Forest Ecology and Management 462 (2020) 117953

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Forest Ecology and Management

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

Variation in whole-rotation yield among Eucalyptus genotypes in response to T

water and heat stresses: The TECHS project
⁎
Dan Binkleya, , Otavio C. Campoeb,c, Clayton Alcarde Alvaresc, Rafaela Lorenzato Carneirod,
Jose L. Stapec,d
a
School of Forestry, Northern Arizona University, Flagstaff, AZ 86011, USA
b
Department of Forest Sciences, University of Lavras (UFLA), Lavras, MG 37200, Brazil
c
Department of Forest Science, São Paulo State University – UNESP, 18600 Botucatu, SP, Brazil
d
Forestry Science and Research Institute (IPEF), Piracicaba, SP 13400, Brazil

A B S T R A C T

The TECHS project spanned a 3500 km gradient from the Amazon to Uruguay, examining the influence of stresses from temperature and water supply on clonal
plantations of Eucalyptus, with and without rain reduction, and across a stocking gradient. The whole-rotation mean annual increment (MAI) showed a humped
pattern in relation to temperature, rising from about 18 Mg ha−1 yr−1 of stemwood production when mean annual temperatures were near 16 °C, to 27 Mg ha−1 yr−1
at 20 °C, and then falling to less than 15 Mg ha−1 yr−1 above 24 °C. The age trend in growth showed a steeper initial rise in the warmer tropical sites (reaching a peak
current annual increment, CAI, of 27 Mg ha−1 yr−1, at age 2–3 years), but the slower early growth in the cooler subtropical sites had a higher peak (CAI of
32 Mg ha−1 yr−1, at 4 years) and slower decline, giving 15% higher MAI for the cooler region. Whole-rotation MAI declined by about 2.2 Mg ha−1 yr−1 for each 1 °C
increase in temperature (in the range between 19.5 and 23.5 °C), and MAI declined by 0.5 Mg ha−1 yr−1 for each 100 mm yr−1 decline in rain. The effect of reducing
ambient rain was also a loss of 0.5 Mg ha−1 yr−1 for each 100 mm yr−1 reduction in rain, though the effect was small on low productivity sites (< 0.1 Mg ha−1 yr−1
for sites with MAI of 10 Mg ha−1 yr−1), and large on high productivity sites (1.4 Mg ha−1 yr−1 for sites with MAI of 40 Mg ha−1 yr−1). In the stocking portion of the
project, growth of individual trees decreased (and stand-level growth increased) with increases in stocking, and water deficits led to decline in both measures of
growth. Under favorable environments for Eucalyptus, stem growth in intensively managed plantations is about five-times the rates reported for non-plantation
forests. The higher growth in plantations declines under warmer and drier conditions, matching productivity of non-plantation forests below about 900 mm yr−1
rainfall and 26 °C annual average temperature. The potential productivity of forests depends more strongly on management systems (genetic selection, site pre-
paration, fertilization, spacing, competition control and protection) than on environmental gradients.

1. Introduction 2019; Elli et al., 2019).

Rates of forest growth change through time and across space, and
these patterns have been a core focus since the beginning of forestry
and silviculture (Puettmann et al., 2008). Explanations based on pro-
duction ecology became more common beginning in the 1980s
(Waring, 1983; Linder, 1987; Cannell, 1989), leading to experiments
that varied the water and nutrient supplies to test responses to changing
resource supplies and efficiencies of resource use (for example, Pereira
et al., 1989; Gower et al., 1992; Albaugh et al., 2004). Most studies in
production ecology have been designed for individual sites, though
comparisons among studies have shown some interesting patterns
(Binkley, 2011; Waring et al., 2016; Albaugh et al., 2016). High rates of
production and profitability spurred intensive examination of the pro-
duction ecology of Eucalyptus plantations in Brazil, including compar-
isons across geographic gradients and the application of production
ecology models (Almeida et al., 2004; Stape et al., 2004a; Attia et al.,
The allocation of land to forest plantations depends on an array of
economic, market, and social factors (Gonçalves et al., 2020). Costs of
land may be lower in areas that have lower value for competing agri-
cultural land uses, but only if tree survival and growth are acceptably
high. The interactions of stresses from temperatures, water supplies,
and pests and pathogens differ substantially among Eucalyptus geno-
types (clones). The TECHS project (Binkley et al., 2017) was designed
as a follow up to the Brazil Eucalyptus Potential Productivity (BEPP)
Project which examined whole-rotation carbon budgets in relation to
nutrition and water supply (Stape et al., 2010). Within-site irrigation is
challenging and expensive, so the TECHS project examined the influ-
ence of water supply in two ways: a geographic gradient of more than
3500 km, and within-site rain reduction treatments. In this paper we
used the TECHS Project, at the end of a full Eucalyptus rotation (about
6 years), to examine three questions:

⁎
Corresponding author.
E-mail address: dan.binkley@alumni.ubc.ca (D. Binkley).

https://doi.org/10.1016/j.foreco.2020.117953
Received 4 December 2019; Received in revised form 29 January 2020; Accepted 30 January 2020
Available online 28 February 2020
0378-1127/ © 2020 Elsevier B.V. All rights reserved.
D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

Fig. 1. The TECHS sites were distributed across a 3500-km gradient from the Amazon to Uruguay (subtropical sites are underlined), with ranges in rainfall of 600 to
1800 mm yr−1, temperature of 16 to 28 °C, and water deficit of 0 to 700 mm yr−1. The subset of 22 sites with full-rotation data available for this paper are indicated
on the map and graph, and an additional site (#13) was used only in the stocking analysis.

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D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

Fig. 2. Aerial view of Site 26, 33 months after planting (upper). The rain reduction subplots used troughs to capture about 30% of incoming rain and move the water
out of the plantation (Site 31, 20 months).

• How does tree survival and whole-rotation mean annual increment The mean annual temperatures of the sites ranged from 17 to 28 °C,
(MAI) relate to temperature and water supplies across sites? with average annual precipitation (during the years of the project) from
• How sensitive is survival and production to within-site changes in 620 to 1650 mm yr−1 (Table 1). Accounting for rainfall, temperature,
water supply and stocking? humidity and soil water storage, the annual water deficit
• How do these responses differ among genotypes (clones)? (Thornthwaite-Mather water balance in control plots) ranged from 0 to
700 mm yr−1.
Tree diameters were measured every 6 months at 1.3 m height, and
2. Project description and methods
stemwood biomass was calculated based on clone-specific allometric
equations developed using destructive sampling data from 22 sites at
The TECHS project began in 2011, with an experimental design
approximately 6 years after planting. Stemwood biomass equations
applied at 36 sites from northern Brazil to Uruguay (Fig. 1, for details
were fitted using Schumacher-Hall model with at least 12 trees per
see Binkley et al., 2017). The project used 18 clones, with 7 oper-
clone. Coefficients of determination ranged from 0.94 to 0.99, and all
ationally used clones for the tropical sites (B2, D4, E5, G7, H8, P7, R9),
equations had P values < 0.001 (Table 2). The clones differed sub-
7 other operationally used clones for the cooler subtropical sites (F6, I9,
stantially in leaf area and growth rates (Fig. 3), with the best clone at
J1, L3, M4, N5, O6), and 4 “plastic” clones that were used for all sites
each site producing about twice as much wood as the worst clone,
(A1, C3, K2, Q8). Each site had one replicate of 11 clones, plus an
though the absence of within-site replication confounds true differences
additional clone chosen for the local site (not included in this analysis).
among clones with unrelated variation among plots (see Discussion). At
Trees were planted at 3 × 3 m spacing (~1110 trees ha−1) in 0.2 ha
the time of this analysis, twenty-two sites had full rotation data avail-
plots, and each plot was split to include a rainfall removal treatment
able. Mean annual increments were calculated for the oldest measure-
(about a 30% reduction in canopy throughfall beginning during the
ment period, which ranged from 61 to 77 months (the typical length of
second year after planting; Fig. 2). The influence of stocking was ex-
a rotation, except for cool sites that may extend one or two years
amined at each site with an additional plot for each clone, with dis-
longer). The MAI was nearly constant during this period, so no age-
tances between planting rows giving tree densities equivalent to 450 to
related trends in MAI would confound our analyses. The focus of the
14,000 trees ha−1. The population of inference was the geographic
project was on MAI, but we also applied the end-of-rotation biomass
range of Eucalyptus plantations in Brazil and Uruguay, so the unit of
equations to earlier measurements of diameter and height to estimate
replication was site (rather than plots within sites). The TECHS project
trends with stand age. Stem density is lower in younger trees, so the age
had a number of clones and sites where rain reduction treatments were
trends slightly overestimate growth in early years.
not installed, giving 2 plots for each clone at a site. These 83 replicate
Survival at the end of the rotation averaged 89.5% across all clones
pairs of plots provided an estimate of the size of random differences
and sites, with no significant effect of site temperature, rainfall, or
between plots with no treatment differences within sites. Among these
rainfall exclusion at the 3 × 3 m spacing. Across the environmental
pairs, the average difference in current annual increment at the end of
gradient in this study, well-selected clones (with 9 m2 tree−1) did not
the rotation was 16% (1.9 Mg ha−1 yr−1), with a standard deviation of
have a high mortality risk for the weather experienced during this
the differences between the pairs of 1.9 Mg ha−1 yr−1.

3
 D. Binkley, et al.

Table 1
Environmental characteristics of the sites tropical and subtropical (underlined) used in this paper, updated from Binkley et al. (2017) for the entire period of the rotation.
Site Latiitude Longitude Eleva-tion -Temperature °C – AGDD Cold days < 2 °C Hot days > 35 °C Relative Incident VPD (24 hr) VPD (day-light) Rain PET AET Water deficit
humidity radiation
−1
S W m Mean Min. Max. % MJ m−2 yr−1 ————kPa————— ——————————mm yr ———————

2 24.21 49.97 770 18.3 14.5 23.4 33,479 11 14 77.1 17.1 0.6 0.8 1652 841 834 7
4 19.31 42.42 243 22.7 18.0 29.3 44,799 0 223 76.2 16.1 0.8 1.1 1102 1141 856 284
5 18.58 42.93 873 20.7 16.1 27.2 40,020 0 21 74.8 17.6 0.7 1.1 952 960 771 189
7 18.02 50.90 681 22.5 16.7 30.1 44,220 1 105 70.4 17.1 1.1 1.5 1515 1129 829 300
8 11.86 38.37 218 25.4 20.3 30.7 49,319 0 169 69.2 18.5 1.1 1.5 729 1427 729 698
9 18.73 47.92 969 23.4 18.8 29.1 45,508 0 37 60.9 19.1 1.2 1.7 1390 1180 957 223
11 18.71 52.59 783 22.6 17.9 29.1 44,404 0 40 67.3 19.2 1.0 1.4 1106 1103 820 283
12 19.76 40.13 36 24.4 20.8 30.0 49,020 0 85 75.9 18.0 0.8 1.2 1010 1306 936 370
13 20.90 51.90 361 25.0 19.6 32.3 50,269 0 578 65.9 17.7 1.3 1.8 1215 1406 1150 256
14 19.96 51.59 480 24.5 18.8 31.3 48,241 0 351 67.5 18.8 1.1 1.6 1173 1335 1038 297
19 12.17 48.50 255 26.7 21.4 33.7 54,026 0 748 66.8 20.1 1.3 1.8 1229 1603 1068 534

4
20 22.35 46.97 633 21.7 15.8 29.6 42,512 0 146 72.6 17.4 0.8 1.2 1187 1055 947 108
22 24.23 50.53 888 19.1 15.3 23.8 34,928 8 0 80.2 17.0 0.5 0.7 1531 874 866 8
24 22.73 49.00 656 21.5 16.8 28.2 42,052 0 69 73.2 15.7 0.8 1.2 1308 1030 948 82
26 16.78 44.31 926 23.1 17.6 29.9 45,107 0 90 65.3 19.5 1.1 1.6 802 1154 662 493
29 3.44 43.07 81 27.5 23.0 34.1 56,580 0 863 71.4 17.3 1.2 1.7 1369 1740 1101 639
30 17.32 43.77 848 23.0 17.3 29.8 44,630 0 115 62.5 21.7 1.2 1.7 619 1152 591 561
31 16.34 39.60 200 23.6 20.3 28.5 46,589 0 23 86.4 18.4 0.4 0.7 1172 1206 994 211
6 30.19 51.62 150 18.4 14.4 23.6 33,648 21 38 79.1 16.0 0.5 0.8 1212 871 821 50
10 23.03 48.53 869 20.8 16.2 26.7 39,509 0 17 72.8 17.8 0.8 1.1 1546 973 933 40
23 27.53 50.10 870 16.7 12.5 22.6 30,247 72 0 80.1 16.2 0.4 0.7 1668 780 769 11
28 26.11 50.21 812 17.1 12.4 23.5 31,241 93 0 84.2 14.6 0.4 0.6 1273 799 759 40
33 23.85 48.70 695 20.3 14.9 25.9 36,918 1 2 77.9 16.4 0.6 0.9 1175 940 818 122

Site 33 is both tropical and subtropical. Temperatures are mean annual temperature, and the coldest and warmest month. Cold and hot days are number of days with average temperature below 2 °C or above 35 °C.
AGDD = accumulated growing degree days considering temperatures between 5 and 35 °C. Incident radiation is average global radiation, PET = potential evapotranspiration by Penman–Monteith equation;
AET = actual evapotranspiration by water balance of Thornthwaite-Mather and using a specific water holding capacity of the soils; water deficit is by water balance of Thornthwaite-Mather and using a specific water
holding capacity of each site’s soil and AET. All values are for the period of the project.
Forest Ecology and Management 462 (2020) 117953
D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

Table 2 rotation.
Biomass regression equations for stem wood mass in the form: ln(stem mass, The patterns of stemwood production across gradients in time,
kg) = b0 + b1ln(DBH, cm) + b2ln(height, m). precipitation, and temperature were evaluated with CurveExpert ver-
Clone b0 b1 b2 r2 Standard Minimum Maximum sion 2.6 (http://www.curveexpert.net/), for all clones within a site, and
error DBH (cm) DBH (cm) in some cases for whichever three clones had the highest growth in each
site. The best-fit equations were chosen based on the lowest AIC
A1 −5.1213 2.1142 1.1904 0.96 14.26 10.2 23.0
(Akaike’s Inormation Criterion) corrected for small sample sizes.
B2 −3.6947 2.3361 0.5474 0.98 11.02 12.7 24.0
C3 −4.3563 1.9002 1.1107 0.97 14.18 6.5 22.9
D4 −4.0626 1.7995 1.1588 0.96 12.23 9.2 27.9
E5 −3.9496 1.9995 0.9348 0.97 10.10 10.6 22.5 3. Results
F6 −4.8363 1.5095 1.6217 0.98 12.33 12.4 26.4
G7 −4.0187 2.0550 0.9372 0.98 12.35 9.2 23.6 The accumulated stem mass at the end of the rotation was about
H8 −3.3154 1.6001 1.0936 0.97 9.05 9.5 21.6
15% greater for subtropical sites than for tropical sites (Fig. 4). The
I9 −4.1872 1.8534 1.1414 0.98 13.30 8.4 23.2
J1 −4.8363 1.5095 1.6217 0.98 12.33 12.4 26.4 peak in current annual increment at 4 years in the subtropical sites was
K2 −4.0298 1.9330 1.0240 0.97 10.92 8.6 24.5 about 15% greater than the tropical-site peak at 2.5 years. The post-
L3 −4.2801 1.8534 1.1414 0.98 3.72 9.7 14.5 peak decline was greater for the tropical sites, leading to diverging
M4 −4.1872 1.8534 1.1414 0.98 13.30 8.4 23.2 yields beyond 4.5 years.
N5 −4.1872 1.8534 1.1414 0.98 13.30 8.4 23.2
O6 −6.0410 1.2330 2.2280 0.98 9.27 14.0 23.2
Fig. 4 has a continuous representation of age (rather than binning
P7 −4.1187 2.2249 0.8225 0.96 7.38 6.4 17.4 into “young” and “old”), but the information on stem mass and growth
Q8 −5.1083 1.6901 1.5640 0.95 12.32 8.2 23.8 are clumped into two bins (tropical and subtropical). This can be useful
R9 −3.8471 2.0752 0.8756 0.99 8.49 9.3 21.9 for showing overall differences between classes, but some information
about the responses to temperature and age is discarded in the binned
summaries. A fuller use of information would allow site temperature

Fig. 3. Examples of differences between clones along plot boundaries. Four years after planting at site 11, Clone Q8 (upper plot in upper picture, left plot in lower
picture) had higher leaf area and light interception than Clone C3. Higher light interception was associated with 80% larger mean annual increment for the rotation
for clone Q8 (see Mattos et al., 2020).

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D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

A response was clearly curvilinear at the end of the rotation, with highest
Subtropical:
MAI occurring at an average annual temperature of 19 to 20 °C (Fig. 6).
Stem mass = 261.8/(1+(age/6.07)^-2.46), r2 = 0.64
Stem mass (Mg ha-1)

At this optimal temperature, the three most productive clones had

Tropical: about 40% greater MAI than the average of all 11 clones. The decline in
Stem mass = 145.9/(1+(age/3.85)^-2.40), r2 = 0.52
current increment at mid-rotation was about 2.9 Mg ha−1 yr−1 for each
1 °C increase in average annual temperature (Binkley et al., 2017). The
response was similar for the average of all clones for the whole rotation,
with a drop of about 2.2 Mg ha−1 yr−1 for each degree of warming in
the range of 19.5 to 23.5 °C for all clones. The temperature response
appeared even stronger for the best three clones at each site, with MAI
dropping by about 3.5 Mg ha−1 yr−1 for each degree of warming be-
tween 19.5 and 23.5 °C. This “best-three clones” pattern may be more
Stand age (years)
B accurately described as the “best-three plots,” because the high growth
30 in these plots resulted from either (or both) genotypes or simply among-
Stem increment (Mg ha-1 yr-1)

plot variation in growth (see Discussion).

25
20
15
10
5 increment (Binkley et al., 2017).
0
0 1 2 3
Age (years)
Fig. 4. The whole-rotation yields were lower for warmer tropical sites than for
cooler subtropical sites (A, shaded regions are 95% confidence bands for site
averages (n = 659 observations for all ages, clones and sites). The current
annual increment peaked at age 2 to 3 for the tropical sites, and year 4 for the
subtropical (lower graph).
also to be a continuous variable in the relationships. The rapid decline
in current growth rate at higher temperatures is apparent in Fig. 5A.
The physiology underlying the responses in growth might relate more
to the accumulation of stem mass over time rather than age, and indeed
stem mass provided a much stronger pattern than age (Fig. 5B).
Subtropical sites Mean annual increment decreased with decreasing rainfall, though
the strength of the pattern was weaker than the response to tempera-
ture. A 100 mm yr−1 decrease in rainfall across sites was associated
with a 0.5 Mg ha−1 yr−1 decrease in MAI (Fig. 6). This response was
more moderate than at mid-rotation, when a 100 mm yr−1 decrease in
Tropical sites rainfall was associated with a 1.5 Mg ha−1 yr−1 decrease in current
The effects of temperature and rainfall can be combined into a
single variable of a water deficit, based on rainfall, potential evapo-
4 5 6 7
transpiration, and soil water storage capacity. The decline in growth in
relation to increasing water deficit was more precise than the effect of
rainfall alone, but the water deficit did not account for more of the
variation in MAI among sites than temperature alone. The effects of
temperature and water stresses can also be examined independently
rather than constrain the shape of the relationship by the calculation of
a water deficit. The effect of rainfall on MAI differed substantially
among sites in relation to site differences in temperature (Fig. 7). The
simple relationship between temperature and MAI had a lower AIC
(about 3 units lower) than the combined relationship with temperature
and rainfall on MAI, indicating that the simple temperature relationship
matched the evidence of MAI across sites better than the combined
information for temperature and rainfall.

3.1. MAI responses to geographic gradients in temperature and rainfall

At mid-rotation, the TECHS plantations showed a large growth re-
duction across all sites with increasing temperature (Binkley et al.,
2017). The subset of the coolest sites may have shown a different trend,
with growth increases with increasing temperature, but at mid-rotation
a curved response of growth versus temperature was not statistically
more consistent with the data than a simple linear response. The
3.2. MAI responses to stocking
Stand-level growth increased with increasing stocking (trees ha−1),
as the growth rates of individual trees declined (Fig. 8). The influence of
water deficits was similar in magnitude to the effect of stocking, and the
combination of both variables gave the best prediction of growth rates.
We expected that mortality would increase at high stocking and high

A B

r2 = 0.19 r2 = 0.33

Fig. 5. Using site temperatures to examine growth with age (A, n = 3308 plot-level observations of all ages, clones, and sites) makes fuller use of the information in
the experiment than binning the sites into two temperature classes as in Fig. 4. The physiology underlying the age trend may relate more to stem mass than to age per
se, and substituting stem mass for age provided a much stronger fit to the data (B).

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D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

A 80
Best 3 clones per site

Mean annual increment

Mean annual increment
MAI = 1/0.534 – (0.0518 x temp) +
(0.00134 x temp2) r2 = 0.64 60

(m3 ha-1 yr-1)

(Mg ha-1 yr-1)
40

Average of all clones

20
MAI = 1/0.751 – (0.0724 x temp) +
(0.00184 x temp2) r2 = 0.60
0

Mean annual temperature (oC)

B
Best 3 clones per site
Mean annual increment

MAI = 2.63 + 0.017 x rain, r2 = 0.31

(Mg ha-1 yr-1)

Average of all clones

MAI = 2.94 + 0.012 x rain, r2 = 0.25

Mean annual precipitation (mm)

C

Best 3 clones
Mean annual increment

MAI = 30.8 - 0.028 x deficit, r2 = 0.66

(Mg ha-1 yr-1)

All clones
MAI = 23.8 - 0.022 x deficit, r2 = 0.63

Water deficit (mm yr-1)

Fig. 6. Mean annual increment showed a curvilinear trend with temperature across the 22 sites (A), with maximum growth occurring at about 19 to 20 °C (shaded
regions are 95% confidence bands). The best-3 clones at each site grew about 30% faster than the average of all clones (see Discussion). Growth increased with
increasing rainfall (B), similar to the black line representing the pattern from over 33,000 inventory sites of Suzano S.A. A strong decline in growth was also evident
in relation to water deficit (C), though the relationship was not as strong as the effect of temperature (AIC for water deficit was about 2 higher than for temperature).

water deficits; the average tend fit this pattern but it accounted for little
of the observed variation and was not close to being significant
(P = 0.35).
3.3. Production responses to within-site reductions in water supply
The reduction in rain lowered average CAI in the final year of the
rotation by 2.0 Mg ha−1 yr−1, with greater reductions on more pro-
ductive sites (Fig. 9). Sites with MAI of 10 Mg ha−1 yr−1 lost only
0.8 Mg ha−1 yr−1 (6%) with reduced rain, whereas a site with MAI of
40 lost an average of 5.5 Mg ha−1 yr−1 (18%). The rain reduction effect
did not relate to patterns of rain or temperature across sites (P = 0.4).
The removal of 30% of rain would be an average reduction of about
340 mm yr−1, so a 100 mm yr−1 reduction in rain gave an average
reduction in CAI of 0.6 Mg ha−1 yr−1. The loss of CAI in response to
rain reduction increased with increasing site productivity,
from < 0.1 Mg ha−1 yr−1 on sites with MAI of 10 Mg ha−1 yr−1 to
1.6 Mg ha−1 yr−1 for sites with MAI of 40 Mg ha−1 yr−1. The addition

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D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

All clones 3 best clones

Fig. 7. The response surface of MAI in relation to temperature and rainfall across the 22 sites showed a clear curvilinear effect of temperature, and how the effect of
rainfall varied with temperature (r2 = 0.71 for all clones, 0.72 for best three clones, P < 0.0001).

of site rainfall as a second independent variable improved the predic-
tion of the loss of CAI in response to rain reduction. Once the CAI (with
full rain) was accounted for, a change of 100 mm yr−1 in ambient rain
changed CAI by 0.6 Mg ha−1 yr−1.
3.4. Responses among genotypes
The plastic clones (those chosen to include across the full
geographic gradient) showed the greatest MAI at sites with mean an-
nual temperatures between about 19 and 20.5 °C (Fig. 10). With the
exception of one clone, all tropical clones showed declining MAI with
increasing temperature. Clone E5 had a slight hump in temperature
response, with maximum MAI occurring at 20.5 °C. The plastic clones
had been chosen for expected strong performance across the broad
range of sites, and indeed they grew almost as well as the average of the
tropical clones. The importance of genotype by environment

A. Individual B. Stand
tree growth growth

C. Tree
survival

Fig. 8. The growth of individual trees (A) declined with increasing stocking and water deficit (r2 = 0.41, P < 0.01). Stand growth increased with increasing
stocking and decreasing water deficit (r2 = 0.40, P < 0.01). Survival at the end of the rotation appeared to decline with increasing stocking and water deficits (C),
but the effect was not significant (r2 = 0.10, P = 0.35).

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D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

CAI reduction = -.8 + 0.158 * MAI, r2 = 0.08

reduction (Mg ha-1 yr-1)

CAI reduction with rain

MAI with full rain (Mg ha-1 yr-1)

Fig. 9. Current annual increment (CAI) declined by an average of 2.0 Mg ha−1 yr−1 (12%) with a 30% reduction in rainfall (left, larger numbers mean larger
reduction in growth (117 dots are each clone in each site, P < 0.001, shaded region is the 95% confidence band for the average trend). The inclusion of water deficit
along with MAI improved the prediction of growth reduction (large values are large reductions, r2 = 0.16, AIC reduced by 8 with inclusion of water deficit).

40 points in the rankings (Fig. 11). An early selection of the best-per-

A1
forming clones would not have matched a selection made at the end of
Mean annual increment

C3 the rotation. The ranking for site productivity was somewhat more
30
K2
(Mg ha-1 yr-1)

robust; poor sites remained in the lower rankings, but rankings shuffled
Q8 for the more productive sites.
20 B2
D4
E5 4. Discussion
10
G7
H8 The effect of genotypes was examined in two ways: Figs. 6 and 7
0 showed the average growth responses in comparison with growth of
P7
16 18 20 22 24 26 28
R9 whichever three clones showed the greatest growth in each site, and
Mean annual temperature (oC) Fig. 10 traced the responses of each clone across sites. The patterns in
40 Figs. 6 and 7 are not attributable to genotype alone, however, because
Mean annual increment

the three plots with the highest growth (identified at the end of the
30 rotation) may have resulted from better-adapted genotypes or from
(Mg ha-1 yr-1)

random variation among plots (see discussion of null expectations for

20
Plastic clones
10
0
16 18 20
Mean annual temperature (oC)
Fig. 10. All clones except P7 showed strong responses to temperature (upper);
P7 performed well only on the warmest sites. The average MAI for plastic and
tropical genotypes showed similar responses to temperature (lower). The
ranking of the best-performing clones varied with site temperature.
interactions was evident in the comparison of average clone perfor-
mance versus the average of the whichever three clones grew best at
each site (Fig. 7). The apparent size of the genotype effect may be
overestimated in Fig. 7, however, because the lack of replication within
sites means that post-facto plots with higher production rates included
any true genotype effects plus random variations in site conditions
(Binkley, 2018). A “clone” could be among the best either for genetic
reasons or because the microsite for a particular plot is unusually
productive.
The subtropical clones showed no significant relationship between
MAI and temperature. The lack of a clear trend may relate to the small
number of sites (5) that reached full rotation at the time of this analysis,
along with the relatively narrow range of site temperatures sites (mean
temperatures between 16 and 22 °C).
The relative performance of clones changed substantially from mid-
rotation to the end of the rotation, with many clones shifting several
within-site interpretations of unreplicated studies, Binkley, 2018). The
Tropical clones
TECHS project was not designed to separate clonal (genotype) effects
within sites from random variation among plots. As noted in the
Methods, adjacent plots of the same clone (with no rain exclusion
treatment) differed by an average of about 16% in MAI (1.9 Mg ha−1
yr−1, with a standard deviation 1.9 Mg ha−1 yr−1). The MAI of the
22 24 26 28
three best clones within each site averaged 30% (5.3 Mg ha−1 yr−1,
standard deviation 2.3 Mg ha−1 yr−1) greater than the average of all 11
clones, double the null expectation of 1.9 Mg ha−1 yr−1 based on
variance among replicate plots of the same genotypes (P < 0.0001).
We can be confident that the identification of the best 3 clones in each
site was largely a genotype difference rather a microsite-variation
among plots within sites. This issue would not apply to Fig. 9, because
random within-site effects were incorporated in the average trend ob-
served for each clone across sites.
Eucalyptus stem production declined by about 10 to 15% for a 1 °C
increase in mean annual temperature (in the range of 19.5 to 23.5 °C),
which is similar to a 15% increase in leaf respiration per degree found
for similar clones and sites in Brazil (J.L. Stape, unpublished data). We
suspect that the decline in stem production at high temperatures re-
sulted from an increase in whole-plant respiration that exceeded any
increase in photosynthesis (Drake et al., 2019a, 2019b; Ryan and Asao,
2019). If whole-plant respiration increased with increasing tree size,
this might also account for the larger decline in growth with age for
faster-growing tropical sites (Figs. 4 and 5). Unfortunately, funding was
not available to test this expectation.
The differences in growth among genotypes (and plots) within sites
can be examined with a production ecology approach. A comparison of
all 18 clones at Site 33 found that the mid-rotation leaf area index

9
D. Binkley, et al. Forest Ecology and Management 462 (2020) 117953

Clones Fig. 11. The ranking of clone performance across

sites at mid rotation differed substantially from
L3 P7
10 Subtropical Q8 10 Tropical C3
rankings at the end of the rotation (1 = highest
growth rate), for clones planted in both subtropical
and plastic M4 and plastic K2 (upper left) and tropical sites (upper right), in part

End rotation rank

8
End rotation rank

8 F6 Q8 because of changes in pest and pathogen problems

I9 R9 over time within some sites. The MAI for the whole
6 J1 6 E5 rotation was lower than the current annual incre-
N5 G7 ment at mid-rotation (lower left), and the ranking
4 C3 4 H8 of sites was also very inconsistent (lower right).
K2 D4
2 O6 2 B2
A1 A1
0 0
0 2 4 6 8 10 0 2 4 6 8 10
Sites Mid-rotation rank Mid-rotation rank
31
30 10 4
End rotation MAI (Mg ha-1 yr-1)

22
11 2
End rotation rank

8 7
24 5
20 20 9 75 6 20
4 24
12 4 11
10 2
8 2 22
33
0 0
0 10 20 30 40 50 60 0 2 4 6 8 10
Mid-rotation CAI (Mg ha-1 yr-1) Mid-rotation rank

ranged from about 3 to 6 among clones, and light interception by the of the responses of production to temperature have typically shown that
canopies ranged only from about 2.0 to 2.7 GJ m−2 yr−1 (Mattos et al., productivity increases with increasing temperatures, with the highest
2020). The clones differed by as much as 2-fold in the amount of wood rates occurring in the warmest locations. We plotted the response of all
produced per unit of light intercepted, so clonal differences in this index clones to temperature and ambient rain (Fig. 7) with the pattern for
of light use efficiency were more important than differences in light non-plantation forests in the database compiled by Taylor et al. (2017).
interception. An investigation of carbon (C) fluxes at four TECHS sites The effect of plantation versus non-plantation on stem growth was large
showed strong differences in the partitioning of C to stems versus be- (2-fold or more) under optimal environmental conditions for Eucalyptus
lowground: the more productive clones and sites allocated substantially plantations (Fig. 12). The productivities of more marginal sites were
greater proportions of their total C to stem production (Campoe, 2020). similar for plantation and non-plantation forests. Intensive silviculture
The response of Eucalyptus productivity to environmental gradients applied to highly selected genotypes of Eucalyptus increase stemwood
across sites might confound important driving factors with other cov- production by more than 12 Mg ha−1 yr−1 (a 5-fold increase) in the
arying factors. At mid-rotation the geographic pattern of production best locations. The Eucalyptus plantations were also more responsive
and rain indicated that a 100 mm yr−1 increase in rain was associated along environmental gradients, showing 2-fold changes in productivity
with a 1.5 Mg ha−1 yr−1 increase in CAI (Binkley et al., 2017). The over domains where non-plantation forests tended to show changes of
strong influence of temperature confounded the simple relationship about 50%. The striking decline in productivity of Eucalyptus planta-
between growth and rain, and the apparent benefit of 100 mm yr−1 of tions with temperatures above 22 °C revealed a limit of temperature
rain dropped to 0.4 Mg ha−1 yr−1 when the effect of temperature was adaptability that was much less pronounced in the non-plantation for-
also included. The rain reduction treatments allowed for the direct ef- ests. The broader “optimum” temperature for the non-plantation forests
fect of rain to be separated from interactions with temperature. The may result from the shifting genetics across the geographic gradients, as
end-of-rotation CAI increased by an average of 0.5 Mg ha−1 yr−1 for a the diverse array of species changes.
100 mm yr−1 increase in rain (Fig. 9), which is similar to the mid- Although the patterns of temperature and rainfall effect on pro-
rotation value of 0.4 Mg ha−1 yr−1. The CAI declined from 16 Mg ha−1 ductivity were very highly significant, these factors still left most of the
yr−1 at mid-rotation to 11 Mg ha−1 yr−1 in the final year of the ro- variance across sites and clones unexplained. This is not surprising, as
tation. An overall conclusion from the TECHS project is that the within- the influence of temperature and rain on production are also influenced
site sensitivity to rain was similar to the across-site response, as long as at finer scales than annual averages, and by factors such as vapor
the strong effect of temperature across sites was also accounted for. pressure deficit, soil water holding capacity, and silvicultural details.
We note that changes in growth and in rankings for sites and clones The general conclusions presented in this paper are both robust and
resulted from both the normal patterns expected in relation to stand fuzzy. More precise insights can be developed from more detailed in-
age, and to variable effects of insects and diseases. The impacts of in- vestigations, such as the parameterization of ecophysiological growth
sects and diseases were highly variable among clones, and problems models. Models such as 3PG and G’day typically yield correlations be-
became more pronounced later in the rotation. tween predicted and observed mean annual increments with r2 of 0.8 or
The broader implications of the growth patterns in the TECHS higher (Almeida et al., 2004, 2010; Stape et al., 2004b; Attia et al.,
Project relate to issues of scale, and how insights may or may not be 2019; Elli et al., 2019; Caldeira et al., 2020). Intensive experiments
suitable for extension to other situations. Regional and global analyses across large geographic gradients (and diverse genotypes) provide both

10
D. Binkley, et al.
TECHS all clones
Non-plantation
forests
Fig. 12. The TECHS clones responded to geographic gradients in temperature and ambient rain (left, upper blue surface from Fig. 7) more strongly than non-
plantation forests (left, lower red surface, Taylor et al., 2017, updated 2017–03-27, n = 143, P < 0.0001, doi:https://doi.org/10.5063/F19021QT). The greater
productivity of Eucalyptus plantations dropped to near 0 (or below) for very dry sites and for very hot sites. The best-performing clones at each site would have a
larger difference than the trend plotted here.
insights into broad averages, and opportunities to parametrize and
validate models that can provide detailed site-specific insights.
Declaration of Competing Interest
The work has not been published elsewhere, and is not under con-
sideration by any other journal. We declare no personal or financial
interests that could conflict and bias the science of our paper.
Acknowledgements
This overview of forest production was possible because of the work
of over 200 people involved in TECHS. The contributions of Ítalo R.
Cegatta, Luiz Erivelto de Oliveira Júnior and Renata Abranches
Junqueira were fundamental for the TECHS database. The project was
possible only through the coordination provided by Forestry Science
and Research Institute (IPEF, ipef.br/techs/en, Directors Luiz Ernesto
George Barrichelo and José Otávio Brito). TECHS was funded by the
following 26 companies: Anglo American (Andre Machado), Arauco
(Rodrigo Coutinho), Arborgen (Gabriela Bassa), ArcelorMittal
(Roosevelt Almado), Cenibra (Fernando Leite), CMPC (Elias
Araujo),Comigo (Ubirajara Oliveira), Copener (Jacyr Alves), Duratex
(Raul Chaves), Eldorado (Vinicius Silva), Fazenda Campo Bom
(Jacqueline Pirez), Fibria (Rodolfo Loos), Florestal Itaquari (Admir
Mora), Forestal Oriental (Ricardo Methol), Gerdau (Francisco Gomes),
GMR (Paulo Leite), International Paper (Cristiane Lemos), Jari (Katia
Silva), Klabin (James Stahl), Lwarcel (Marcela Capoani), Montes del
Plata (Alejandro Gonzalez), Plantar (David Fernandes), Rigesa (Ricardo
Paim), Suzano (Luiz Fabiano and Leandro de Siqueira), Vallourec
(Helder Andrade, in memoriam) and Veracel (Helton Lourenço).
Several universities and institutes also supported TECHS: University of
Sao Paulo, Sao Paulo State University, Federal University of Lavras,
Federal University of Rio Grande do Norte, Colorado State University,
North Carolina State University, and the USDA Forest Service.
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