MURDOCH RESEARCH REPOSITORY€¦ · As even mild deficiencies of micronutrients such as copper,...
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MURDOCH RESEARCH REPOSITORY
This is the author’s final version of the work, as accepted for publication following peer review but without the publisher’s layout or pagination.
The definitive version is available at http://dx.doi.org/10.2989/SOUTH.FOR.2008.70.2.8.536
Dell, B., Hardy, G.E.St.J. and Burgess, T. (2008) Health and nutrition of plantation eucalypts in Asia. Southern Forests,
70 (2). pp. 131-138.
http://researchrepository.murdoch.edu.au/2326/
Copyright: © NISC Pty Ltd.
It is posted here for your personal use. No further distribution is permitted.
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Health and nutrition of plantation eucalypts in Asia
B Dell*, G Hardy and T Burgess
Program for Australian Tree Health, School of Biological Sciences and Biotechnology,
Murdoch University
Abstract
Understanding site fertility constraints has lifted productivity of eucalypt plantations in East
and South-east Asia through targeted application of inorganic fertilisers. Whilst most
attention has been given to the supply of nitrogen and phosphorus in silvicultural
prescriptions, and inoculation with mycorrhizal fungi is being undertaken by some
companies, more attention is required to diagnose and correct micronutrient disorders. For
example, the incidence of boron deficiency remains a concern across the region affecting tree
growth, tree form, fibre quality and possibly the extent of damage from some eucalypt
pathogens. As even mild deficiencies of micronutrients such as copper, boron and manganese
can alter the phenol pathway, limitations in the uptake of these elements have implications
for tree defence against some fungal pathogens. Due to the high humidity and temperatures
throughout the year, fungal leaf diseases such as Cylindrocladium quinqueseptatum have had
a huge impact on the eucalypt plantation industry in South-east Asia. Often poor nutrition
seems to be an underlying cause in disease susceptibility. However, in spite of the size of the
eucalypt plantation sector, there is not an integrated understanding of biotic and abiotic
causes and their interactions by the industry. Furthermore, the emergence of new eucalypt
pathogens such as Kirramyces destructans and their rapid spread threatens the viability of
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plantations unless selection of resistant clones and breeding for resistance can keep up with
the incursion of new and invasive eucalypt pathogens. In a number of countries, there is low
genetic diversity in eucalypt plantations and breeding programs are yet to embrace screening
for disease resistance. The challenge to a successful eucalypt plantation industry in Asia is to
optimise genetics and nutrition in order to produce trees with the greatest chance of
overcoming pest and disease threats.
Keywords: invasive pathogens, Kirramyces, Leptocybe invasa, management, micronutrient
deficiencies, soil infertility
Introduction
There are now over 2.5 million ha of eucalypts in East and South-east Asia and, together with
Australia, this region is experiencing great expansion in plantation area. Hence there are vast
areas of first- and second-rotation eucalypts. Productivity range is broad, with mean annual
increment from less than 10 to over 45 m3 ha-1 y-1. In many parts of Asia, productivity is
constrained by soil infertility coupled with inadequate fertilisation and the incursion of
damaging diseases. Although macronutrients (e.g. potassium [K], nitrogen [N] and
phosphorus [P] in particular; Figure 1a) still limit productivity of wood in some plantations,
micronutrients remain problematic in many geographical regions and there is not a good
understanding of micronutrient deficiencies by some plantation managers (Xu and Dell,
2002). The continuing incidence of micronutrient disorders in plantations in Asia indicates
that this constraint to productivity has not been adequately recognised in the past. This has
resulted in operational fertiliser practices that inadvertently exclude micronutrients and hence
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some fertiliser operations remain suboptimal (Dell et al., 2001). Furthermore, during harvest
in some countries, forest farmers remove the main stem of the trees and local villagers then
collect the harvest residue (leaves and branches) and cut the understorey as fuel. Tree roots
may also be collected after soil tillage. These practices result in degradation and loss of soil
fertility (Xu et al., 2002). Retention of slash on site and fertilisation are essential to maintain
soil fertility and productivity of eucalypt plantations on these degraded lands.
In the majority of countries where eucalypts have been established as exotics, plantations
have initially been relatively pest and pathogen free. However, over time disease and pest
problems have generally increased (Wingfield, 1999; Wingfield et al., 2001; Burgess and
Wingfield, 2002). An increase in the incidence and impact of such problems is evident in the
Asian region. For example, eucalypt plantations were established over three decades ago in
Thailand and two waves of foliar diseases became problematic after several rotations. Now
the industry faces a new threat from a small wasp, Leptocybe invasa, that causes severe
damage from galls in young tissues in susceptible clones. In Vietnam, one company turned
away from eucalypts in favour of Acacia mangium due to the impact of leaf blights. Chinese
plantations, which have been largely free of disease and insect pests, are now experiencing
damage from recent introductions of fungal pathogens most likely on germplasm acquired
from elsewhere in Asia.
This paper examines some of the major constraints facing commercial eucalypt plantations in
Asia using published and unpublished data from the authors’ laboratories and observations
from recent visits to plantations in the region.
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Soil fertility constraints and plantation health
Micronutrient disorders can be induced by fertilisation with macronutrients, which promote
rapid growth of eucalypts that exceeds the supply characteristics of the plantation soil. This
has been observed by us in the case of boron (B) and Zinc (Zn) in China and the Philippines,
and copper (Cu), manganese (Mn), Zn and B in Australia. However, in parts of Yunnan, the
soils are so deficient in B that this element is the primary nutrient limiting productivity in
some districts. In central Thailand, iron (Fe) deficiency is common where eucalypts have
been planted on alkaline soils (Figure 1b). In southern Australia, land that previously was
used for agricultural pursuits of cropping and animal production on annual pasture is being
turned over to short-rotation (10 year) eucalypt plantations. Application of superphosphate
over many decades, and the use of N-fixing forage plants, has led to the build up of a bank of
P and N in these soils that are of low natural fertility. Even though some micronutrients have
been applied in the past, the amount of residual Cu for example, though adequate for annual
crops, appears to be inadequate for fast-growing eucalypts.
Eucalypts are particularly sensitive to B deficiency resulting in loss of crown vigour, shoot
death, poor stem form and reduced wood quality. Boron deficiency is a major constraint to
the productivity of eucalypt plantations in many parts of Asia (Dell et al., 2001), especially in
new plantations in China, Indonesia, Laos, the Philippines, Thailand and Vietnam (Figure
1c). Deficiencies have also been reported on the eastern seaboard of Australia, on a few
pockets of sandy soils in south-western Australia and in New Zealand. In Yunnan, E.
globulus develops a prostrate habit when NPK fertiliser lacking B is applied (Dell et al.,
2001). Symptoms of deficiency are more severe during the dry season and mildly affected
trees may partially recover during the following wet season. Sites with sandy soils derived
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from granite and sandstones are especially vulnerable. Foliar B concentrations in recently
expanded leaves lower than 9–12 mg B kg–1 dry weight are associated with deficiency
symptoms (Dell et al., 2001). However, foliar analysis is rarely undertaken in Asia to monitor
the nutritional status of industrial plantations. Furthermore, unless symptoms are so severe
that trees have a prostrate habit, the disorder is often unrecognised, with seasonal dieback and
poor form being attributed to biotic or climatic events. This is surprising given that the impact
of B deficiency on eucalypts has been known for nearly 40 years.
Foliar analysis has proven to be an effective tool in the diagnosis and prevention of
micronutrient disorders in eucalypt plantations in Australia. Since monitoring commenced c.
10 years ago, the incidence of micronutrient deficiencies in E. globulus in south-western
Australia declined by 80% in the two years after corrective fertiliser was applied (BD unpubl.
data). Micronutrient disorders have become uncommon in Australian plantations, in contrast
to those in Asia where foliar analysis is rarely used as a monitoring tool.
Inoculation with beneficial ectomycorrhizal fungi has the potential to improve fertiliser
utilisation efficiency and to increase nutrient retention onsite (Xu et al., 2001; Dell et al.,
2002). However, it is unknown whether these fungi can increase the supply of B for tree
growth as B is not essential for the growth of mycelium in culture. As a result of a decade of
trials in South China, planting stock is now being inoculated with fungi in parts of China
(mainly Yunnan, Hainan and Guangzhou). Other laboratories in the region are beginning to
examine this technology for their plantations. Whilst the principles of inoculation in the
nursery are well understood (Brundrett et al., 2005), the selection of fungi for different site
constraints is a factor that must be considered as there is no effective universal inoculum for
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eucalypts. Care must also be taken to ensure that any introduced fungi that are compatible
with eucalypts do not become invasive in native forests.
Biotic factors impacting on plantation health
Disease
A number of pathogens threaten the productivity of plantation eucalypts in tropical and
subtropical regions of East and South-east Asia (Hardy et al., 2003; Barber, 2004) and these
are summarised in Table 1. The most serious is probably Kirramyces destructans (Figure 2a,
b), first identified by Wingfield et al. (1996) on E. grandis in northern Sumatra causing
serious defoliation of the lower canopy. It quickly spread to Thailand in 2000 and Vietnam in
2002 (Old et al., 2003; Thu, 2005) and is now problematic in South China (Burgess et al.,
2006a). Movement of the pathogen long distances on infested mother and planting stock
explains the recent appearance of disease in nurseries and plantations in widely dispersed
geographical regions in China. Disease caused by this pathogen was first observed in 2003 in
central Guangdong Province and in Hainan Dao on planting stock originating from Guangxi
Province. In one Guangxi nursery visited in 2004, K. destructans had been a serious problem
since 2002. The production of cuttings from infested mother stock and poor nursery hygiene
resulted in a high proportion of diseased planting stock. Infested material has been, and
continues to be, widely distributed across South China, reaching isolated parts of south-
western Yunnan in 2003, and Fujian Province in 2006. Recent molecular analysis of eight
gene regions showed that there was no genetic variation within K. destructans isolates found
in Thailand and Vietnam, and very low genetic variation within populations from Indonesia
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and China (V Andjic pers. comm.). This suggests that the pathogen has been recently
introduced into the region. Kirramyces destructans has recently been reported on planted E.
urophylla hybrids growing on Melville Island, Northern Territory and on mature foliage of an
unknown Eucalyptus species in Derby, Western Australia (Burgess et al., 2007). From these
observations it has been concluded that K. destructans is native to Australia and has been
introduced into Asia.
Other Kirramyces species have the potential to cause partial or complete defoliation of 1–2-
year-old trees. For example, K. epicoccoides caused complete defoliation (Figure 2c, d) of
two E. urophylla clones in northern and eastern Guangdong in 2005 and 2006. Many other
Mycosphaerella spp. occur in Asia but their impact is minimal (Burgess et al., 2006b). By
contrast, in Australia Mycosphaerella has caused defoliation in several plantations (Carnegie
and Ades, 2003).
So far, pathogens causing stem cankers are having less impact than leaf and shoot blights in
Asia. Kirramyces zuluensis, an anamorph of Mycosphaerella (Andjic et al., 2007), is a
serious canker disease (Figure 3a) in many parts of the world (e.g. South Africa and Ethiopia)
and so far causes minor damage in Thailand (E. camaldulensis), Vietnam (E. urophylla and
E. camaldulensis) and China (E. urophylla) (Cortinas et al., 2006). Chrysoporthe cubensis
(previously Cryphonectria) is considered a big threat to eucalypts everywhere, including
Australia. It has been isolated from South-east Asia and China but does not appear to be
having a major impact as experienced in South Africa and South America.
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Various endophytic, stem fungal pathogens, including Holocryphia eucalypti (= Endothia
gyrosa), Cytospora spp. and various Botryosphaeriaceae tend to occur on trees that are under
stress from a range of factors including water stress (drought and waterlogging), insect attack
and from nutrient deficiencies such as Cu and B. In the tropics, Lasiodiplodia theobromae is
commonly isolated from cankers. However, in general the incidence is low and sporadic.
Bacterial wilt caused by Ralstonia solanacearum race 1, biovar III is an ongoing problem in
Eucalyptus plantations in South China (Figure 3b) and causes minor damage in other
countries. The disease mainly attacks young eucalypt trees resulting in rapid death.
Generally, death is sporadic in plantations and can occur on sites that were previously
forested as well as on sites of mixed land use. The practice of many nurseries of using field-
collected soil and then growing rooted cuttings in plastic bags on the ground may exacerbate
the spread of the pathogen into new plantations, especially in upland areas. The pathogen is
widely distributed and has an extensive host range including field and horticultural crops. No
effective control measures are available.
Pests
Unlike in Australia, where eucalypts in young plantations are often heavily attacked by
insects, most plantations in Asia have very low incidence of insect damage. Recently,
however, Leptocybe invasa (Mendel et al., 2004) has become a major problem in Thailand
(Figure 3c, d) and Vietnam. This introduced wasp induces galls on petioles, midribs and
young stems, causes deformation, reduced leaf size, loss of apical dominance and stunting of
mother plants, cuttings in nurseries, in outplanted stock and coppice of susceptible clones of
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E. camaldulensis. Some resistant clones have been identified but resistance can break down
quickly and insecticides are ineffective on mother plants and impractical in the field. No
biological control agent has been identified. With the exception of the Asian gypsy moth, it is
unlikely for native insects in Asia to become a major problem on eucalypts in that region as
these tend to have difficulty coping with secondary metabolites of eucalypts.
Micronutrients and disease
As indicated earlier, we have observed many instances of increased disease impact in
eucalypts stressed by abiotic factors such as nutrient imbalance, drought and adverse weather
conditions. As even mild deficiencies of micronutrients such as Cu, B and Mn can alter
phenol metabolism, limitations in the uptake of these elements have implications for tree
defence against some fungal pathogens. For example, there has been an increase in cankers of
opportunistic fungi such as Neofusicoccum eucalyptorum in stems of coppiced E. globulus
plantations in south-western Australia on soils that are deficient in Cu (Figure 4a, b).
In 2003, Dell and Xu (2006) were surprised to observe extensive B deficiency symptoms
associated with a new pattern of leaf necrosis (Figure 4c) in eucalypt plantations in South
China where previously corrective B treatments had only been required on highly weathered
soils derived from granite and sandstone in the region (Dell et al., 2001). Generally, some 5–
10% of trees were also showing symptoms from bacterial wilt. The incidence of bacterial wilt
and B deficiency suggested a causal relationship. It was proposed that the B deficiency
symptoms were due to reduction in B uptake caused by loss in function of near-surface roots
due to invasion of the xylem by the pathogen. Two factors that may have exacerbated the
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problem were the likely reduced uptake of B from the surface soils due to the below-average
rainfall in that year and the occurrence of two typhoons, which seriously damaged some
plantations and caused extensive root breakage, opening up infection points to bacteria
resident in the soil. Reduced strength in the wood of roots and stems resulting from incipient
B deficiency would have predisposed the trees to typhoon damage and subsequent infection.
In a B rates trial undertaken in south-western China, even mild B deficiency lowered the
lignin content of wood and reduced wood density and fibre length by up to 10% of healthy
trees (BD and D Xu unpubl. data).
Discussion
The key to managing the nutrient status of eucalypts in plantations is to have reliable
information on the micro-and macronutrient status of the trees and the soil, and a working
knowledge of biotic factors that may interfere with nutrient acquisition, utilisation or
retranslocation. Often, micronutrients are ignored until severe defects in form are reported.
By this stage, not only has there been considerable loss of potential yield but reduced fitness
can result in increased incidence of pathogen attack. Foliar analysis has proven to be effective
in identifying marginal nutrient disorders prior to visible signs being present. In Australia,
plantations are routinely monitored for nutrient status. This relies on analytical laboratories
having access to recommended foliar nutrient concentrations for healthy and deficient trees.
Data sets have been compiled so far for only a limited number of plantation species (e.g. E.
globulus, E. grandis and E. urophylla; Dell et al., 2001), and generally are for unimproved
genetics. A huge effort is required to develop new data sets for other plantation species such
as E. dunnii and various hybrids, but so far this has not been a priority for the industry even
though nutrient disorders are frequently encountered in plantations in Asia.
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In our experience, correction of a micronutrient deficiency is not difficult if the problem is
diagnosed early in the rotation and its cause is understood (Dell et al., 2003). Generally, as
micronutrient deficiencies result from tree growth outstripping the supply capacities of
plantation soil, application of fertiliser is recommended. Dramatic growth recovery has been
achieved by topdressing early with B in parts of East and South-east Asia where soils are
acutely low in B. Similarly, addition of Cu has improved tree form and growth in south-
western Australia and reduced the incidence of disease by taxa in the Botryosphaeriaceae.
However, there are no long-term prescriptions for the management of micronutrients across
rotations. Not surprisingly, therefore, disorders continue to be reported even when corrective
measures have been taken in a previous crop.
Various pathogens present in Asia impact heavily on establishment and productivity.
Management for these diseases is essential for the continued success of the industry. As the
climate is extremely favourable for disease development, inoculum levels are often high.
Thus, tree health within plantations primarily will depend upon breeding and selecting for
resistance or tolerance to a range of diseases. The tolerance of the trees will be increased by
the optimisation of site-specific fertiliser prescriptions. Besides the known pathogens, there is
also the threat of the introduction of a new disease such as eucalyptus rust (Glen et al., 2007).
In conjunction with foliar monitoring for nutrient status, strategically placed sample plots can
be used for assessment of biotic stresses, inventory information on survival and growth rates
and monitoring of soil indices. In Asia, until recently, there has been little systematic
monitoring of tree health in plantations. One of the values of having monitoring programs is
that it allows the early detection of pests and pathogens, and provides information to the
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breeders on any differential response of clones. For known diseases that exist elsewhere,
important clonal lines can be taken and screened where the disease is present. However,
regular monitoring of the current plantation estate is the only way to discover and potentially
combat unknown pests and disease.
There has been little attempt to assess differences in genotypes of plantation eucalypts for
their capacity to do well on soils where micronutrient disorders are problematic. Indeed, most
eucalypt screening trials in Asia and Australasia in the past have been undertaken without
regard for either the fertiliser inputs that are possible in well-managed plantations or have
failed to recognise the importance of micronutrients by the application of so-called complete
fertilisers that are often inadequate in their contents of micronutrients (Dell et al., 2003). Yet,
there are several clonal field trials where differences in growth appear to be related either to
improved micronutrient acquisition or more efficient micronutrient utilisation, which
suggests possibilities for selection and breeding in the future. Recently, a few Fe-efficient and
B-efficient clones, predominantly of E. camaldulensis, have been identified in Thailand.
In Thailand a number of laboratories are actively developing screening methods for eucalypt
clones resistant to Cryptosporiopsis eucalypti (e.g. Singchada et al., 2006), Cylindrocladium
quinqueseptatum and Kirramyces destructans. The impact of the recent incursion of
Leptocybe invasa has resulted in some fast-growing clones being abandoned. The early
indication is that resistance in some other clones is beginning to break down.
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By contrast with Thailand, the magnitude of disease and pest problems in China remains
relatively small but the threats continue to mount as new pathogens spread into that part of
Asia. This is of great concern as the number of eucalypt clones that have been mass planted is
very small (Arnold, 2005) and there has been little progress so far in breeding for tolerant
lines. Pests and diseases present risks to commercial plantations, especially clonal plantations
based on relatively few clones. Therefore, it can be argued that dependency on such a narrow
diversity of clones in China places the industry at risk of pests and disease in the future. A
great deal of effort is required to breed fast-growing clones that are tolerant to abiotic and
biotic stresses and have desirable fibre qualities. The potential benefits gained by improved
breeding and silvicultural practices will need to be balanced by the increased risks associated
with nutrient imbalances, pests and diseases. This is particularly true in plantations, which are
usually of a uniform age, densely planted and of a narrow genetic range. Consequently, it
must be expected that fungal and bacterial diseases together with pests will, with time,
become important economic constraints to long-term plantation health management.
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Figure 1. Common nutrient disorders in eucalypt plantations in Asia. (a) Symptoms of
macronutrient deficiencies taken from trees in one compartment in Yunnan. Left to right:
healthy, low K, low N and low P. (b) Fe deficiency in Thailand. (c) B deficiency in Yunnan
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Figure 2. Impact of Kirramyces destructans (a, b) and K. epicoccoides (c, d) on canopy loss
of Eucalyptus urophylla in Guangdong Province, China in 2005–2006. Total leaf shedding
occurred in (c) and the emergent new growth was severely impacted by B deficiency, hence
the multiple epicormics at the nodes (d)
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Figure 3. (a) Canker caused by Kirramyces zuluensis in Vietnam, 2006. (b) Sporadic death
due to Ralstonia solanacearum in China, 2006. Impact of Leptocybe invasa on cuttings (c)
and coppice (d) of Eucalyptus camaldulensis in Thailand, 2007
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Figure 4. Examples of micronutrient × disease interactions. Neofusicoccum eucalyptorum
cankers in the mid-trunk (a) and near the crown apex (b) in Cu-deficient Eucalyptus globulus
in Australia, 2007. Leaf necrosis (pockets of dead tissue) and B-deficiency symptoms
(interveinal chlorosis, cupping, brittleness) in E. urophylla in which the xylem of the main
lateral roots and trunk were infested with Ralstonia solanacearum in China, 2003
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Table 1. Current and potential major pathogens that threaten productivity of eucalypt
plantations in Asia and Australasia
Type Name Status Impact* Leaf and shoot blight Cryptosporiopsis eucalypti Well established in South-east Asia Severe in Thailand and Vietnam Cylindrocladium quinqueseptatum Well established in the region Severe in Thailand and Vietnam; moderate
in China Kirramyces destructans Spreading rapidly in the region,
recently recorded for mainland Australia
Severe in Sumatra, Thailand and Vietnam; moderate in China; potentially a serious threat to tropical/subtropical plantations in Australia
Kirramyces epicoccoides Well established in the region Moderate in parts of China and Indonesia
Rust Puccinia psidii Not present in the region Unknown but potentially severe
Canker Chrysoporthe cubensis Common in South-east and East Asia Minor impact
Kirramyces zuluensis Common in South-east and East Asia Minor impact, potentially a threat to Australia
Bacterial wilt Ralstonia solanacearum Some biovars common in Asia Moderate in Vietnam and China
* Impact is sometimes species or hybrid sensitive