American Journal of Plant Sciences, 2013, 4, 896-904
http://dx.doi.org/10.4236/ajps.2013.44110 Published Online April 2013 (http://www.scirp.org/journal/ajps)
Growth of Shorea contorta Vid. Inoculated with Eucalypt
Ectomycorrhizal Fungi in the Nursery and in a
Logged-Over Dipterocarp Forest in Surigao, Philippines
Nelly S. Aggangan1*, Mitzi Ann T. Pollisco2, Jeremias B. Bruzon3, Joan S. Gilbero3
1National Institute of Molecular Biology and Biotechnology, University of the Philippines Los Baños, College, Philippines; 2Eco-
systems Research and Development Bureau, Department of Environment and Natural Resources, Los Baños, Philippines; 3Depart-
ment of Environment and Natural Resources, Ecosystems Research and Development Services, Surigao City, Philippines.
Received February 29th, 2013; revised April 1st, 2013; accepted April 9th, 2013
Copyright © 2013 Nelly S. Aggangan et al. This is an open access article distributed under the Creative Commons Attribution Li-
cense, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
ABSTRACT
This study was conducted to determine the compatibility of ectomycorrhizal (ECM) fungi associated with eucalypts on
dipterocarps. Two nodal cuttings of Shorea contorta (Vid.) were rooted in a non-mist system for two months, and later
inoculated with vegetative mycelia of three strains of Pisolithus collected under eucalypts and a strain of Scleroderma
from dipterocarps. Inoculated rooted cuttings were planted in irradiated potting mix and raised in the nursery for five
months. Root colonization prior to outplanting ranged from 31% to 38% and ECM fungi did not colonize the uninocu-
lated ones. The cuttings were outplanted in a logged-over dipterocarp forest in Bislig, Surigao Sur following a Ran-
domized Complete Block Design with four blocks; each block was planted with 50 seedlings. Two years after outplant-
ing, Pisolithus strain from New Caledonia (H6394) promoted the highest height (46 cm) and diameter (0.48 cm) incre-
ment. Height was increased by 17% and diameter by 7% relative to the uninoculated control. By contrast, Pisolithus
strain from the Philippines (H615) gave the shortest (26 cm) height increment and smallest (0.42 cm) diameter. Diame-
ter growth of cuttings inoculated with Scleroderma D01 (from the Philippines) and a Pisolithus strain from Australia
(H445) was bigger (0.47 cm) than the uninoculated treated cuttings (0.45 cm). In terms of survival, the uninoculated
cuttings gave the highest (90%) percent survival while the lowest (60%) was those inoculated with Pisolithus strain
from New Caledonia. It is possible that the indigenous ECM fungi infected easily the roots of the uninoculated cuttings
thus contributing to the high survival rate (90%). During outplanting, fruit bodies of S. columnare were present in the
field site. The results, however, show that ECM fungi were able to colonize the roots and that some strains promoted the
growth and survival of S. contorta planted in a logged-over dipterocarp forest in Bislig, Surigao, Philippines.
Keywords: Ectomycorrhiza; Pisolithus; Scleroderma; White Lauan; Rooted Cuttings
1. Introduction
The lowland dipterocarp forests of Southeast Asia in-
cluding Malaysia, Indonesia and the Philippines, are
dominated by trees belonging to the family Dipterocar-
paceae. They are of considerable environmental and
economic value at the local, regional and global scale,
providing many goods and services to a growing popula-
tion [1]. The forests of this region are among the fastest
disappearing in the world and restoration is urgently re-
quired. The importance of species-site matching, my-
corrhizal fungi and post-planting maintenance for resto-
ration are apparent [1].
Dipterocarpaceae consist of 17 genera and approxi-
mately 500 species worldwide and the largest genera are
Shorea (196 species), Hopea (104 species), Dipterocarpus
(70 species), and Vatica (65 species) [2,3]. Shorea con-
torta Vid. (white lauan) is one of the most important
dipterocarp species because of its economic value and
fast growth. Dipterocarp trees are reported to be strictly
associated with ectomycorrhizal (ECM) fungi [4-8]. The
fungal species involved are Basidiomycetes and Asco-
mycetes [9]. These ECM fungi play important roles in
nutrient and biogeochemical cycle in forest ecosystems
[9,10]. Ectomycorrhizal fungi reported to be associated
with dipterocarps are: Amanita, Boletus, Cantharellus,
Russula, Lactarius, Laccaria and Scleroderma [4,11-15].
*Corresponding author.

Growth of Shorea contorta Vid. Inoculated with Eucalypt Ectomycorrhizal Fungi in the
Nursery and in a Logged-Over Dipterocarp Forest in Surigao, Philippines
897
Many reports have shown that ECM fungi increase the
growth and nutrient uptake of dipterocarps seedlings
[16-18], can act as biological control against root patho-
gens [19] and increase drought tolerance [20].
Pisolithus and Scleroderma are the well-studied gen-
era of ECM fungi and the most common in young planta-
tions of pines, eucalypts and acacias in the Philippines.
The spores of these ECM fungi are the components of
Mycogroe tablets developed at the National Institute of
Molecular Biology and Biotechnology (BIOTECH),
University of the Philippines Los Baños (UPLB), Col-
lege, Laguna, which are now commercially distributed in
the country [21]. Under dipterocarps, Sclerodermas are
relatively common but no Pisolithus so far was observed.
Mycogroe tablets containing spores of Scleroderma and
Pisolithus from pines and eucalypts were previously re-
ported to increase growth of dipterocarps planted in a
marginal grassland [15].
The function of ECM in promoting tree growth and
survival in marginal sites depends on the compatibility of
the host and the fungus [22]. It is therefore crucial to
determine compatibility of ECM fungi on dipterocarps.
Compatibility is defined as the merging of symbionts to
form a structurally defined mycorrhiza in which physio-
logical exchange of metabolites between the partners,
improve nutrition of host plant and the fungus. In a com-
patible association, infected roots have a fungal mantle, a
well-developed Hartig net and a radially elongated epi-
dermal cells in the case of eucalypts [23]. On the other
hand, signs of incompatibility include, the production of
phenolics in root tissue and tissue disorganization which
produces deleterious effects on the fungus or hosts or
both [24].
Ectomycorrhizal plants vary widely in compatibility
with ECM fungi [23] and the number of fungal associates
differs strongly among ECM hosts [24]. Smits [4] and
Becker [26] reported that ECM fungi of some diptero-
carp species in Malaysia are restricted to single host spe-
cies. Soil from beneath the ECM tree Dipterocarpus
grandiflorus failed to provide ECM fungi for S. obtusa of
the same family [4]. Likewise, S. leprosula and S. max-
welliana seedlings growing in the same vicinity only
shared two of twenty ECM types [26].
The idea of inoculating dipterocarps with ECM fungi
associated with eucalypts was based on the observation
in Indonesia and in the Philippines that S. columnare was
found underneath eucalypts and dipterocarps. Three spe-
cies of the genus Pisolithus have been described from
dipterocarp species. Pisolithus aurantioscabrosus has
been reported from Malaysia under Shorea macroprera
[27], P. adbitus collected under Dipterocarpus alatus in
Thailand [28] and P. indicus associated with Vatica in-
dica in India [29]. In the Philippines, no basidiome of
Pisolithus was ever observed under dipterocarps [12-14].
Since Pisolithus is cosmopolitan in both tropical and
temperate regions and forms ECM association with wide
range of woody plants, the aim of this study was to de-
termine the growth performance of S. contorta rooted
cuttings inoculated with three isolates of eucalypts’ ECM
fungus Pisolithus and a dipterocarps’ Scleroderma iso-
late under nursery and in a logged-over dipterocarp for-
est in Bislig, Surigao City, in the island of Mindanao,
Philippines.
2. Materials and Methods
2.1. Experimental Design
There were five inoculation treatments which was con-
ducted following a Randomized Complete Block Design
with five blocks in the nursery and four blocks in the
field. Under nursery conditions, there were 100 plants
per treatment. The nursery phase was conducted at the
screenhouse of the National Institute of Molecular Biol-
ogy and Biotechnology (BIOTECH), University of the
Philippines Los Baños (UPLB) for five months and
shipped to Surigao a month prior to outplanting. Each
block contained ten seedlings planted in a row plot with a
spacing of one meter and 2 meters in between plots.
2.2. Field Site
The site is located within the Dipterocarp Forest Re-
search Center, Barangay Maharlika, Bislig, Surigao del
Sur. The experimental area is about 0.30 hectare and it is
contiguous with rolling terrain and elevation ranging
from 100 - 200 m asl. The area is a second growth forest
supporting lesser known tree species but dominated by
dipterocarp trees with diameter ranging from 50 to 80 cm.
The climatic type is 4 (with no distinct dry and wet sea-
son). Maximum rainfall is from November to March. The
average monthly rainfall is 3472 mm, mean temperature
is 24.5˚C and mean relative humidity of 86%. The soil is
sandy clay loam with pH of 5.9, 2.48% organic matter,
0.06% total nitrogen and 0.21 mg·kg−1 available phos-
phorus.
2.3. Production of Rooted Cuttings
Cutting propagation as an alternate techniques for the
mass production of genetically and phenotypically supe-
rior dipterocarp planting stocks has been practiced [30,
31]. Rooted cuttings used in this experiment were pro-
duced at the Ecosystems Research and Development Bu-
reau (ERDB), Department of Environment and Natural
Resources (DENR), Los Banos, Laguna using two nodal
cuttings technique of Pollisco [30]. Cuttings were ob-
tained from a clonal hedge garden established near the

Growth of Shorea contorta Vid. Inoculated with Eucalypt Ectomycorrhizal Fungi in the
Nursery and in a Logged-Over Dipterocarp Forest in Surigao, Philippines
898
ERDB building. Two nodal cuttings of S. contorta were
rooted in a non-mist system for two months.
2.4. ECM Production and Inoculation of Rooted
Cuttings
Cultures of Pisolithus H6394 from New Caledonia and
H445 from Australia were provided by Dr. Bernie Dell
of Murdoch University, Perth, Western Australia. Pi-
solithus H615 was collected in Bukidnon in the island of
Mindanao, Philippines. The three Pisolithus isolates were
associated with eucalypts. A local strain of Scleroderma
(DO1) was collected from under dipterocarp forest
growing in Zambales, Luzon. The four ECM fungi were
mass produced and maintained at the National Institute
of Molecular Biology and Biotechnology, University of
the Philippines Los Baños, College, Laguna on a Modi-
fied Melin Norkran’s (MMN) agar medium [32].
Rooted cuttings were inoculated with one-month old
vegetative mycelia of three isolates of Pisolithus, and an
isolate of Scleroderma. Mycelial disks (3 cm diameter)
were placed in contact with the roots. The inoculated
rooted cuttings were transplanted in 4" × 8" PE bags
filled with irradiated potting mix 1:1:1 garden soil, sand
and coir dust), raised for five months in the nursery
where growth was monitored monthly. Each treatment
consisted of 100 seedlings. The seedlings were later
shipped to Mindanao. After one month of acclimatization,
the seedlings were outplanted in a logged-over diptero-
carp forest under the jurisdiction of the Dipterocarp For-
est Research Center, Ecosystems Research and Devel-
opment Services-DENR building, Bislig, Surigao Sur.
Prior to outplanting, the initial height and stem diameter
was measured.
2.5. Plant Growth Measurement
Height of nursery grown seedlings were measured prior
to outplanting in the field. Height and diameter were
measured at two, four and eight years after transplanting.
Height was measured using a meter stick and later a
bamboo stick at 10 cm above the soil surface up to the
shoot tip. Diameter was measured using a vernier caliper
and later using a diameter tape at 10 cm above the soil
surface. Wood volume was calculated using the formula
(Stan Rance, personal communication):

2
Wood volumeDGLEHeight3.141612
where


Representative eight-year-old trees (height ranged
from the smallest to the tallest) were harvested (cut 10
cm above the ground) at random covering the whole ex-
perimental plot similar to that done on eucalypts [33].
Wood volume of trees was computed using the equation
derived from a regression analysis.
2.6. Assessment of Mycorrhizal Infection
Destructive sampling was done prior to shipping of dip-
terocarp seedlings. The root system of plants (5 seedlings
per treatment) in the nursery was examined under a mi-
croscope for the presence of ectomycorrhizas. Root sam-
ples were also collected at two and eight years to assess
root colonization by the inoculant fungi. Surface soil was
scraped off using a hand trowel and fine roots (0.3 - 0.5
m away from the base of the tree) were detached from
the main roots growing on two sides of randomly se-
lected trees (2 trees per treatment per block). Preliminary
examination of fresh collected fine roots indicated dis-
tinctive yellow Pisolithus mycorrhizas. Root samples
were washed in water then later preserved in 50% etha-
nol until these were cleared and stained with lactic-glyc-
erol trypan blue [34]. Stained roots were spread evenly
on Petri plates with grid lines and the presence or ab-
sence of ectomycorrhizas (fully colonized root tips) were
recorded on roots that crossed the grid lines [35]. The
presence of fruit bodies of ectomycorrhizal fungi was
assessed after outplanting.
2.7. Statistical Analysis
Height, diameter and wood volume were analyzed statis-
tically using an Analysis of Variance of a Randomized
Complete Block Design [36]. Treatment means were
compared using a Duncan Multiple Range Test at p <
0.05 [37]. Statistical analysis was done using MSTATC
computer program [38].
3. Results
3.1. Mycorrhizal Root Colonization and Fruit
Bodies of ECM Fungi
Prior to outplanting, the roots tips were colonized (11% -
38%) by ECM fungi (Figure 1). The uninoculated plants
were not mycorrhizal. Two years after outplanting, root
colonization by the different ECM fungi studied ranged
from 11% to 38% and no ectomycorrhizal root tips was
observed on the uninoculated plants.


DGLE Diameter at ground level
Diameter at 10 cm above the groundHeight
Height10 cm



During outplanting, fruit bodies of Scleroderma spe-
cies were observed in the nursery close to the experi-
mental area. Fruit bodies were sessile as well as stalked
Sclerodermas most likely S. columnare. Eight years after
outplanting, root samples collected from the root zone

Growth of Shorea contorta Vid. Inoculated with Eucalypt Ectomycorrhizal Fungi in the
Nursery and in a Logged-Over Dipterocarp Forest in Surigao, Philippines
899
Figure 1. Roots of 6-month-old dipterocarp seedlings colo-
nized by different ectomycorrhizal fungi prior to outplant-
ing in a logged-over dipterocarp forest in Bislig, Surigao
City, Mindanao Island, Philippines.
revealed that only those inoculated with the Philippine
Pisolithus (H615) had mycorrhizal association. The roots
were golden yellow and 10% to 20% of the root tips were
colonized with Pisolithus H615.
3.2. Seedling Growth and Mycorrhizal
Colonization Prior to Outplanting
Prior to outplanting, seedlings inoculated with a eucalypt
Pisolithus strain from Australia (H445) and from New
Caledonia (H6394) promoted the largest leaf size/area,
greener and healthier than those inoculated with Philip-
pine ECM fungus (H615) and much more with the uni-
noculated control seedlings (Figure 2). By contrast, cut-
tings inoculated with Philippine Scleroderma (D01) iso-
late from dipterocarps and the Australian eucalypt Pi-
solithus gave the highest height increment (Figure 3).
Height was increased by 22% and 20%, respectively.
3.3. Seedling Survival under Field Conditions
The uninoculated plots gave the highest (90%) seedling
survival two years after outplanting (Figure 4(a)). By
contrast, plots inoculated with the Australian Pisolithus
H445 gave the lowest (63%). Among the mycorrhizal
plants, those inoculated with the New Caledonian Pi-
solithus H6394 gave a seedling survival of 83%, Philip-
pine Pisolithus of 70% and the dipterocarp Scleroderma
of 67%. Four years after outplanting, the uninoculated
treatment had a great drop of the number of surviving
plants from 90% at two years to 40% at four years after
outplanting (Figure 4(b)). Seedlings inoculated with the
Philippine Pisolithus gave the highest number of surviv-
ing seedlings (53%). Eight years after outplanting, seed-
lings inoculated with the New Caledonian Pisolithus
gave the lowest (13%) survival while the Philippine Pi-
solithus and Scleroderma gave the highest with seedling
survival of 37% and 40%, respectively (Figure 4(c)). The
uninoculated treatment gave the second lowest (23%)
Figure 2. Six-month-old S. contorta rooted cuttings due to
inoculation with three isolates of Pisolithus (H6394, H615,
H445) and a Scleroderma (D01) isolates prior to outplanting
in Bislig, Surigao, Mindanao.
Figure 3. Height increment of six-month-old S. contorta
rooted cuttings due to inoculation with three isolates of Pi-
solithus (H6394, H615, H445) and a Scleroderma (D01) iso-
lates prior to outplanting in Bislig, Surigao, Mindanao is-
land, Philippines. Bar represe nts LSD value at p < 0.05.

Growth of Shorea contorta Vid. Inoculated with Eucalypt Ectomycorrhizal Fungi in the
Nursery and in a Logged-Over Dipterocarp Forest in Surigao, Philippines
900
Figure 4. Seedling survival of S. contorta inoculated with
eucalypt and dipterocarp ectomycorrhizal fungi Pisolithus
(H445, H6394 and H615) and Scleroderma (D01) isolates 2
(a), 4 (b) and 8 (c) years after outplanting in a logged over
dipterocarp forest in Bislig, Surigao City, Mindanao island,
Philippines. Bars represe nt LSD value at p < 0.05.
seedling survival.
3.4. Seedling Growth under Field Conditions
Two years after outplanting, height growth was highest
in cuttings inoculated with Pisolithus H6394 (Figure
5(a)). Pisolithus H6394 increased height by 20% over the
control, two years after outplanting. However, after an-
other two years, those inoculated with Scleroderma out-
grew all the rest of the treated counterpart (Figure 5(b)).
Scleroderma D01 increased height by 23% relative to the
uninoculated one. Eight years after outplanting, the Phil-
ippine Pisolithus was the only ECM fungi that promoted
height of S. contorta (Figure 5(c)). Philippine Pisolithus
promoted a 23% taller height than the control (453 cm).
On the other hand, the Australian inoculated cuttings
gave the shortest height eight years after outplanting.
In terms of stem diameter, ECM inoculation did not
significantly increase stem diameter two years after out-
planting (Figure 6(a)). However, after another two years
in the field, Scleroderma D01 treated cuttings outgrew
the other ECM inoculated seedlings by 29% relative to
the control (Figure 6(b)). The Philippine Pisolithus gave
the lowest stem diameter during the four-year growth
(Figure 5(b)), but promoted the highest eight weeks after
Figure 5. Height of S. contorta inoculated with eucalypt and
dipterocarp ectomycorrhizal fungi Pisolithus (H445, H6394
and H615) and Scleroderma (D01) isolates 2 (a), 4 (b) and 8
(c) years after outplanting in a logged over dipterocarp
forest in Bislig, Surigao City, Mindanao island, Philippines.
Bars represent LSD value at p < 0.05.
planting (Figure 6(c)). This Pisolithus isolate promoted
stem diameter of S. contorta by 18%.
3.5. Wood Yield under Field Conditions
The New Caledonian eucalypt Pisolithus H6394 pro-
moted the highest (10 cm3) wood volume two years after
outplanting while those inoculated with the other ECM
fungi gave a wood volume lower than the uninoculated
ones (Figure 7(a)). Two years later, wood volume ob-
tained by those inoculated with the Australian Pisolithus
was lower (74 cm3) than the uninoculated ones (83 cm3)
and those inoculated with Scleroderma gave the highest
(164 cm3). Scleroderma doubled the wood volume ob-
tained by the control plants (Figure 7(b)) whereas, plants
with the Philippine Pisolithus gave the lowest (35 cm3).
Eight years after outplanting, the Philippine Pisolithus
isolate was the only ECM that promoted wood volume.
This local eucalypt isolate increased wood volume by
88% relative to the control (1829 cm3) (Figure 6(c)).
4. Discussion
The potential for improved establishment of dipterocarp
seedlings or cuttings with the inoculation with ECM

Growth of Shorea contorta Vid. Inoculated with Eucalypt Ectomycorrhizal Fungi in the
Nursery and in a Logged-Over Dipterocarp Forest in Surigao, Philippines
901
Figure 6. Stem diameter of S. contorta inoculated with
eucalypt and dipterocarp ectomycorrhizal fungi Pisolithus
(H445, H6394 and H615) and Scleroderma (D01) isolates 2
(a), 4 (b) and 8 (c) years after outplanting in a logged over
dipterocarp forest in Bislig, Surigao City, Mindanao island,
Philippines. Bars represe nt LSD value at p < 0.05.
fungi has been demonstrated for several species [16,39-
44]. Survival of dipterocarp seedlings was proven to be
more dependent on ECM association than on light inten-
sity and soil properties [45]. In this study, the four ECM
fungi studied differentially influenced growth response
of S. contorta under nursery and field conditions. Plants
inoculated with the exotic eucalypt Pisolithus H445 was
the best in promoting growth, robust vegetative growth
with larger leaf area and greener in color as compared
with the other plants treated with the other exotic euca-
lypt Pisolithus (H6394) isolate and the native eucalypt
Pisolithus (H615) and dipterocarp Scleroderma (D01)
isolates. In an earlier study, it was concluded that, the
exotic eucalypt Pisolithus H6394 isolate and native dip-
terocarp Astraeus isolate can be used to inoculate Ani-
soptera thurifera. Furthermore, Pisolithus H6394 can
also be used for S. guiso rooted cuttings grown in Philip-
pine red soil and under screenhouse conditions [44].
Growth of Hopea odorata and Hopea helferi was also
reported to be stimulated by inoculation with an ECM
strain of Pisolithus tinctorius under aseptic conditions
[40]. This implies that Hopea odorata and Hopea helferi
seedlings respond to ECM infection, and exhibit a low
level of host-fungus specificity [24,40].
Figure 7. Wood volume of S. contorta inoculated with euca-
lypt and dipterocarp ecto mycorrhizal fungi Pisolithus (H445,
H6394 and H615) and Scleroderma (D01) isolates 2 (a), 4 (b)
and 8 (c) years after outplanting in a logged-over diptero-
carp forest in Bislig, Surigao City, Mindano Island, Philip-
pines. Bars represent LSD value at p < 0.05.
Inocula isolated from one tree species can be success-
fully inoculated into other species and even genera, lead-
ing to improved mineral nutrition and drought tolerance
[43]. The Australian eucalypt Pisolithus H445 was in-
troduced in the Philippines in 1992 by the senior author.
This fungus promoted the growth of Eucalyptus uro-
phylla grown in the nursery and in marginal grassland in
Malaybalay, Bukidnon, Mindanao [46]. The plant growth
promoting effect was evident even up to five years after
outplanting [22]. However, this fungus never produced
fruiting bodies in spite of the presence of infection in the
roots [22]. In the present study, Pisolithus H445 was the
most promising ECM fungus during the first two years of
S. contorta outplanted in a logged-over dipterocarp forest
in Bislig, Surigao Sur, Mindanao. Thus implies that this
fungus is a good potential inoculant for S. contorta at
least during early seedling stage.
At two years, the uninoculated cuttings outgrew those
inoculated with a Scleroderma isolate from dipterocarp
(DO1), the Pisolithus from New Caledonia (H6394) and
also the Philippine Pisolithus (H615). It is possible that
the uninoculated plants were colonized with growth pro-
moting indigenous ECM fungi present in the site. The
experimental site is a logged over secondary dipterocarp

Growth of Shorea contorta Vid. Inoculated with Eucalypt Ectomycorrhizal Fungi in the
Nursery and in a Logged-Over Dipterocarp Forest in Surigao, Philippines
902
forests so, definitely there are indigenous ECM fungi
associated with the remaining dipterocarp trees. Fruiting
bodies of ECM fungi were observed in the nursery and
the adjacent dipterocarp forests about fifty meters away
from the experimental site during planting. ECM fungi
were different species of Scleroderma and ectomycor-
rhizal mushrooms. Moreover, since the seedlings were
made to acclimatize in the nursery for one month, the
seedlings were possibly contaminated with spores from
the nursery.
The Australian eucalypt Pisolithus H445 was the most
promising ECM isolate at two years. However, growth
and yield of S. contorta due to inoculation with this fun-
gus was lower than the control during the four and eight
year observation period. This implies that this fungus
may not have persisted in a logged over dipterocarp for-
est after two years. The root samples collected eight
years after outplanting were not colonized with ECM
fungi. Scleroderma species from a dipterocarp forest in
Zambales doubled the wood volume in relation to the
uninoculated ones at four years but it was no longer ef-
fective at eight years. The Philippine Pisolithus from
eucalypt forests was less effective in promoting growth
and yield during early seedling stages. However, eight
years after outplanting, the Philippine Pisolithus isolate
promoted the highest height increment and the roots have
distinct brilliant golden yellow color. This indicates that,
the indigenous Pisolithus, which was collected from
eucalypts, survived in the roots of S. contorta for eight
years even though the growth promoting effects during
the early seedling stage was not so prominent.
A similar study was conducted by [47] where an exotic
strain of Pisolithus was inoculated into two dipterocarp
species. They found that although inoculation enhanced
the growth of the dipterocarps, the performance and per-
sistence of the inoculum in logged-over forest was not
successful. Kikuchi examined the role of mycorrhizas of
dipterocarp seedlings in plantation and in forests by
quantification of mycorrhizas [48]. He found that seed-
lings of 2 to 4 m high have numerous mycorrhizas and
that the amount of mycorrhizas increased as the seedlings
became bigger both in plantation and in natural regenera-
tion forests. Abundant mycorrhizas were found in big
seedlings. In the present study, it seems that only those
inoculated with the Philippine Pisolithus H615 isolate
had few (10% to 15%) mycorrhizas (but no fruiting bod-
ies) and gave the highest wood yield eight years after
outplanting. The other ECM inoculated plants did not
have any mycorrhizal roots in spite of the presence of
fruit bodies of Scleroderma (S. columnare) in the nearby
plantations and nursery. The thick organic matter on the
soil surface may have prevented the formation of fruit
bodies. Basidiomes of Sclerodermas in the nearby plan-
tations were observed in areas with no or very little soil
covering.
In conclusion, the results indicate that Pisolithus H445
from eucalypts growing in Australia was the most effec-
tive in promoting growth and wood yield of S. contorta
two years after out planting in a logged over dipterocarp
forest in Bislig, Surigao Sur. Scleroderma D01 from dip-
terocarp forest in the Philippines was the most effective
in promoting growth and wood yield at four years after
out planting while Pisolithus from the eucalypt planta-
tions in the Philippines was the most effective in pro-
moting growth and wood yield of S. contorta eight years
after out planting in a logged over dipterocarp forest. It is
therefore recommended, that Pisolithus H445, Sclero-
derma D01 and Pisolithus H615 can be mass produced to
address growth promotion of S. contorta under nursery
and field conditions such as those in logged-over dip-
terocarp forests in Bislig, Surigao City in the island of
Mindanao, Philippines. Moreover, it is important to iso-
late ectomycorrhiza inoculum that persists on diptero-
carps and show signs of improving seedling growth and
survival in the early stages of establishment.
5. Acknowledgements
This project was partly and jointly funded by the INCO-
DC; International Cooperation with Developing Coun-
tries Contract # ERBIC18CT-98319 entitled “Harnessing
mycorrhizal symbiosis in mixed Acacia mangium and
dipterocarp plantations in Malaysia and in the Philip-
pines”; National Institute of Molecular Biology and Bio-
technology, University of the Philippines Los Banos and
the Department of Environment and Natural Resources,
Caraga Region, Butuan City, Mindanao island, Philip-
pines.
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