 27, 637–648. density on branching characteristics of Eucalyptus pilularis and E. grandis. Aust. For.
70, 262–268.
13
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
Kerr, G., 2003. Effects of spacing on the early growth of planted Fraxinus excelsior L. Can. McCarthy, M.C., Enquist, B.J., 2007. Consistency between an allometric approach and
J. For. Res. 33, 1196–1207. optimal partitioning theory in global patterns of plant biomass allocation. Funct.
Khanna, P.K., 1997. Comparison of growth and nutrition of young monocultures and Ecol. 21, 713–720.
mixed stands of Eucalyptus globulus and Acacia mearnsii. For. Ecol. Manage. 94, McClain, K.M., Morris, D.M., Hills, S.C., Buse, L.J., 1994. The effects of initial spacing on
105–113. growth and crown development for planted northern conifers: 37-year results. For.
Kholdaenko, Y.A., Belokopytova, L.V., Zhirnova, D.F., Upadhyay, K.K., Tripathi, S.K., Chron. 70, 174–182.
Koshurnikova, N.N., Sobachkin, R.S., Babushkina, E.A., Vaganov, E.A., 2022. Stand McElwain, J.C., Yiotis, C., Lawson, T., 2016. Using modern plant trait relationships
density effects on tree growth and climatic response in Picea obovata Ledeb. between observed and theoretical maximum stomatal conductance and vein density
plantations. For. Ecol. Manage. 519, 120349. to examine patterns of plant macroevolution. New Phytol. 209, 94–103.
Kigalu, J.M., 2007a. Effects of planting density on the productivity and water use of tea Medeiros, P.L., Silva, G.G.C., Oliveira, E.M.M., Ribeiro, C.O., Silva, J.M.S., Pimenta, A.S.,
(Camellia sinensis L.) clones. I. measurement of water use in young tea using sap flow 2020. Efficiency of nutrient use for biomass production of a Eucalyptus clone as a
meters with a stem heat balance method. Agric. Water Manage. 90, 224–232. function of planting density in short-rotation cropping. Aust. For. 83, 66–74.
Kigalu, J.M., 2007b. Effects of planting density and drought on the productivity of tea Mehari, A., Habte, B., 2006. Influence of initial spacing on growth and branching
clones (Camellia sinensis L.): Yield responses. Phys. Chem. Earth Parts ABC 32, characteristics of Cordia africana trees established on Eritrean highland. New For. 31,
1098–1106. 185–193.
Kirongo, B.B., Mbelase, A.S., Senelwa, K., Hitimana, J., Etiegni, L., 2012. Spacing and Meinzer, F.C., Fownes, J.H., Harrington, R.A., 1996. Growth indices and stomatal control
genotype on height and diameter growth of four Eucalyptus under short rotation. of transpiration in Acacia koa stands planted at different densities. Tree Phys. 16,
J. Trop. For. Manag. 18, 1–9. 607–615.
Klein, T., Hartmann, H., 2018. Climate change drives tree mortality. Science 362, 758. Mendieta-Araica, B., Spörndly, E., Reyes-Sánchez, N., Salmerón-Miranda, F., Halling, M.,
Kumar, R., Lata, K., Khadda, B.S., Jadav, J.K., Rai, A.K., Khajuria, S., 2017. Influence of 2013. Biomass production and chemical composition of Moringa oleifera under
planting density on vegetative morphomatrix, yield and economics of drumstick different planting densities and levels of nitrogen fertilization. Agrofor. Syst. 87,
(Moringa oleifera) cv. PKM 1 under rainfed condition of semi-arid ecosystem. Indian 81–92.
J. Agric. Sci. 87, 866–869. Metsaranta, J.M., 2019. Long-term tree-ring derived carbon dynamics of an experimental
Küpers, J.J., Oskam, L., Pierik, R., 2020. Photoreceptors regulate plant developmental plantation in relation to species and density in Northwestern Ontario. Canada. For.
plasticity through auxin. Plants 9, 940. Ecol. Manage. 441, 229–241.
Lam, T.Y., Guan, B.T., 2020. Modeling stand basal area growth of Cryptomeria japonica D. Milyutina, I.L., Sudachkova, N.E., Romanova, L.I., 2013. Response of the antioxidant
Don under different planting densities in Taiwan. J. For. Res. 25, 174–182. system of light-demanding and shade-bearing pine species to phytocenotic stress.
Larsen, S.U., Jørgensen, U., Lærke, P.E., 2019. Harvest interval and row spacing of SRC Contemp. Probl. Ecol. 6, 149–155.
willow influence yield and nutrient content. Biomass and Bioenergy 126, 181–189. Miranda, I., Gominho, J., Pereira, H., 2015. Heartwood, sapwood and bark variation in
Lasserre, J.P., Mason, E.G., Watt, M.S., 2005. The effects of genotype and spacing on coppiced Eucalyptus globulus trees in 2nd rotation and comparison with the single-
Pinus radiata [D. Don] corewood stiffness in an 11-year old experiment. For. Ecol. stem 1st rotation. Silva Fenn. 49, 1141.
Manage. 205, 375–383. Moher D, Liberati A, Tetzlaff J, Altman DG, Group P 2009 Preferred reporting items for
Lee, Y.-J., Lenhart, J.D., Temple, A., 1997. Influence of planting density on diameter and systematic reviews and meta-analyses: The PRISMA statement. PLoS Med. 6:
height in East Texas pine plantations. South. J. Appl. For. 22, 241–244. e1000097.
Lima RLS, Campos ARF, Azevedo CAV de, Calado JAW, Silva S de S, Nascimento R do Moore, G.W., Bond, B.J., Jones, J.A., 2011. A comparison of annual transpiration and
2016 Effects of planting density on vegetative growth and production components of productivity in monoculture and mixed-species Douglas-fir and red alder stands. For.
jatropha (Physic nut L). Aust. J. Crop Sci. 10:632–636. Ecol. Manage. 262, 2263–2270.
Lin, Y., Yang, H., Ivković, M., Gapare, W.J., Colin Matheson, A., Wu, H.X., 2013. Effect of Nagar, B., Rawat, S., Rathiesh, P., Sekar, I., 2015. Impact of initial spacing on growth and
genotype by spacing interaction on radiata pine genetic parameters for height and yield of Eucalyptus camaldulensis in arid region of India. World Appl. Sci. J. 33,
diameter growth. For. Ecol. Manage. 304, 204–211. 1362–1368.
Liu, F., Gao, C., Chen, M., Li, K., 2018. Above- and below-ground biomass relationships Naji, H.R., Bakar, E.S., Soltani, M., Ebadi, S.E., Abdul-Hamid, H., Javad, S.K.S., Sahri, M.
of Leucaena leucocephala (Lam.) de Wit in different plant stands. PLoS One 13, H., 2014. Effect of initial planting density and tree features on growth, wood density,
e0207059. and anatomical properties from a Hevea brasiliensis trial plantation. For. Prod. J. 64,
Liu, L., Li, Y., Zhang, J., Sun, H., 2019. Impact of initial planting density on the optimal 41–47.
economic rotation of Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) in an Naji, H.R., Nia, M.F., Kiaei, M., Abdul-Hamid, H., Soltani, M., Faghihi, A., 2016. Effect of
experimental forest plantation. Forests 10, 713. intensive planting density on tree growth, wood density and fiber properties of
Liziniewicz, M., Ekö, P.M., Agestam, E., 2012. Effect of spacing on 23-year-old lodgepole maple (Acer velutinum Boiss.). iForest –. Biogeosci. For. 9, 325–329.
pine (Pinus contorta Dougl. var. latifolia) in southern Sweden. Scand. J. For. Res. 27, Nan, W., Ta, F., Meng, X., Dong, Z., Xiao, N., 2020. Effects of age and density of Pinus
361–371. sylvestris var. mongolica on soil moisture in the semiarid Mu Us Dunefield, northern
Loewe-Muñoz, V., Balzarini, M., Del Río, R., Delard, C., 2019. Effects of stone pine (Pinus China. For. Ecol. Manage. 473, 118313.
pinea L.) plantation spacing on initial growth and conelet entry into production. New Narducci, T.S., Yared, J.A.G., Júnior, S.B., 2016. Growth and survival of Sclerolobium
For. 50, 489–503. paniculatum Vogel and the relationship between rainfall and the increment in
Loewe-Muñoz, V., Del Río, R., Balzarini, M., 2020. Stone pine (Pinus pinea) growth and diameter at different planting spacings. Rev. Árvore 40, 447–454.
cone yield as a function of planting density in Chile. Bosque 41, 381–386. Negrave, R.W., Prescott, C.E., Barker, J.E., 2007. Growth and foliar nutrition of juvenile
Lokmal, N., Zaki, A.M., Fazwa, M.A.F., Suhaimi, W.C., Azmy, Y., Zakaria, I., Tan, H., western hemlock and western redcedar plantations on low- and medium-
Khoo, S.K., Wan-Akil, T.A., 2008. Growth of several rubber clones for timber productivity sites on northern Vancouver Island: response to fertilization and
production. J. Trop. For. Sci. 20, 175–180. planting density. Can. J. For. Res. 37, 2587–2599.
Long, J.N., Smith, F.W., 1988. Leaf area - sapwood area relations of lodgepole pine as Nelson, A.S., Wagner, R.G., Day, M.E., Fernandez, I.J., Weiskittel, A.R., Saunders, M.R.,
influenced by stand density and site index. Can. J. For. Res. 18, 247–250. 2016. Light absorption and light-use efficiency of juvenile white spruce trees in
Looney, C.E., Zhang, J., 2022. Site quality and intensive early stand management natural stands and plantations. For. Ecol. Manage. 376, 158–165.
practices affect growth dominance, structural complexity, and tree growth in Newton, P.F., 2015. Occurrence of density-dependent height repression within jack pine
ponderosa pine plantations. For. Ecol. Manage. 519, 120318. and black spruce populations. Forests 6, 2450–2468.
Luo, J., He, M., Qi, S., Wu, J., Gu, X.S., 2018. Effect of planting density and harvest Nilsson, U., 1994. Development of growth and stand structure in Picea abies stands
protocol on field-scale phytoremediation efficiency by Eucalyptus globulus. Environ. planted at different initial densities. Scandinavian J. For. Res. 9, 135–142.
Sci. Pollut. Res. 25, 11343–11350. Nilsson, U., Albrektson, A., 1994. Growth and self-thinning in two young Scots pine
Ma, S., Wang, X., Miao, W., Wang, X., Sun, H., Guo, Z., 2021. Relative influence of stands planted at different initial densities. For. Ecol. Manage. 68, 209–215.
environmental, stand factors and functional traits on allocation of forest productivity Ning K, Ding C, Huang Q, Zhang W, Yang C, Liang D, Fan R, Su X 2019 Transcriptome
during the restoration of subtropical forests in central China. For. Ecol. Manage. 482, profiling revealed diverse gene expression patterns in poplar (Populus ×
118814. euramericana) under different planting densities. PLoS ONE, 14.
Mabapa, M.P., Ayisi, K.K., Mariga, I.K., 2018. Seasonal effect on Moringa oleifera gaseous Nirsatmanto, A., 2016. Early growth of improved Acacia mangium at different planting
exchange and water use efficiency under diverse planting densities. J. Appl. Bot. densities. Jurnal Manajemen Hutan Tropika 22, 105–113.
Food. Qual. 91, 219–225. O’Hara, K.L., York, R.A., 2014. Leaf area development and crown architecture in a giant
Marziliano, P.A., Coletta, V., Menguzzato, G., Nicolaci, A., Pellicone, G., Veltri, A., 2015. sequoia spacing study. For. Sci. 60, 776–783.
Effects of planting density on the distribution of biomass in a douglas-fir plantation Okojie, J.A., Bailey, R.L., Borders, B.E., 1988. Spacing effects in an unthinned 11-year-
in southern Italy. iForest – Biogeosci. For. 8, 368–376. old Terminalia superba plantation in the dry lowland rainforest area of Nigeria. For.
Marziliano, P.A., Menguzzato, G., Scuderi, A., Scalise, C., Coletta, V., 2017. Biomass Ecol. Manage. 23, 253–260.
conversion and expansion factors in Douglas-fir stands of different planting density: Ola-Adams, B.A., 1993. Effects of spacing on biomass distribution and nutrient content of
variation according to individual growth and prediction equations. For. Syst. 26, Tectona grandis Linn. f. (teak) and Terminalia superba Engl. Diels. (afara) in south-
e003. western Nigeria. For. Ecol. Manage. 58, 299–319.
Matos FA, Daniel O, Serra AP, Heid DM, Nascimento JM do, Nogueira IMB, Ensinas SC, Oliveira, N., del Río, M., Forrester, D.I., Rodríguez-Soalleiro, R., Pérez-Cruzado, C.,
Altomar PH, Conrad V do A, Potrich DC, Martinez MA 2015 Effects of intra-specific Cañellas, I., Sixto, H., 2018. Mixed short rotation plantations of Populus alba and
tree competition on dendrometric parameters of Peltophorum dubium. Aust. J. Crop Robinia pseudoacacia for biomass yield. For. Ecol. Manage. 410, 48–55.
Sci. 9:1003–1009. Olmos, V.M., 2022. Forestry and the forest products sector: Production, income and
Mazza, G., Cutini, A., Manetti, M.C., 2015. Influence of tree density on climate-growth employment, and international trade. Forest Policy Econ. 135, 102648.
relationships in a Pinus pinaster Ait. forest in the northern mountains of Sardinia Özbayram, A.K., Çiçek, E., 2020. Initial planting density experiments of narrow-leaved
(Italy). IForest 8, 456–463. ash in Turkey: ten-year results. Sumar. List 144, 269–278.
14
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
Pachas, A.N.A., Shelton, H.M., Lambrides, C.J., Dalzell, S.A., Murtagh, G.J., Hardner, C. Roth, B.E., Li, X., Huber, D.A., Peter, G.F., 2007b. Effects of management intensity,
M., 2018. Effect of tree density on competition between Leucaena leucocephala and genetics and planting density on wood stiffness in a plantation of juvenile loblolly
Chloris gayana using a Nelder Wheel trial II. Belowground interactions. Crop Pasture pine in the southeastern USA. For. Ecol. Manage. 246, 155–162.
Sci. 69, 733–744. Russell, M.B., Burkhart, H.E., Amateis, R.L., 2009. Biomass partitioning in a miniature-
Pachas, A.N.A., Sakanphet, S., Soukkhy, O., Lao, M., Savathvong, S., Newby, J.C., scale loblolly pine spacing trial. Can. J. For. Res. 39, 320–329.
Souliyasack, B., Keoboualapha, B., Dieters, M.J., 2019. Initial spacing of teak Schwerz, F., Eloy, E., Elli, E.F., Caron, B.O., 2019. Reduced planting spacing increase
(Tectona grandis) in northern Lao PDR: impacts on the growth of teak and companion radiation use efficiency and biomass for energy in black wattle plantations: towards
crops. For. Ecol. Manage. 435, 77–88. sustainable production systems. Biomass Bioenergy 120, 229–239.
Pallett, R.N., Sale, G., 2004. The relative contributions of tree improvement and cultural Schwerz, F., Neto, D.D., Caron, B.O., Nardini, C., Sgarbossa, J., Eloy, E., Behling, A.,
practice toward productivity gains in Eucalyptus pulpwood stands. For. Ecol. Elli, E.F., Reichardt, K., 2020a. Biomass and potential energy yield of perennial
Manage. 193, 33–43. woody energy crops under reduced planting spacing. Renew. Energy 153,
Pannacci, E., Bartolini, S., Covarelli, G., 2009. Evaluation of four poplar clones in a short 1238–1250.
rotation forestry in central Italy. Ital. J. Agron. 4, 191–198. Schwerz, F., Neto, D.D., Caron, B.O., Tibolla, L.B., Sgarbossa, J., Eloy, E., Elli, E.F.,
Parker, G.G., 2020. Tamm review: Leaf Area Index (LAI) is both a determinant and a Carvalho, L.G., 2020b. Carbon stocks, partitioning, and wood composition in short-
consequence of important processes in vegetation canopies. For. Ecol. Manage. 477, rotation forestry system under reduced planting spacing. Ann. For. Sci. 77, 67.
118496. Scolforo, J.R.S., Altoé, T.F., Scolforo, H.F., de Mello, J.M., Silva, C.P.C.e., Ferraz Filho, A.
Paula, R.R., Reis, G.G., Reis, M.G.F., Oliveira Neto, S.N., Leite, H.G., Melido, R.C.N., C., 2016. Management strategies of Eremanthus erythropappus (DC.) MacLeish under
Lopes, H.N.S., Souza, F.C., 2013. Eucalypt growth in monoculture and silvopastoral different initial spacing. Ciência e Agrotecnologia 40, 298–304.
systems with varied tree initial densities and spatial arrangements. Agrofor. Syst. 87, Scott, W., Meade, R., Leon, R., Hyink, D., Miller, R., 1998. Planting density and tree-size
1295–1307. relations in coast Douglas-fir. Can. J. For. Res. 28 (1), 74–78.
Penner, M., Robinson, C., Burgess, D., 2001. Pinus resinosa product potential following Scott, R.E., Mitchell, S.J., 2005. Empirical modelling of windthrow risk in partially
initial spacing and subsequent thinning. For. Chron. 77, 129–139. harvested stands using tree, neighbourhood, and stand attributes. For. Ecol. Manage.
Peracca, G.G., O’Hara, K.L., 2008. Effects of growing space on growth for 20-year-old 218, 193–209.
giant sequoia, ponderosa pine, and Douglas-fir in the Sierra Nevada. West. J. Appl. Scott, D.A., Tiarks, A., 2008. Dual-cropping loblolly pine for biomass energy and
For. 23, 156–165. conventional wood products. South. J. Appl. For. 32, 33–37.
Pérez, I., Pérez, J., Carrasco, J., Ciria, P., 2014. Siberian elm responses to different Shainsky, L.J., Newton, M., Radosevich, S.R., 1992. Effects of intra- and inter-specific
culture conditions under short rotation forestry in Mediterranean areas. Turkish J. competition on root and shoot biomass of young Douglas-fir and red alder. Can. J.
Agric. For. 38, 652–662. For. Res. 22, 101–110.
Pinkard, E.A., Neilsen, W.A., 2003. Crown and stand characteristics of Eucalyptus nitens Sharma-Natu, P., Ghildiyal, M.C., 2005. Potential targets for improving photosynthesis
in response to initial spacing: implications for thinning. For. Ecol. Manage. 172, and crop yield. Curr. Sci. 88, 1918–1928.
215–227. Shi, K., Cao, Q., 1997. Predicted leaf area growth and foliage efficiency of loblolly pine
Pons, T.L., Jordi, W., Kuiper, D., 2001. Acclimation of plants to light gradients in leaf plantations. For. Ecol. Manage. 95, 109–115.
canopies: evidence for a possible role for cytokinins transported in the transpiration Silva, W.G., dos Santos, D.M.C.L., Lima, A.P.L., Mattos, F.D., Lima, S.F., de Paula, R.C.M.,
stream. J. Exp. Bot. 52, 1563–1574. 2019. Growth and volumetric production of eucalyptus clones in different planting
Prasad, J.V.N.S., Korwar, G.R., Rao, K.V., Mandal, U.K., Rao, G.R., Srinivas, I., spaces. Rev. Agric. Neotrop. 6, 38–47.
Venkateswarlu, B., Rao, S.N., Kulkarni, H.D., 2011. Optimum stand density of Silva PSL, Paiva HN de, Oliveira VR de, Siqueira PL de OF, Soares EB, Monteiro AL,
Leucaena leucocephala for wood production in Andhra Pradesh, Southern India. Tavella LB 2014 Biomass of three species as a response to planting density and
Biomass Bioenergy 35, 227–235. interspecific competition. Rev. Árvore 38:319–329.
Quer, E., Baldy, V., DesRochers, A., 2020. Ecological drivers of root grafting in balsam fir Silva, P.S.L., Holanda, A.E.R., de Paiva, H.N., de Oliveira, F.H.T., de Oliveira, O.F., 2012.
natural stands. For. Ecol. Manage. 475, 118388. Planting density and initial growth of two tree species adapted to the semi-arid
Quicke, H., Glover, G., Glover, R., 1999. Loblolly pine growth response to herbaceous region. Rev. Árvore 36, 951–959.
vegetation control at different planting densities. Can. J. For. Res. 29, 960–967. Smith, R.G.B., Glencross, K., Nichols, J.D., Palmer, G., Viranamangga, R., 2016. Effect of
Radtke, P.J., Burkhart, H.E., 1999. Basal area growth and crown closure in a loblolly pine initial spacing on first thinning product recovery, and financial outcomes in
spacing trial. For. Sci. 45, 35–44. whitewood (Endospermum medullosum) plantations in Vanuatu. Small-scale For. 15,
Rai, R.S., Srinivasan, V.M., 1990. High density short rotation studies in Eucalyptus 45–59.
tereticornis and Casuarina equisetifolia. Int. Crops J. 6, 113–122. Snowdon, P., Woollons, R.C., Benson, M.L., 1998. Incorporation of climatic indices into
Ramalho, F.M.G., Pimenta, E.M., Goulart, C.P., de Almeida, M.N.F., Vidaurre, G.B., models of growth of Pinus radiata in a spacing experiment. New For. 16, 101–123.
Hein, P.R.G., 2019. Effect of stand density on longitudinal variation of wood and Srivastava, N., Goel, V.L., Behl, H.M., 1999. Influence of planting density on growth and
bark growth in fast-growing Eucalyptus plantations. iForest – Biogeosci. For. 12, biomass productivity of Terminalia arjuna under sodic soil sites. Biomass and
527–532. Bioenergy 17, 273–278.
Ramos-Trejo OS, Canul-Solís JR, Alvarado-Canché A del R, Castillo-Sánchez LE, Stape, J.L., Binkley, D., 2010. Insights from full-rotation Nelder spacing trials with
Sandoval-Gío JJ, Campos-Navarrete MJ, Piñero-Vázquez AT, Chay-Canul AJ, Eucalyptus. South. For. 72, 90–97.
Casanova-Lugo F (2020) Growth, forage yield and quality of Morus alba L. and Stape, J.L., Binkley, D., Ryan, M.G., 2004. Eucalyptus production and the supply, use and
Gliricidia sepium (Jacq.) Walp. in mixed and pure fodder bank systems in Yucatan, efficiency of use of water, light and nitrogen across a geographic gradient in Brazil.
México. Agrofor. Syst. 94:151–157. For. Ecol. Manage. 193, 17–31.
Rédei, K., 2009. Effects of initial spacing on the stand structure and yield of young black Štefančík, I., 2020. Development of the Norway spruce (Picea abies /L./ Karst.) stand
locust (Robinia pseudoacacia L.) stands. Int. J. Hortic. Sci. 15. established by various spacings and affected by abiotic harmful factors and ungulate
Remaury, A., Guittonny, M., Rickson, J., 2019. The effect of tree planting density on the game. J. For. Sci. 66, 117–131.
relative development of weeds and hybrid poplars on revegetated mine slopes Stolarski, M.J., Szczukowski, S., Tworkowski, J., Krzyżaniak, M., Załuski, D., 2017.
vulnerable to erosion. New For. 50, 555–572. Willow biomass and cuttings’ production potential over ten successive annual
Resende, R.T., Soares, A.A.V., Forrester, D.I., Marcatti, G.E., dos Santos, A.R., harvests. Biomass and Bioenergy 105, 230–247.
Takahashi, E.K., Silva, F.F.e., Grattapaglia, D., Resende, M.D.V., Leite, H.G., 2018. Stolarski, M.J., Szczukowski, S., Tworkowski, J., Krzyżaniak, M., 2019a. Extensive
Environmental uniformity, site quality and tree competition interact to determine willow biomass production on marginal land. Polish J. Environ. Stud. 28,
stand productivity of clonal Eucalyptus. For. Ecol. Manage. 410, 76–83. 4359–4367.
Resquin F, Navarro-Cerrillo RM, Rachid-Casnati C, Hirigoyen A, Carrasco-Letelier L, Stolarski, M.J., Szczukowski, S., Tworkowski, J., Krzyżaniak, M., Załuski, D., 2019b.
Duque-Lazo J 2018 Allometry, growth and survival of three eucalyptus species Willow production during 12 consecutive years—The effects of harvest rotation,
(Eucalyptus benthamii Maiden and Cambage, E. dunnii Maiden and E. grandis Hill ex planting density and cultivar on biomass yield. GCB Bioenergy 11, 635–656.
Maiden) in high-density plantations in Uruguay. Forests 9:745. Subedi, S., Kane, M., Zhao, D., Borders, B., Greene, D., 2012. Cultural intensity and
Resquin, F., Navarro-Cerrillo, R.M., Carrasco-Letelier, L., Casnati, C.R., 2019. Influence planting density effects on aboveground biomass of 12-year-old loblolly pine trees in
of contrasting stocking densities on the dynamics of above-ground biomass and the Upper Coastal Plain and Piedmont of the southeastern United States. For. Ecol.
wood density of Eucalyptus benthamii, Eucalyptus dunnii, and Eucalyptus grandis for Manage. 267, 157–162.
bioenergy in Uruguay. For. Ecol. Manage. 438, 63–74. Sun, H., Diao, S., Liu, R., Forrester, D., Soares, A., Saito, D., Dong, R., Jiang, J., 2018.
Resquin, F., Navarro-Cerrillo, R.M., Carrasco-Letelier, L., Casnati, C.R., Bentancor, L., Relationship between size inequality and stand productivity is modified by self-
2020. Evaluation of the nutrient content in biomass of Eucalyptus species from short thinning, age, site and planting density in Sassafras tzumu plantations in central
rotation plantations in Uruguay. Biomass and Bioenergy 134, 105502. China. For. Ecol. Manage. 422, 199–206.
Rodríguez-García, E., Bravo, F., 2013. Plasticity in Pinus pinaster populations of diverse Tanabe, J., Endo, R., Kuroda, S., Ishiguri, F., Narisawa, T., Takashima, Y., 2019. Variance
origins: Comparative seedling responses to light and nitrogen availability. For. Ecol. components and parent–offspring correlations of growth traits vary among the initial
Manage. 307, 196–205. planting spacings in Zelkova serrata. Silvae Genet. 68, 45–50.
Rončević, S., Andrašev, S., Ivanišević, P., Kovačević, B., Klašnja, B., 2013. Biomass Tarroux, E., Desrochers, A., 2011. Effect of natural root grafting on growth response of
production and energy potential of some eastern cottonwood (Populus deltoides Bartr. jack pine (Pinus banksiana; Pinaceae). Am. J. Bot. 98, 967–974.
ex Marsh.) clones in relation to planting spacing. Sumar. List 137, 33–42. Tarroux, E., Desrochers, A., Krause, C., 2010. Effect of natural root grafting on growth
Roth, B.E., Jokela, E.J., Martin, T.A., Huber, D.A., White, T.L., 2007a. Genotype × response of jack pine (Pinus banksiana) after commercial thinning. For. Ecol.
environment interactions in selected loblolly and slash pine plantations in the Manage. 260, 526–535.
Southeastern United States. For. Ecol. Manage. 238, 175–188. 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.
15
M.M. Gabira et al. Forest Ecology and Management 534 (2023) 120882
Tenorio, C., Roque, R.M., Valverde, J.C., Arias-Aguilar, D., 2019. Biomass production Will, R.E., Barron, G.A., Burkes, E.C., Shiver, B., Teskey, R.O., 2001. Relationship
and characteristics of short rotation plantations of clones of Gmelina arborea in three between intercepted radiation, net photosynthesis, respiration, and rate of stem
spacings. Silvae Genet. 68 (1), 92–100. volume growth of Pinus taeda and Pinus elliottii stands of different densities. For. Ecol.
Thoranisorn, S., Sahunalu, P., Yoda, K., 1990. Density effects and sel-thinning in even- Manage. 154, 155–163.
aged pure stands of Eucalyptus camaldulensis Dehn. The Botanical Magazine 103, Will, R.E., Narahari, N.V., Shiver, B.D., Teskey, R.O., 2005. Effects of planting density on
283–295. canopy dynamics and stem growth for intensively managed loblolly pine stands. For.
Thyroff, E.C., Burney, O.T., Mickelbart, M.V., Jacobs, D.F., 2019. Unraveling shade Ecol. Manage. 205, 29–41.
tolerance and plasticity of semi-evergreen oaks: Insights from maritime forest live Will, R.E., Hennessey, T., Lynch, T., Holeman, R., Heinemann, R., 2010. Effects of
oak restoration. Front. Plant Sci. 10, 1526. planting density and seed source on loblolly pine stands in Southeastern Oklahoma.
Toillon, J., Fichot, R., Dallé, E., Berthelot, A., Brignolas, F., Marron, N., 2013. Planting For. Sci. 56, 437–443.
density affects growth and water-use efficiency depending on site in Populus deltoides Wilson, J.B., Steel, J.B., Steel, S.L.K., 2007. Do plants ever compete for space? Folia
× P. nigra. For. Ecol. Manage. 304, 345–354. Geobot. 42, 431–436.
Tong, Q.J., Zhang, S.Y., 2005. Impact of initial spacing and precommercial thinning on Woodruff, D.R., Bond, B.J., Ritchie, G.A., Scott, W., 2002. Effects of stand density on the
jack pine tree growth and stem quality. For. Chron. 81, 418–428. growth of young Douglas-fir trees. Can. J. For. Res. 32, 420–427.
Tonini, H., Schwengber, D.R., Morales, M.M., Magalhães, C.A.S., de Oliveira, J.M.F., Xue, L., Jacobs, D.F., Zeng, S., Yang, Z., Guo, S., Liu, B., 2012. Relationship between
2018. Growth, biomass, and energy quality of Acacia mangium timber grown at above-ground biomass allocation and stand density index in Populus xeuramericana
different spacings. Pesqui. Agropecu. Bras. 53, 791–799. stands. Forestry 85, 611–619.
Torreano, S.J., Smalley, G.W., Nunley, A.N., Rennie, J.C., 2012. Structure and growth of Yan, Z., Zhou, Q., Teng, M., Ji, H., Zhang, J., He, W., Ye, Y., Wang, B., Wang, P., 2019.
40-year-old unthinned loblolly pine planted at three spacings in south-central High planting density and leaf area index of masson pine forest reduce crown
Tennessee. South. J. Appl. For. 36, 136–140. transmittance of photosynthetically active radiation. Glob. Ecol. Conserv. 20,
Truax, B., Fortier, J., Gagnon, D., Lambert, F., 2018. Planting density and site effects on e00759.
stem dimensions, stand productivity, biomass partitioning, carbon stocks and soil Yáñez, M.A., Fox, T.R., Seiler, J.R., 2015. Early growth responses of loblolly pine
nutrient supply in hybrid poplar plantations. Forests 9, 293. varieties and families to silvicultural intensity. For. Ecol. Manage. 356, 204–215.
Tsamir, M., Gottlieb, S., Preisler, Y., Rotenberg, E., Tatarinov, F., Yakir, D., Tague, C., Yáñez, M.A., Zamudio, F., Espinoza, S., Ivković, M., Guerra, F., Espinosa, C., Baettig, R.
Klein, T., 2019. Stand density effects on carbon and water fluxes in a semi-arid M., 2019. Genetic variation and growth stability of hybrid poplars in high-density
forest, from leaf to stand-scale. For. Ecol. Manage. 453, 117573. short-rotation coppice stands in central Chile. Biomass Bioenergy 120, 84–90.
Tun, T.N., Guo, J., Fang, S., Tian, Y., 2018. Planting spacing affects canopy structure, York RA, O’hara KL, Battles JJ 2013 Density effects on giant sequoia (Sequoiadendron
biomass production and stem roundness in poplar plantations. Scand. J. For. Res. 33, giganteum) growth through 22 years:Implications for restoration and plantation
464–474. management. West. J. Appl. For. 28:30–36.
Uhl, E., Biber, P., Ulbricht, M., Heym, M., Horváth, T., Lakatos, F., Gál, J., Steinacker, L., Załuski, D., Mielniczuk, J., Bronowicka-Mielniczuk, U., Stolarski, M.J., Krzyżaniak, M.,
Tonon, G., Ventura, M., Pretzsch, H., 2015. Analysing the effect of stand density and Szczukowski, S., Tworkowski, J., 2018. Survival analysis of plants grown in long-
site conditions on structure and growth of oak species using Nelder trials along an term field experiments. Agron. J. 110, 1791–1798.
environmental gradient: experimental design, evaluation methods, and results. For. Zhang, X., Cao, Q.V., Duan, A., Zhang, J., 2017. Modeling tree mortality in relation to
Ecosyst. 2, 1–19. climate, initial planting density and competition in Chinese fir plantations using a
Ungar, E.D., Rotenberg, E., Raz-Yaseef, N., Cohen, S., Yakir, D., Schiller, G., 2013. Bayesian logistic multilevel method. Can. J. For. Res. 47, 1278–1285.
Transpiration and annual water balance of Aleppo pine in a semiarid region: Zhang, J., Finley, K.A., Johnson, N.G., Ritchie, M.W., 2019. Lowering stand density
implications for forest management. For. Ecol. Manage. 298, 39–51. enhances resiliency of ponderosa pine forests to disturbances and climate change.
Waghorn, M.J., Mason, E.G., Watt, M.S., 2007. Influence of initial stand density and For. Sci. 65, 496–507.
genotype on longitudinal variation in modulus of elasticity for 17-year-old Pinus Zhang, Y., Tian, Y., Ding, S., Lv, Y., Samjhana, W., Fang, S., 2020c. Growth, carbon
radiata. For. Ecol. Manage. 252, 67–72. storage, and optimal rotation in poplar plantations: a case study on clone and
Waghorn, M.J., Watt, M.S., 2013. Stand variation in Pinus radiata and its relationship planting spacing effects. Forests 11, 842.
with allometric scaling and critical buckling height. Ann. Bot. 111, 675–680. Zhang, X., Wang, Z., Chhin, S., Wang, H., Duan, A., Zhang, J., 2020b. Relative
Wang, C.-S., Zeng, J., Hein, S., Zhao, Z.-G., Guo, J.-J., Zeng, J., 2016. Crown and branch contributions of competition, stand structure, age, and climate factors to tree
attributes of mid-aged Betula alnoides plantations in response to planting density. mortality of Chinese fir plantations:Long-term spacing trials in southern China. For.
Scand. J. For. Res. 32 (8), 679–687. Ecol. Manage. 465, 118103.
Wang, C.-S., Tang, C., Hein, S., Guo, J.-J., Zhao, Z.-G., Zeng, J., 2018. Branch Zhang, J., Zhang, J., Mattson, K., Finley, K., 2020a. Effect of silviculture on carbon pools
development of five-year-old Betula alnoides plantations in response to planting during development of a ponderosa pine plantation. Forests 11, 997.
density. Forests 9, 42. Zhao, D., Bullock, B.P., Montes, C.R., Wang, M., Greene, D., Sutter, L., 2019. Loblolly
Wang, C.-S., Guo, J.-J., Hein, S., Wang, H., Zhao, Z.-G., Zeng, J., 2019. Foliar morphology pine outperforms slash pine in the southeastern United States – A long-term
and spatial distribution in five-year-old plantations of Betula alnoides. For. Ecol. experimental comparison study. For. Ecol. Manage. 450, 117532.
Manage. 432, 514–521. Zhao, D., Bullock, B.P., Montes, C.R., Wang, M., Westfall, J., Coulston, J.W., 2020. Long-
Wang, Z., Yu, N., Li, R., Zou, W., Yang, J., Yin, G., Peng, T., Wang, Z., Yu, N., Li, R., term dynamics of loblolly pine crown structure and aboveground net primary
Zou, W., Yang, J., Yin, G., Peng, T., 2020. Branch development of eight-year-old production as affected by site quality, planting density and cultural intensity. For.
Mytilaria laosensis plantations in response to planting density. Austrian J. For. Sci. 4, Ecol. Manage. 472, 118259.
247–266. Zhao, D., Kane, M., 2012. Differences in growth dynamics of loblolly and slash pine
Węgiel, A., Polowy, K., 2020. Aboveground carbon content and storage in mature Scots plantations in the southeastern United States. For. Ecol. Manage. 281, 84–92.
pine stands of different densities. Forests 11 (2), 240. Zhao, D., Kane, M., Borders, B.E., 2011. Growth responses to planting density and
Weiner, J., Freckleton, R.P., 2010. Constant final yield. Annu. Rev. Ecol. Evol. Syst. 41, management intensity in loblolly pine plantations in the southeastern USA Lower
173–192. Coastal Plain. Ann. For. Sci. 68, 625–635.
Wells, E.D., 1994. Effects of planting spacing and refertilization on growth and nutrition Zhao, D., Kane, M., Borders, B., Subedi, S., Akers, M., 2012. Effects of cultural intensity
of black spruce (Picea mariana (Mill.) B.S.P.) planted on a minerotrophic peatland in and planting density on stand-level aboveground biomass production and allocation
Newfoundland. Canada. Can. J. For. Res. 24, 1302–1311. for 12-year-old loblolly pine plantations in the upper coastal plain and piedmont of
West, P.W., Smith, R.G.B., 2019. Inter-tree competitive processes during early growth of the southeastern United States. Canadian Journal of Forest Research 42 (1),
an experimental plantation of Eucalyptus pilularis in sub-tropical Australia. For. Ecol. 111–122.
Manage. 451, 117450. Zheng, Y., Zhang, Y., Wu, J., 2016. Yield and quality of Moringa oleifera under different
West, P.W., Smith, R.G.B., 2020. Effects of tree spacing on branch-size development planting densities and cutting heights in southwest China. Ind. Crops Prod. 91,
during early growth of an experimental plantation of Eucalyptus pilularis in 88–96.
subtropical Australia. Aust. For. 83, 39–45. 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.
16