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Susceptibility of Corymbia species and hybrids to arthropod herbivory

inAustralian subtropical hardwood plantations

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DOI: 10.2989/20702620.2010.547247

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 Southern Forests 2010, 72(3/4): 147–152 Copyright © NISC (Pty) Ltd
Printed in South Africa — All rights reserved SOUTHERN FORESTS
ISSN 2070–2620 EISSN 2070–2639
doi: 10.2989/20702620.2010.547247

Susceptibility of Corymbia species and hybrids to arthropod herbivory in

Australian subtropical hardwood plantations

HF Nahrung1*, R Waugh1, DJ Lee2,3 and SA Lawson1

Agri-Science Queensland, Department of Employment, Economic Development and Innovation, Ecosciences Precinct,
1

41 Boggo Rd, Dutton Park, Queensland 4102, Australia

2Agri-Science Queensland, Department of Employment, Economic Development and Innovation, Locked Bag 16,

Fraser Road, Gympie, Queensland 4570, Australia

3 Faculty of Science, Health and Education, University of the Sunshine Coast, Maroochydore DC, Queensland 4558, Australia

* Corresponding author, e-mail: helen.nahrung@deedi.qld.gov.au

Corymbia hybrids are becoming significant plantation varieties in subtropical and tropical Australian plantation forestry.
Although primarily developed for disease resistance and amenability to clonal propagation, they have also proven to have
good growth rates and site plasticity. Here we examined the susceptibility of pure Corymbia species and hybrids to pest
attack. Three trial sites containing C. citriodora subsp. variegata, C. torelliana, and the hybrids C. torelliana × C. citriodora
subsp. variegata, C. torelliana × C. citriodora subsp. citriodora and C. torelliana × C. henryi were assessed for pest identity,
incidence and severity. Pests caused about three-quarters of the visible crown damage to trees in these trials. At the site that
had the most arthropod damage, hybrid trees had higher damage scores and higher growth scores (height, diameter at breast
height over bark, and volume) than pure species. Site was more important than taxon in explaining damage scores, and taxa
performed differently for most traits between sites. Tree growth was negatively correlated with general crown damage, while
arthropod damage alone showed no significant relationship with growth. Our results highlight the importance of establishing
taxa trials across a range of sites when selecting for pest resistance.

Keywords: Corymbia citriodora, Corymbia torelliana, Corymbia variegata, Corymbia henryi, eucalypt, hardwood, heritability, pest

Introduction

Eucalypt (Eucalyptus and Corymbia) plantations are
a relatively new but rapidly expanding sector of the
hardwood forest industry in subtropical Australia. Almost
106 000ha of hardwood plantations are established in
Queensland and New South Wales (NSW) (Parsons and
Gavran 2007), many of which suffer significant productivity
losses from pests and diseases (Wylie and Peters 1993,
Nahrung 2006, Carnegie et al. 2008). With hardwood
plantations in Queensland and NSW being established at
a rate of over 12 000 ha y−1, it is essential that manage-
ment strategies are adopted to minimise rotation length
and maximise production to ensure the long-term viability
of the industry.
Development of interspecific eucalypt hybrids for forest
plantations is a silvicultural strategy adopted in many
eucalypt-growing regions to maximise tree performance
by combining the desirable traits of different species (de
Assis 2000). Traits for improvement through hybridisation
include growth rate, coppicing and propagation ability, pulp
yield, wood density and resistance to frost, drought, salinity
(Dale and Dieters 2007) and pests and diseases (Potts and
Dungey 2004). However, hybrid eucalypts are usually more
susceptible to insect herbivores than their parental species
(Whitham et al. 1994, Dungey and Potts 2003, Potts and
Dungey 2004), and they tend to support more insect
herbivores than pure species in native eucalypt forests
(Morrow et al. 1994, Potts and Dungey 2004). Herbivores
can impact heavily on seedling establishment, tree growth
rates, tree form and wood quality, and their management is
critical to successful plantation productivity.
The spotted gums, C. citriodora subsp. variegata
(F.Muell.) K.D.Hill and L.A.S.Johnson (CCV), C. citriodora
subsp. citriodora (Hook.) K.D.Hill and L.A.S.Johnson (CCC),
and C. henryi (S.T.Blake) K.D.Hill and L.A.S.Johnson
(CH) are molecularly homogeneous (Ochieng et al. 2008;
Shepherd et al. 2008; Ochieng et al. 2010), and their hybrids
with C. torelliana (F.Muell.) K.D.Hill and L.A.S.Johnson
(CT) (cadaga) are among the preferred plantation species
in subtropical Queensland and NSW (Lee 2007, Lee et al.
2009, 2010).
Here, we assessed arthropod herbivory on two Corymbia
pure species, CCV and CT, and three hybrids, CT × CCC,
CT × CCV and CT × CH, determining the incidence and
severity of arthropod damage on each taxon, and examined
possible relationships with tree growth. Pests commonly
associated with these Corymbia were recorded. We further
sought to test the notion that hybrid eucalypts are more
susceptible to pests than pure species, and to ascertain
whether there may be potential for including selection for
pest resistance into advanced hybrid breeding programs.
148 Nahrung, Waugh, Lee and Lawson

Methods Table 1: Site details for the taxa trials monitored for pest incidence
and severity. Temperature and rainfall data refer to the 12-month
Existing hybrid progeny trials designed for the selection period prior to sampling; data are the mean ± SE and values in
of superior family lines based on growth rate, tree form, parentheses are the temperature range or total rainfall, respectively
height and diameter at breast height over bark (DBHOB)
and to allow estimation of genetic parameters (Lee et al. Year planted
2009) were used to assess pest susceptibility of pure and Location Temperature Daily rainfall
Site (months at
hybrid Corymbia taxa. Pure species (CCV and CT) were (nearest town) (°C) (mm)
sample)
grown from undomesticated open-pollinated seed, while I 26º35′41.9″ S, 2004 (46) 18.3 ± 0.3 1.8 ± 0.3
the hybrids (CT × CCV, CT × CCC and CT × CH) were the 151º54′53.9″ E (5.5–29) (673)
product of a controlled-cross-pollination program initiated (Kingaroy)
in 2001 using field selects of the parental species (Lee II 25º31′55.2″ S, 2005 (34) 20.4 ± 0.3 2.0 ± 0.3
2007). Three trial sites located in Queensland (Tables 1 151º28′19.2″ E (8.0–29.3) (726)
and 2) containing CCV, CT and the hybrids CT × CCV, (Mundubbera)
CT × CCC and CT × CH were assessed for pest incidence III 26º6′3.5″ S, 2004 (46) 19.3 ± 0.3 1.8 ± 0.3
151º37′22.7″ E (6.5–30) (654)
and severity. All three sites were located on predominantly
(Proston)
red ferrosol soil. Families and seedlots were not explicitly
considered within our taxa level treatments because our aim
was simply to examine and compare species and hybrids. Table 2: Number of trees of each taxon assessed at each trial site
Trees less than 1 m tall were excluded from analyses.
Taxon Site I Site II Site III Total
General crown damage
Corymbia citriodora subsp. 123 141 9 273
At each site, individual trees were given a general crown
variegata (CCV)
damage score (GCD) based on Stone et al.’s (2003) crown
C. torelliana × C. henryi 59 77 0 136
damage index, between zero (no damage) and 100 (all (CT × CH)
foliage damaged/removed) in approximate 5% increments, C. torelliana × C. citriodora 104 62 43 209
as an overall indicator of crown health. Restricted maximum subsp. citriodora
likelihood (REML) was used to detect differences in GCD (CT × CCC)
between taxa within and between sites, with taxa as the C. torelliana × C. citriodora 315 192 53 560
fixed effect and plot as random effect in the model using subsp. variegata
GenStat 11 (VSN International, Hemel Hempstead). (CT × CCV)
C. torelliana (CT) 186 160 45 391
Arthropod damage
A separate score for pest-damage only (PDS) using the
same scale (estimated proportion of foliage damaged by For the three taxa that represented parent and hybrid
arthropods) was recorded as for GCD. The contribution of species (CT, CT × CCV and CCV), a summary table of all
pests to overall crown damage was estimated using PDS/ measured parameters and the strength of their relation-
GCD. As above, REML was used to identify differences ships was made to examine possible patterns of heritability
between sites and taxa. for these traits (based on terminology in Fritz et al. 1999).
To identify taxa that were most ‘resistant’ to pest damage, Significance levels between these three taxa were checked
the proportion of trees that sustained the lowest category using analysis of variance (ANOVA) and Fisher’s protected
of damage (<5%) was compared between taxa using least significant difference (PLSD) post hoc tests.
chi-squared pairwise comparisons. The 10 most commonly
encountered pests were recorded as being nil, minor, Results
moderate or major contributors to damage at each site.
General crown damage
Relationships with growth General crown damage levels differed significantly between
Height (m) and DBHOB (cm) for trees were measured as all three sites (REML, F2,1234 = 743, P < 0.0001). Overall,
part of the taxa performance trials reported by Lee et al. site I received the highest levels of damage (mean ± SE:
(2009) five months after GCD and PDS scores were taken at 36.4 ± 0.9), and site II the lowest (16.6 ± 0.7), with site III’s
sites I and II. Site III was not measured for growth as it was average GCD score 21.6 ± 1.9. There was a significant
not part of Lee et al’s (2009) mensuration schedule. Volume taxa–site interaction (REML, F7,1234 = 3.7, P < 0.001), but
was calculated from height and DBHOB using the standard GCD only differed between taxa at site II, where CT and CT
conical approximation: volume = basal area × height × 1/3, × CH received the least crown damage (Table 3).
where basal area = 3.14 × (1/2 × DBHOB × height / (height
– 1.6))2. Site and taxa differences in growth were identi- Arthropod damage
fied using REML with the same fixed and random effects Sites II and III received statistically similar levels of defoli-
as crown damage analyses. A correlation matrix was ation from pests (9.1 ± 0.2 and 9.7 ± 0.7, respectively),
constructed using growth measures and defoliation scores while site I had significantly higher levels of pest damage
for both crown damage types using GenStat 11. (15.1 ± 1) (REML, F2,1251 = 347, P < 0.001). Again, there was
Southern Forests 2010, 72(3/4): 147–152 149

Table 3: Average ± SE general crown damage scores (range in
parentheses) for five Corymbia taxa at each of three field sites.
Different superscript letters within a column indicate means that
are statistically different. No letters within a column indicate no
significant differences within that site
Table 4: Average ± SE arthropod damage scores (range in
parentheses) for five Corymbia taxa at each of three field sites.
Different superscript letters within columns indicate means that are
statistically different at each site

Taxon Site I Site II Site III

Taxon Site I Site II Site III CCV 10.1 ± 1.6c 12.8 ± 0.7a 13.3 ± 3.2a
CCV 31 ± 2.3 18.6 ± 1.4a 23.9 ± 8.7 (5–70) (3–50) (5–30)
(10–90) (3–80) (5–90) CT × CH 14.5 ± 2.1bc 9.3 ± 0.7b –
CT × CH 33.8 ± 2.7 13.3 ± 1.5ab – (5–70) (3–30)
(15–90) (3–70) CT × CCC 23.6 ± 4.0a 9.0 ± 0.6bc 10.5 ± 1.6a
CT × CCC 40 ± 5.1 18.0 ± 1.9a 33.3 ± 4.9 (2–90) (3–30) (3–50)
(10–95) (3–90) (5–95) CT × CCV 18.3 ± 2.2ab 8.8 ± 0.4b 7.3 ± 1.2a
CT × CCV 38.5 ± 2.8 17.7 ± 1.2a 15.4 ± 3.0 (1–90) (3–60) (3–60)
(5–95) (3–95) (3–95) CT 12.3 ± 1.7c 7.2 ± 0.4c 11.2 ± 0.9a
CT 39.0 ± 2.3 13.7 ± 1.2b 17.0 ± 1.4 (1–80) (3–30) (3–30)
(10–90) (3–95) (5–50) REML results F4,308.5 = 4.59 F4,767.9 = 19.5 F4,148 = 0.8
REML results F4,304.2 = 1.71 F4,767 = 2.97 F3,14.5 = 3.2 for taxa P = 0.001 P < 0.001 P = 0.52
for taxa P = 0.15 P = 0.02 P = 0.06

0.9 a a
a significant taxa-site interaction (F7,1234 = 145, P < 0.001), PROPORTION OF DAMAGE 0.8 a a
with different taxa receiving different levels of attack at CAUSED BY PESTS b
sites I and II (Table 4). 0.7
Across all sites and taxa, pests were responsible for 0.6
an average 75.8 ± 1% of the damage sustained by trees.
The proportion of defoliation attributed to pests (AD/GCD) 0.5
differed between sites (REML, F2,735.7 = 19.7, P < 0.001), and 0.4
taxa (F4,930.9 = 16.2, P = 0.003), with CT × CCC receiving 0.3
less overall damage, as a proportion, from arthropod attack
0.2
(Figure 1).
The proportion of trees of each taxon that received less 0.1
than 5% defoliation from arthropods differed between taxa
CCV CT × CH CT × CCC CT × CCV CT
(Figure 2), with CT being the least favoured by arthropod
pests, and CCV the most ‘susceptible’; hybrids all performed TAXON
intermediately to these parental species.
Figure 1: Mean proportion of total crown damage caused by
Pest species composition and abundance varied between
pests on each of five Corymbia taxa. Error bars represent the
sites (Table 5). Lee et al. (2009) mentioned red-shouldered SE. Different letters above bars represent statistical differences
leaf beetle and longicorn beetle resistance to hybrids, but between means
these were not encountered at the time of our surveys.
Patterns of herbivory between taxa were primarily
influenced by which pests were present at each site. For c
PROPORTION WITH AD <5%

example, the high damage levels recorded for CCV at site 0.25
II was due to the presence of mites at that site, while the
higher PDS overall at site I appeared to be predominantly
0.2 b
influenced by a high population of the leaf beetle Paropsis
atomaria; individual pest patterns are examined elsewhere
(Nahrung et al. unpublished data). 0.15 b
b

Relationships with growth 0.1

Sites I and II were the only sites for which growth data
were available. Although the parameters measured were 0.05 a
correlated with each other (see below), significance patterns
varied between taxa (Table 6), except that CT consist-
CCV CT × CH CT × CCC CT × CCV CT
ently the scored the lowest at both sites for all measures.
Trees were significantly taller at site I than site II (REML, TAXON
F1,1110.3 = 315.9, P < 0.001), with significantly larger DBHOB
Figure 2: The proportion of trees of different Corymbia taxa with
and volumes (F1,1108.7 = 35.8, P < 0.001, and F1,1112.4 = 45.9, less than 5% defoliation by arthropods (AD). Different letters
P < 0.001, respectively). There was a significant site–taxa above bars denote significant differences identified by chi-squared
interaction for all growth measures. pairwise comparisons between taxa
150 Nahrung, Waugh, Lee and Lawson

General crown damage was significantly negatively had equal highest arthropod damage scores at site I, but
correlated with all growth parameters (which were correlated equal lowest at site II. At site I, hybrid trees had significantly
with each other) (Table 7). Arthropod damage, however, higher levels of arthropod defoliation than pure species,
had no significant relationship with growth. while the pure species CCV had the highest damage at
site II, and all taxa were equivalently attacked at site III.
Comparative hybrid and parental taxa performance Hybrid eucalypts are generally more susceptible to pests
Performance of the hybrid taxon (CT × CCV) for which than their parents (Whitham et al. 1994, Dungey and Potts
both parents were represented in our field sites varied 2003, Potts and Dungey 2004), but this pattern was only
relative to performance of parental taxa (CT, CCV) at each evident at site I where pest pressure (PDS) was highest.
site (Table 8). Patterns checked using ANOVA were the So-called ‘hybrid superiority’ was suggested by growth
same as in the REML analysis above (results not shown). measures at site I, where hybrids outperformed parents for
Arthropod damage scores suggested hybrid susceptibility all growth parameters. Hybrid superiority is dependent upon
at site I, additive (intermediate) resistance at site II, and no environmental conditions (Potts and Dungey 2004) and this
difference between parents at site III (see Fritz et al. 1999). may also be reflected in the intersite variation in perfor-
At all sites, hybrid growth traits were superior or interme- mance between taxa shown here. Although general climatic
diate to parent taxa. conditions and soil types were similar, and plant taxa were
identical, intersite pest distribution was not. The inclusion
Discussion of more study sites may assist in explaining this; arthropod
distribution and movement into plantations is likely to be
Across all sites, arthropods caused about three-quarters influenced by the distribution and availability of suitable
(76%) of all visible crown damage, with the remainder attrib- hosts in surrounding habitat (Strauss 2001).
utable largely to Quambalaria shoot blight, and possibly The three hybrid taxa generally performed similarly to
nutrient imbalance and drought. Only CT × CCC differed each other for GCD and PDS, and at all sites were statisti-
from this pattern, with more of its general crown damage cally inseparable, with the exception of CT × CH at site I,
caused by non-arthropod factors. Despite this, CT × CCC which had similar PDS to the parental species. This pattern
is consistent with recent molecular studies by Ochieng
et al. (2008, 2010) and Shepherd et al. (2008), which
Table 5: The 10 most commonly encountered pest species/groups
demonstrated that CCV, CCC and C. henryi are geneti-
on Corymbia taxa at three field sites and their relative abundance:
cally indistinguishable. Despite this molecular homogeneity
nil (–), minor (+), moderate (++), major(+++)
of male parents, the three hybrid taxa performed differently
for all growth measures at each site, possibly due to the
Pest group Site I Site II Site III specific combining ability of the limited number of parents
Erinose mites + +++ +++ of the hybrids planted at each site (DJL pers. obs.). Lee et
Leaf beetles +++ + +++
al. (2009) found that taxonomic identity of the male parent
Leaf hoppers +++ +++ +++
+++ + +
(CCC, CCV or C. henryi) did not explain variation in growth
Leaf blister sawfly
Orthoptera + +++ – in Corymbia hybrid progeny trials. When hybrids derived from
Swarming scarabs ++ ++ + a broader genetic base of spotted gum species/subspecies
Flea beetles ++ + + are available this will be re-examined. However, herbivore
Weevils – + ++ susceptibility (not examined by Lee et al. 2009) may be
Skeletonising Lepidoptera + +++ + more influenced by broad taxon-related preferences (CT
Amorbus sp. – ++ – versus CCC, CCV and C. henryi) of the herbivores species.

Table 6: Growth measures (mean ± SE; range in parentheses) for Corymbia taxa at sites I and II. Different superscript letters within columns
indicate means that differ significantly. DBHOB = Diameter at breast height over bark

Site I Site II
Taxon
DBHOB (cm) Height (m) Volume (m3) DBHOB (cm) Height (m) Volume (m3)
CCV 6.9 ± 0.3c 6.7 ± 0.2c 0.02 ± 0.001c 6.4 ± 0.1a 6.4 ± 0.1a 0.014± .001ab
(1–14.6) (1.9–11.5) (0.001–0.07) (1–10.5) (1.7–9.4) (0.002–0.04)
CT × CH 10.6 ± 0.3a 8.2 ± 0.2a 0.04 ± 0.002a 7.1 ± 0.3a 5.8 ± 0.1b 0.017 ± 0.001a
(4.6–14.9) (3.1–10.9) (0.01–0.08) (1–11.7) (1.4–8.9) (0–0.05)
CT × CCC 9.6 ± 0.3b 7.7 ± 0.2ab 0.03 ± 0.002b 6.9 ± 0.1a 5.9 ± 0.1b 0.016 ± 0.001a
(2.6–13.4) (3.4–10.1) (0.002–0.007) (1–11.7) (1.1–9.10) (0–0.05)
CT × CCV 9.3 ± 0.2b 7.3 ± 0.1b 0.03 ± 0.002b 5.7 ± 0. 1b 4.9 ± 0.1c 0.013 ± 0.001b
(2.7–15.7) (2.5–11.1) (0.002–0.1) (1–12) (1.1–9.3) (0–0.05)
CT 6.8 ± 0.2c 4.9 ± 0.1d 0.015 – 0.001d 3.3 ± 0.1c 3.1 ± 0.1d 0.005 ± 0.001c
(1–12.1) (1.5–9.3) (0.001–0.05) (1–9.5) (1–7) (0–0.03)
REML results for F4,623.4 = 46.6 F4,619.9 = 67.4 F4,622.5 = 51.8 F4,779.5 = 64.3 F4,80.3 = 112.8 F4,772.2 = 46.3
taxa P < 0.001 P < 0.001 P < 0.001 P < 0.001 P < 0.001 P < 0.001
Southern Forests 2010, 72(3/4): 147–152
Alternatively, another explanation for similar GCD and PDS,
but differential growth despite their genetic similarity, is
that growth may simply be a plastic trait influenced by site
factors, while herbivore susceptibility may be more closely
linked with genetically determined plant chemistry.
Heritability patterns were further examined using only the
parental-species–hybrid set (CCV, CT and CT × CCV). Even
here, inferred heritability differed between sites for all traits
(Table 7), with all of Fritz et al.’s (1999) inheritance patterns
exhibited. However, Lee et al. (2009) found a trend towards
stronger additive control of growth traits in Corymbia hybrids
in older trials; our sites may have been too young to detect
clear patterns. Using the CT, CCV, CT × CCV parent/taxa
set, Nahrung et al. (2009) found that foliar traits including
specific leaf weight, lamina thickness, leaf glabrousness and
leaf chemistry all differed between taxa, with the hybrid often
intermediate (additive inheritance pattern) between parent
taxa. Leaf physical and chemical properties affect herbivore
feeding, and P. atomaria exhibited different preference and
performance between these taxa (Nahrung et al. 2009).
With respect to general arthropod damage in the current
study, hybrids showed decreased resistance compared with
parent species (site I), an intermediate (additive) pattern
(site II) and no difference (site III). These patterns may
relate to pest pressure (site III had the lowest PDS overall,
while site I had the highest levels of pest damage), which in
turn is dependent upon pest species and abundance, which
also differed between sites in this study. The importance of
conducting taxa trials to identify pest resistance traits across
multiple sites and geographic areas cannot be overstated.
Selecting for resistance to insect pests has not been an
explicit consideration in the Corymbia breeding program;
the development of hybridisation of CCV with CT was
primarily in response to the former species’ susceptibility to
Quambalaria shoot blight, and its low amenability to vegeta-
tive propagation (Lee 2007, Shepherd et al. 2007). Overall,
Table 7: Correlation matrix showing relationships between growth
parameters and general crown damage (GCD) and arthropod
damage (AD). R-values highlighted in bold indicate a significance
level of P < 0.001. DBHOB = diameter at breast height over bark
GCD AD Height DBHOB
AD 0.50
Height –0.29 0.04
DBHOB –0.37 –0.06 0.92
volume –0.25 –0.02 0.85 0.93
Table 8: Summary of traits measured for the matched Corymbia series, CCV, CT and CT × CCV, at sites I, II and III. Relative performance
is shown as: highest score (white), intermediate (grey) and lowest score (black). All differences are statistically significant at the P ≤ 0.05
level or lower. Possible heritability patterns (H): D = dominance, Ei = epistasis for hybrid inferiority, Es = epistasis for hybrid superiority, A =
additive inheritance, n.d. = no difference
GCD AD
Taxon
I II III I II III
CCV
CT × CCV
CT
H D n.d. n.d. Ei A n.d.
151
CT was the most ‘resistant’ to arthropod damage, with about
one-quarter of trees sustaining less than 5% defoliation,
while over 95% of CCV trees exceeded this; the hybrids all
performed intermediately to parental species. Trials have
determined that the hybrids also show vigorous growth
and frost resistance (Lee 2007, Lee et al. 2009). However,
selection for crosses based on vigorous growth rates may
be at the expense of defence and therefore, potentially,
resistance to pests because there is a predicted trade-off
between growth and defence against herbivory (Coley et al.
1985, Coley 1988, Herms and Mattson 1992) and evidence
across plant species for fitness costs in deploying defences
(Koricheva 2002, Strauss et al. 2002, Henery 2006).
General crown damage was significantly negatively
correlated with DBHOB, height and volume, while PDS
showed no relationship with growth, suggesting that leaf
loss due to factors (e.g. Quambalaria shoot blight, nutrient
disorders and drought) additional to pest attack reduces
the growth rate of trees. Further field assessment of clones
derived from these taxa trials will be assessed in future studies
of hybrid susceptibility to pests in subtropical Australia.
Conclusions
Arthropod pests were responsible for about three-quarters of
crown damage, but with no consistent patterns for damage
or for growth parameters across three Corymbia species-
and-hybrid trial sites. Site was more important than taxon in
explaining damage scores, and taxa performed differently
for most traits between sites. Overall, the growth plasticity
shown across sites, increased disease resistance and growth
characteristics exhibited by hybrid Corymbia (Lee 2007)
outweigh arthropod herbivory, which showed no impact
on growth in these trials. However, it should be noted that
because these trials were polycultures, and of relatively small
size, pest risk to these taxa in monocultural commercial-scale
plantations may increase (see e.g. Stamps and Linit 1998).
Our results highlight the importance of establishing taxa trials
across a range of sites when selecting for pest resistance.
Acknowledgements — We sincerely thank Dr Andrew Hayes
(DEEDI) for critical comments on the manuscript, and John Huth,
Peter Pomroy, John Oostenbrink, Alan Ward, Tony Burridge and
Bruce Hogg (all DEEDI) for contribution to the production, establish-
ment, mensuration and documentation of the trial sites. This work
was funded with a grant from the Queensland Department of
Tourism, Regional Development and Industry, Forestry Plantations
Queensland, Forest Enterprises Australia Ltd and Elders Forestry.
DBHOB Height Volume
I II I II I II
Es A Es A Es D
152
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