Fungal diversity on fallen leaves of Ficus in northern Thailand

Abstract

Fallen leaves of Ficus altissima, F. virens, F. benjamina, F. fistulosa and F. semicordata, were collected in Chiang Mai Province in northern Thailand and examined for fungi. Eighty taxa were identified, comprising 56 anamorphic taxa, 23 ascomycetes and 1 basidiomycete. Common fungal species occurring on five host species with high frequency of occurrence were Beltraniella nilgirica, Lasiodiplodia theobromae, Ophioceras leptosporum, Periconia byssoides and Septonema harknessi. Colletotrichum and Stachybotrys were also common genera. The leaves of different Ficus species supported diverse fungal taxa, and the fungal assemblages on the different hosts showed varying overlap. The fungal diversity of saprobes at the host species level is discussed.

Full text

Wang et al. / J Zhejiang Univ Sci B 2008 9(10):835-841 835
Fungal diversity on fallen leaves of Ficus in northern Thailand*
Hong-kai WANG1, Kevin D. HYDE†‡2, Kasem SOYTONG3, Fu-cheng LIN†‡1
(1Biotechnology Institute, Zhejiang University, Hangzhou 310029, China)
(2Fungal Research Group, School of Science, Mae Fah Luang University, Tasud, Chiang Rai 571000, Thailand)
(3Faculty of Agricultural Technology, King Mongkut’s Institute of Technology Ladkrabang, Ladkrabang, Bangkok 10520, Thailand)
†E-mail: kdhyde1@gmail.com; fuchenglin@zju.edu.cn
Received July 28, 2008; revision accepted Aug. 15, 2008
Abstract: Fallen leaves of Ficus altissima, F. virens, F. benjamina, F. fistulosa and F. semicordata, were collected in Chiang
Mai Province in northern Thailand and examined for fungi. Eighty taxa were identified, comprising 56 anamorphic taxa, 23
ascomycetes and 1 basidiomycete. Common fungal species occurring on five host species with high frequency of occurrence were
Beltraniella nilgirica, Lasiodiplodia theobromae, Ophioceras leptosporum, Periconia byssoides and Septonema harknessi. Col-
letotrichum and Stachybotrys were also common genera. The leaves of different Ficus species supported diverse fungal taxa, and
the fungal assemblages on the different hosts showed varying overlap. The fungal diversity of saprobes at the host species level is
discussed.
Key words: Ficus, Fallen leaves, Saprobes, Fungal diversity
doi:10.1631/jzus.B0860005 Document code: A CLC number: Q939.5
INTRODUCTION
Fungi are chemoheterotrophic organisms that
play an important role as decomposers in natural
ecosystems (Duarte et al., 2006; Gadd, 2007). Fungi
have additional economic importance as biocontrol
agents, chemical producers of bioactive compounds
used in the pharmaceutical and many other industries
(Yuen et al., 1999; Bucher et al., 2004; Duarte et al.,
2006). The economic impact of fungi in the envi-
ronment is likely related to the total fungal species
pool (Hyde, 2001; Piepenbring, 2007). Research on
fungal diversity therefore provides a basis for esti-
mating the functional role of fungi in ecosystems,
exploiting these economic fungi and estimating the
total fungal species richness in a region or globally
(Arnold et al., 2001; Mueller and Schmit, 2007).
Although the 1.5 million species estimated by
Hawksworth (2001) are commonly accepted (Hyde,
2001; Hyde et al., 2007), the actual quantity of fungal
species is unclear (Arnold et al., 2001; Hyde, 2001;
Hyde et al., 2007; Schmit and Mueller, 2007). Schmit
and Mueller (2007) estimated that there is a minimum
of 712 000 fungal species worldwide. This estimate
was primarily based on the observed ratio between
vascular plant species diversity and fungal diversity
in a certain area. Meta-analysis of fungal diversity in
small plots (Schmit et al., 2005) demonstrated that the
numbers of tree species and macrofungal species
diversities are correlated (Mueller et al., 2007). The
actual number of fungi is still unknown; however,
only 5%~13% of the total estimated global fungal
species have been described. The undescribed fungi
may occur in poorly studied countries, hosts, habitats,
niches, or tissues, and are mostly microfungi (Hyde,
2001; Schmit and Mueller, 2007). To determine the
accuracy of fungal estimates, more information is
needed concerning species diversity in poorly studied
areas and hosts, especially in poorly studied areas
Journal of Zhejiang University SCIENCE B
ISSN 1673-1581 (Print); ISSN 1862-1783 (Online)
www.zju.edu.cn/jzus; www.springerlink.com
E-mail: jzus@zju.edu.cn
‡ Corresponding authors
* Project supported by the Mushroom Research Centre, Chiang Mai,
Thailand and the National Natural Science Foundation of China (No.
30670072) and the Natural Science Foundation of Zhejiang Province
of China (No. Y350568)

Wang et al. / J Zhejiang Univ Sci B 2008 9(10):835-841
836
(Dulymamode et al., 2001; Hyde, 2001).
There have been several studies on fungi in
poorly studied habitats in Thailand (Jones and Hyde,
2004). This has included studies on larger fungi in the
forests of northern Thailand (Le et al., 2007a; 2007b),
fungi on peat swamp palms (Pinnoi et al., 2006; Pin-
ruan et al., 2007), fungi on other monocotyledons
(Photita et al., 2001; Thongkantha et al., 2008), fungi
on leaf litter of angiosperms (Promputtha et al., 2004;
2005; Duong et al., 2006; 2008), and fungi on decay-
ing wood (Kodsueb et al., 2008a; 2008b). There has
been, however, no study on the litter of Ficus species.
Whether fungi are host-specific or generalists
are significant in estimating species numbers (Zhou
and Hyde, 2001; Hyde et al., 2007), is particularly
important in studying leaf litter. In tropical forests,
plant litter comprises a mixture of diverse host leaves,
unlike that in many temperate forests where stands
contain low tree diversity. Several studies on the
saprobic microfungal diversity associated with plants
have been carried out (Hyde et al., 2001; Yanna et al.,
2001; Dulymamode et al., 2001; Paulus et al., 2006a;
2006b), and to a large extent, fungi tend to be
host-specific at the host genus level (Taylor et al.,
2001). Paulus et al.(2006a; 2006b) studied 6 tree
hosts from 4 families and found very little overlap.
The low overlap may be because the hosts studied
were from different families. The next question to ask
is whether litter of different species of the same host
genus has host-specific fungi or generalists? We,
therefore, chose to study the saprobes on 5 species of
Ficus to establish data on the fungal communities
involved in the decay of litter from different species
of the same host genus.
The genus Ficus (Moraceae) includes some 750
species of woody plants occurring in most tropical
and subtropical forests throughout the world
(Weiblen, 2000). Ficus is an evergreen or deciduous
tree, frequently growing on other trees (strangling)
with smooth pale gray bark and abundant white latex
(Gardner et al., 2000). There is little known con-
cerning the fungal diversity on this host genus
(Suryanarayanan and Vijaykrishna, 2001; Paulus et
al., 2006a). Paulus et al.(2006a; 2006b) reported on
the fungal diversity in leaf litter of 2 species of Ficus
in Australia, while Suryanarayanan and Vijaykrishna
(2001) isolated 28 endophyte fungi from Ficus
benghalensis. Ficus is the second largest tree genus in
northern Thailand after Syzygium with 30 species of
Ficus growing in the region (Gardner et al., 2000).
The purpose of the present study is to assess the
diversity of saprobic fungi on 5 species of Ficus in
northern Thailand in order to (1) investigate fungal
distribution on saprobic fallen leaves of Ficus species;
and (2) evaluate the different fungal communities
involved in the decay of litter from different species
of the same host genus.
MATERIALS AND METHODS
Survey design
To examine the effect of host species on micro-
fungal assemblages, 5 species of Ficus were chosen
for the study. Individual trees of each species were
located at each of the sites selected for sampling
(Table 1). Thirty fallen leaves at various stages of
decay were haphazardly collected from each Ficus
species from each site within 2 d. All samples were
placed in Ziplock plastic bags containing tissue paper
moistened with sterile distilled water and sealed and
incubated at room temperature. Material was exam-
ined in the laboratory on Days 5, 15 and 25 after
sampling.
Assessment of microfungal diversity
For the assessment of fungal presence, all leaves
were cut into 5 cm×5 cm quadrates for observation.
Table 1 Location of Ficus species surveyed in Chiang Mai Province in northern Thailand
Plant sepcies Location
Ficus altissima MRC, T. Pa Pae, A. Mae Taeng, Chiang Mai Province, Thailand, 19°07′12″ N, 98°44′2.64″ E.
F. virens Out door of MRC, T. Pa Pae, A. Mae Taeng, Chiang Mai Province, Thailand, 19°07′12″ N, 98°44′2.64″ E.
F. benjamina About 5 km east of MRC, M. Mae Taeng, Chiang Mai Province, Thailand, 19°05′12″ N, 98°44′2.64″ E.
F. fistulosa Near Mae Malai, M. Mae Taeng, Chiang Mai Province, Thailand, 19°05′0″ N, 98°5′42″ E.
F. semicordata
Tung Cho National Park, forest trail, Tung Joaw Village, Mae Teng Dist., Chiang Mai Province, Thailand,
19°08′4.2″ N, 98°38′ 54″ E.

Wang et al. / J Zhejiang Univ Sci B 2008 9(10):835-841 837
Fruiting bodies of microfungi were observed under a
stereoscope. A slide was then prepared for one rep-
resentative fruiting structure of each taxon and placed
in water and observed under a compound microscope.
Micrographs were taken and stains were added as
appropriate. Slides were made semipermanent by the
addition of 90% (v/v) lactic acid. Species identifica-
tions were made using morphological characters.
Herbarium specimens and slides of all taxa are
maintained at the Mushroom Research Foundation
Herbarium with some duplicates in Zhejiang Univer-
sity Herbarium.
Definitions and statistical analyses
Taxa were recorded as either present or absent
from each leaf. The number of leaves, on which a
particular fungal species was observed, designated
the ‘occurrence of a fungus’ and was used to calculate
the ‘percentage occurrence’ of a taxon on leaves of
one tree species using the following formula (Tsui et
al., 2001; Yanna et al., 2002):
Percent occurrence of taxon A (%)=occurrence of
taxon A/occurrence of all taxa in one tree species×100.
Fungal species diversity on each Ficus species
was calculated using Shannon-Wiener’s index H:
H=−∑PilnPi,
where Pi is the frequency of fungal species i occurring
on specific host leaves (Begon et al., 1993; Wong and
Hyde, 2001).
Sørensen’s index of similarity (S) was applied
(Magurran, 1988; Tsui et al., 2001) to compare the
similarity of the species on leaves of different Ficus
spp.: S=2c/(a+b), where a is the total number of spe-
cies on host A leaves, b is the total number of species
on host B leaves, and c is the number of species on
both host leaves. Similarity is expressed with values
between 0 (no similarity) and 1 (absolute similarity).
RESULTS
Species richness and dominant fungi on Ficus spp.
Examination of decaying leaves of 5 species of
Ficus from northern Thailand yielded 80 fungal taxa,
comprising 56 anamorphic taxa, 23 ascomycetes and
1 basidiomycete. We have identified many taxa to
species level, although in several cases the mor-
phology of the taxa in our study was not identified to
species (unidentified species, Table 2). Five species,
Beltraniella nilgirica, Lasiodiplodia theobromae,
Ophioceras leptosporum, Periconia byssoides and
Septonema harknessi were common species occurring
on all five host species (>3% occurrence, Table 2). In
addition, Colletotrichum and Stachybotrys were
common genera, with several species occurring on the
Ficus leaves (Table 2). Figs.1~5 also show some rare
fungal taxa on the different host species.
(a) (b) (c)
Fig.1 Ellisiopsis gallesiae on fallen leaves of Ficus altis
-
s
ima. (a) Morphology of conidia; (b) and (c) Morpholog
y
of conidiophores. Bar=10 µm
(a) (b) (c) (d)
Fig.2 Morphology of Sagenomella striatispora on fallen
leaves of Ficus fistulosa. (a) Characters of conidiophore;
(b) Characters of conidia and conidiophore; (c) and (d)
developing conidia from conidiophores. Arrow indicates
the young conidium. Bar=10 µm
Fig.3 Morphology of Kirschsteiniothelia spp. on fallen
leaves of Ficus semicordata. (a) Characters of asci. Arrows
indicate the outer layer of the ascus wall; (b) Characters of
ascus; (c) Characters of pseudoparaphyses; (d) Charac-
ters of ascospores. (a) Bar=20 µm; (b)~(d) Bar=10 µm
(a)
(b) (c)
(d)

Wang et al. / J Zhejiang Univ Sci B 2008 9(10):835-841
838
Fig.4 Morphology of Stachybotrys bisbyi on fallen leaves
of Ficus benjamina. (a) Characters conidia. Arrow in-
dicates the smooth wall of the conidium; (b) Characters
of conidiophore. Arrow indicates the rough wall of co-
nidiophore cell; (c) and (d) Characters of conidiophores.
Bar=10 µm
(a)
(b)
(c)
(d)
Fig.5 Morphology of Stachybotrys renispora on fallen
leaves of Ficus virens. (a) Characters of conidiophore;
(b) and (c) Characters of conidia and conidiophores.
Arrow indicates the succession of conidium; (d) and (e)
Characters of conidia. Bar=10 µm
(a) (b)
(c)
(d)
(e)
Table 2 Percent occurrence (%) of fungal taxa on leaves of 5 species of Ficus in northern Thailand
Fungus name FV FF FA FS FB Fungus name FV FF FA FS FB
Ascomycetes Mitosporic fungi
Amphisphaeria fallax 2.3 Ellisiopsis gallesiae 2.3
Anthostomella limitata 2.2 2.4 Fusariella hughesii 4.4 4.7
Anthostomelle nannorrhopis 2.3 3.4 5.8 Fusariella obstipa 2.3
Chaetomium sulphureum 4.7 Fusarium chlamydosporum 4.8
Diaporthe adunca 4.7 Fusarium oxysporum 5.5
Diaporthe perjuncta 2.4 3.4 Fusarium solani 4.4
Eutypella scoparia 2.4 3.4 Giulia tenuis 1.7
Glomerella acutata Kellermania yuccifolia 2.4
Glomerella cingulata 1.1 1.2 Kontospora halophila 1.7
Guignardia spp. 1.1 2.4 1.2 2.9 Koorchaloma jamaicensis 2.4
Hyponectria populi 3.4 Lasiodiplodia theobromae 3.3 4.8 3.6 6.8 5.8
Kirschsteiniothelia spp. 1.7 Menispora ciliata 3.4
Lasiosphaeria breviseta 4.4 Menispora glauca 3.4
Lophiostoma minosporum 6.8 Microdochium phragmitis 4.8 4.7
Massarina hepaticarum 4.7 Mycoenterolobium platysporum 2.2 2.3
Melanospora brevirostris 3.4 Periconia byssoides 4.4 4.8 4.7 6.8 4.3
Munkovalsaria appendenta 1.1 1.7 1.4 Pestalotiopsis gigas 2.2 2.9
Mycoenterolobium platysporium 2.2 2.3 Pestalotiopsis owenii 4.4 2.9
Mycosphaerella dianthi 4.8 2.9 Phaeoisaria clematidis 2.3
Mycosphaerella freycineitae 4.4 3.4 Phoma eupyrena 5.8
Nectria sinopica 1.2 Phoma valerianae 2.2 2.4
Ophioceras leptosporum 4.4 4.8 4.7 6.8 5.8 Phomopsis asparagi 2.2 1.2 1.4
Rhytisma acerinum 3.4 Plectophomella nypae 2.2
Mitosporic fungi Sagenomella striatispora 2.4
Acremonium charticola 2.3 Schwarzmannia goebeliae 1.1
Ajrekarella polychaetriae 2.2 Septonema harknessi 4.4 4.8 4.7 5.1 5.8
Alternaria alternata 4.8 4.3 Sphaeridium candidum 2.4 2.9
Beltraina querna 1.2 Stachybotrys bisbyi 2.9
Beltraniella nilgirica 4.4 4.8 3.6 6.8 5.8 Stachybotrys nephrospora 2.9
Cercospora apii 2.2 2.9 Stachybotrys oenanthes 4.4 2.3 2.9
Chaetospermum artocarpi 4.8 Stachybotrys renispora 3.3
Chaetospermum chaetosporum 5.8Stachybotrys sansevieriae 2.3 3.4
Cladosporium cladosporioides 3.3 3.6 6.8 Torula graminis 5.8
Colletotrichum dematium 2.2 4.8 3.6 2.9 Torula herbarum 4.4
Colletotrichum gloeosporioides 3.6 Trichothecium roseum 2.4 3.6 3.4
Colletotrichum graminicola 4.7 Volutella ciliata 4.4 3.6 6.8
Cylindrocolla urticae 5.8Wiesneriomyces javanicus 2.2 4.8 3.4
Dicyma state of Ascotricha chartarum 2.2 2.3 Xylohypha nigrescens 4.7
Dinemasporium strigosum 2.2 4.8 2.9 Zygosporium chartarum 2.9
Discosia artocreas 2.4 Basidiomycete
Discosia brasiliensis 1.4Halocyphina spp. 1.2
FV: Ficus virens; FF: F. fistulosa; FA: F. altissima; FS: F. semicordata; FB: F. benjamina

Wang et al. / J Zhejiang Univ Sci B 2008 9(10):835-841 839
Fungal diversity
Communities of fungal taxa on different Ficus
species leaf litter were relatively distinct. The number
of taxa varied from 24 (F. semicordata) to 33 (F.
virens and F. altissima) (Table 3). The fungal com-
munities on F. virens and F. altissima were the most
similar but were different from those on F. semicor-
DISCUSSION
Studies of fungal diversity on Ficus have not
been carried out until recently, although this genus is
very common in forests throughout the world
(Weiblen, 2000; Paulus et al., 2006a; 2006b). Sury-
anarayanan and Vijaykrishna (2001) studied fungal
endophyte diversity on different host tissues of F.
benghalensis in India. They found little overlap be-
tween fungal endophyte composition on the leaves
and aerial roots. Paulus et al.(2006a; 2006b) studied
saprobic microfungi in leaf litter of Australia tropical
rainforests, including two Ficus species. They found
that fungal assemblages of F. destruens and F. pleu-
rocarpa grouped closely together. The overlapping
species were Beltraniella portoricensis, Beltrania cf.
concurvispora, Cylindrocladium coulhounii, Dic-
typchaeta cf. novae-guineensis, Cylindrosympodium
cryptocaryae, Idriella acerosa, Idriella lunata, Me-
nisporopsis theobromae, Ophiognomonia elasticae,
Parasypodiella elongate and Trichoderma viride. In
the present study, we investigated fungal diversity on
leaves of five Ficus species. Our results show that the
data (Table 4). The percentages of ascomycetes on F.
virens and F. altissima were similar (24% and 30%,
respectively). F. semicordata had the highest per-
centage of ascomycetes (46%) among the host species.
The percentages of anamorphic taxa on F. virens and
F. altissima were also similar.
overlap of fungal species was much higher than that
in previous studies on fallen leaves from different
hosts in different families. Similarity ranged from
28% to 45%. Some species, such as Ophioceras lep-
tosporum, Beltraniella nilgirica, Lasiodiplodia
theobromae, Periconia byssoides and Septonema
harknessi, were shared by all hosts with high occur-
rence, indicating that a ‘core fungal group’ might be
responsible for the decay of Ficus leaves (Wong and
Hyde, 2001). In this study some fungi are new records
on Ficus. Among these, many species are widespread,
such as Diaporthe adunca, Lasiosphaeria breviseta,
Colletotrichum gloeosporioides and Torula herbarum.
At least one species, Munkovalsaria appendenta, is a
new record for Thailand. Further studies are needed to
find more fungi on the leaves of Ficus, for our inves-
tigation was limited, with one tree of each host being
sampled once.
Some previous research results showed that the
hosts or tissues are important factors that influence
the fungal communities (Ho et al., 2001; Paulus et al.,
2003; Rambelli et al., 2004) and that some saprobes
might be host-specific (Hyde et al., 1997). Wong and
Table 3 Diversity of fungi on different Ficus species
Number of species
Ficus species Ascomycetes Mitosporic fungi Basidomycetes Total Index H
Ficus virens 8 25 33 3.3525
F. fistulosa 6 19 25 3.0887
F. altissima 10 22 1 33 3.3196
F. semicordata 11 13 24 3.0881
F. benjamina 5 21 26 3.1496
Table 4 Index of similarity among different Ficus species
Ficus virens F. fistulosa F. altissima F. semicordata F. benjamina
Ficus virens 0.41 0.45 0.35 0.44
F. fistulosa 0.31 0.41 0.43
F. altissima 0.35 0.31
F. semicordata 0.28
F. benjamina

Wang et al. / J Zhejiang Univ Sci B 2008 9(10):835-841
840
Hyde (2001) investigated fungal diversity on
Gramineae, and their results showed that fungal
composition on host species level varied. Studies on
microfungi of tree leaves in tropical regions sug-
gested that the fungal assemblages on different host
species were variable, and some other terms, such as
‘host exclusivity’, ‘host fidelity’, ‘host affinity’, ‘host
affinity’, ‘host preference’ and ‘host recurrence’,
were used to describe plant-fungal associations with
definitive host (Zhou and Hyde, 2001). Taylor et
al.(2001) noted that saprobic fungi were specific at a
host genus level. In the present study, our results
indicated that saprobic fungal diversity was high at
the host species level, with some rare fungal taxa
occurring on each of the different host species.
ACKNOWLEDGEMENT
We are grateful to Dr. Dhanasekaran Vijayk-
rishna, University of Hong Kong, China and Dr.
Ruilin Zhao, Southwest Forestry University, China
for technical help.
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