Endophytic fungi of wild banana (Musa acuminata) at Doi

Suthep Pui National Park, Thailand*

Wipornpan PHOTITA1†, Saisamorn LUMYONG1, Pipob LUMYONG2 and Kevin D. HYDE3

" Department of Biology, Faculty of Science, Chiang Mai University, Chiang Mai, Thailand 50200.
# Department of Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai, Thailand 50200.
$ Centre for Research in Fungal Diversity, Department of Ecology and Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong.
E-mail : wipornpan!hotmail.com

Received 27 September 2000 ; accepted 9 January 2001.

Endophytic fungi were isolated from 7500 samples of wild Musa acuminata collected from five sites at Doi Suthep Pui National Park,
Thailand during December 1998 to July 1999. Overall colonization rates from surface sterilized tissues were 56n5, 48n9, 48, 47n9 and
41n7 % for the Medicinal Plant Garden, Ban Suthep, Queen Sirikit Botanic Garden, San Gu, and Montatarn waterfall sites
respectively. Sixty-one different fungal taxa were isolated. Fewer isolates were recovered from younger than older samples.
Xylariaceous taxa and Guignardia cocoicola were the most frequently isolated endophytes from leaves and were either absent or rare
in midrib, petiole and pseudostem. Colletotrichum gloeosporioides, C. musae, Guignardia cocoicola, various sterile mycelia and
xylariaceous spp. were common at all sites. The endophyte fungal communities at the five sites were found to differ. Deightoniella
torulosa was the most frequent isolate at the Ban Suthep site and was either absent or rare at other sites. Colletotrichum species were
most common in the midribs and petioles at all sites, while Pyriculariopsis parasitica and Dactylaria sp. were most common in the
pseudostems. The endophyte communities isolated from M. acuminata in this study are compared with those from previous studies
on tropical hosts. Several of the endophytes isolated are established pathogens of banana and provide support for the hypothesis
that some endophytes are latent pathogens. The diversity of fungi on banana is discussed in relation to global estimates of numbers
of fungus species.

INTRODUCTION been introduced throughout the tropics and subtropics, as

The study of endophytes of tropical plants has received much
attention because endophytes are believed to be both diverse
and to provide an excellent potential source of biologically
active novel compounds (Dreyfuss & Petrini, 1984 ; Hyde,
2001). Studies in the tropics have included endophytes in
bamboo (Umali, Quimio & Hyde, 1999 ; Lumyong et al. 2000)
and palms (Rodrigues, 1994 ; Taylor, Hyde & Jones, 1999 ;
Fro$ hlich, Hyde & Petrini, 2000). The only previous study on
the endophytes of Musa sp. was conducted in Australia and
Hong Kong and was mostly confined to cultivated plants
(Brown, Hyde & Guest, 1998).
The Musaceae is a monocotyledonous family comprising
Musa and Ensete species. There are 37 species of Musa and
seven species of Ensete (Price, 1995). This family has its highest
species diversity in South East Asia with representatives
occurring across the Old World Tropics. Due to the
commercial and subsistence value of Musa species they have
* Paper presented at the Asian Mycological Congress 2000 (AMC 2000),
incorporating the 2nd Asia-Pacific Mycological Congress on Biodiversity and
Biotechnology, and held at the University of Hong Kong on 9–13 July 2000.
† Corresponding author.
well as in more marginal climates, such as those in Florida and
Israel. Most fungal disease research has focused on the
commercial cultivars.
The present study was initiated to investigate the ecology
and biodiversity of endophytic fungi in wild banana species at
Doi Suthep Pui National Park, Thailand. This research was
primarily motivated by the desire to establish whether the
endophytes in banana differed from those in other tropical
hosts. We also wanted to establish whether the endophytes in
banana were particularly diverse and whether there were any
evidence that at least some of the endophytes of banana are
latent pathogens.
MATERIALS AND METHODS
Sample collection
Healthy wild banana plants were sampled from four sites at
Doi Suthep Pui National Park at Ban Suthep (site I, alt. 350 m),
Montatarn waterfall (site II, alt. 600 m), Medicinal Plant
Garden (site III, alt. 950 m) and San Gu Region (site IV, alt.
1400 m) and one site at Queen Sirikit Botanic Garden (site V,
alt. 600 m). Samples were collected from each site every three
months from December 1998 to July 1999, so that in total 15
collections were made, i.e. three from each site. Five randomly
selected plants were sampled per site on each collection date.
Two pseudostem parts (outer and inner pseudostem) and two
leaves were removed from each plant : one old leaf, defined as
either the third or fourth most recently opened leaf, and one
young leaf, which was always the youngest opened leaf. The
samples were taken to the laboratory and processed within
24 h.
Isolation of endophytes
The leaves were first washed in running water. Leaf discs
(3 mm diam) were cut to include the vein (10 samples) and
intervein tissues (10 samples) from each leaf sample, using a
sterile cork borer. Ten segments (5i5i5 mm) were cut from
each of the petiole, midrib and two pseudostem samples using
a sterile razor blade. All leaf discs and segments were surface
sterilized in 75 % ethanol for 1 min, 1 % sodium hypochlorite
for 3 min and 95 % ethanol for 0n5 min and then dried in
sterilized paper. This protocol was found to be the optimum
triple sterilized procedure for isolating endophytes from M.
acuminata following a pilot experiment (Photita et al. 1999).
Five surface sterilized leaf discs and segments were evenly
spaced in Petri dishes (9 cm diam) containing 2 % (w\v) malt
extract agar (MEA) with added Rose Bengal (30 mg\l) to slow
down fungal growth and streptomycin sulfate (50 mg\l) to
suppress bacterial growth. Fungi growing out from the plant
tissues were transferred to test tubes containing corn meal
agar slants and incubated at room temperature ("25 mC) to
promote sporulation. Non-sporulating cultures were trans-
ferred to incubation chambers comprising a screw top glass jar
(5n5 cm diam x 10n5 cm high), containing 1 cm of solid potato
dextrose agar, and an 8 cm vertical strip of banana petiole. In
a second protocol the banana petiole to induce sporulation. In
this way it was possible to assign many of the sterile mycelia
to xylariaceous taxa.
Statistical analyses
The results from the three collections for each site are
combined for analysis. Overall colonization rates (CR) were
determined as described by Petrini, Stone & Carroll (1982).
E G
Total number of leaf
discs\petiole\midrib\
pseudostem segments
Colonization F
in a sample yielding  1 isolate H
l E G musae, xylariaceous taxa and Guignardia cocoicola were more
rate Total number of leaf discs\
petiole\midrib\pseudostem
F
segments in that sample H isolated from banana samples collected at Queen Sirikit
Relative frequency of isolation was used for species abundance
and calculated as the number of discs colonized by a given
fungus divided by the total number of discs infected, and
expressed as percentages. A Chi- squared (χ#) goodness-of-fit
test was performed to test whether the colonization rates of
five trials were statistically different. A Kruskal–Wallis test
was used for multisample analysis, such as the investigation of
the number of isolates recovered from different tissue types at
each site.
Identification
Sporulating isolates were identified at the genus and where
possible to species level. xylariaceous taxa produced stromata-
like structures, but did not sporulate. They were sorted into
different taxa based on stromatal and colony morphology.
Non-sporulating isolates were sorted into morphospecies on
the basis of colony surface texture, hyphal pigmentation,
exudates, and growth rates, as described by Fro$ hlich et al.
(2000).
Voucher slides and cultures are held in the Department of
Biology, Chang Mai University.
RESULTS AND DISCUSSION
Overall colonization
Sixty-one different fungal taxa were isolated from Musa
acuminata (Table 1). The most frequently isolated species were
xylariaceous taxa, Colletotrichum musae, C. gloeosporioides,
Guignardia cocoicola and various sterile mycelia (Table. 1).
The number of taxa isolated is similar to the 63 endophytic
taxa isolated from palms (Rodrigues, 1994), but higher than
many previous studies, e.g. Rodrigues & Samuels (1990), 12
taxa on palms, Pereira, Azovedo & Petrini (1993), 13 taxa on
Stylosanthes, Fisher et al. (1994), 23 taxa on Opuntia stricta,
Brown et al. (1998), 25 taxa on Musa sp. and Umali et al.
(1999), 37 taxa from bamboo leaves.
The overall colonization rates (CR) for the 5 sites did not
differ greatly. They were 56n5 % for Medicinal Plant Garden,
48n9 % for Ban Suthep, 48 % for Queen Sirikit Botanic Garden,
47n9 % for San Gu Region, and 41n7 %, for Montatarn
Waterfall (Fig. 1). These results were within the range of other
tropical endophyte studies. A great variation in colonization
rates has been reported for palm endophytes, from as low as
12n5 % (Rodrigues & Samuels, 1990) to as high as 80n8–89n2 %
(Fro$ hlich et al. 2000). In some studies, in which leaflets only
were examined, the colonization rates were 21–30 % on palms
(Rodrigues, 1994) and 20n3 % in palms (Southcott & Johnson,
1997). Colonization rates reported in a previous study on
banana endophytes were 29n6–67 % (Brown et al. 1998).
Site effect
The most common species isolated varied from one site to
another. Banana samples from Ban Suthep were most
frequently colonized by Deightoniella torulosa. Colletotrichum
frequently isolated from banana samples collected at Medicinal
Plant Garden. Cordana musae was the most common species
Botanic Garden. The most common species isolated by Brown
et al. (1998) from Musa sp., which is the only comparable
study, were Colletotrichum gloeosporioides, Pestalotiopsis pal-
marum and Nigrospora oryzae in Hong Kong and Colletotrichum
gloeosporioides, Glomerella cingulata and Phoma sp. in Australia.
Endophytic species abundance also varied according to the
site within each location. Several studies have shown that site
specific factors may influence the level of infection (Carroll,
1995). This variation is probably a reflection of the range of
Table 1. Relative frequency ( %) of endophytic fungi on young and old tissues of Musa acuminata at five sites.

Young Old
Taxa
Site I Site II Site III Site IV Site V Site I Site II Site III Site IV Site V

Ascomycete. sp. 10n7

Chaetomium sp. 1n3
Cladosporium sp. 0n7 2 6 6n7 12n7 4n7
Colletotrichum gloeosporioides 0n7 3n3 10n7 13n3 36n7 10n7
Colletotrichum musae 1n3 4n6 16n7 34n7 54n7 14n7 12
Cordana musae 0n7 21n3 3n3 2 63n3
Curvularia sp. 0n7 2n6 2 1n3 1n3
Dactylaria sp. 0n7 14n7 5n3 17n3
Deightoniella torulosa 0n7 65n3 1n3
Fusarium sp. 1 0n7 1n3 1n3 2n6 2 2n6 4 0n7
Fusarium sp. 2 1n3 2n6 0n7 0n7 1n3
Fusarium sp. 4 1n3 2n6 2 4 4
Guignardia cocoicola 2n6 4 20 40n7 14n7 9n3
Helminthosporium sp. 1n3 0n7
Hyphomycete sp.1 5n3 0n7 0n7 4 16 2 1n3 0n7 4n7 11n3
Hyphomycete sp.5 5n3 4 2n6 0n7
Hyphomycete sp.7 2n6
Hyphomycete sp.10 1n3
Nigrospora oryzae 0n7 0n7 4n7 2n6 1n3 0n7
Periconiella musae 4 0n7
Pestalotiopsis sp. 1n3 0n7 0n7 4
Phialophora sp. 17 10 2n6
Phoma sp. 1 4 0n7 2 10 9n3 5n3 8n7 4n7
Phoma sp. 2 4 0n7 0n7 0n7 2
Phomopsis sp. 1 2n6 1n3 10 2 8n7 2n6
Phomopsis sp. 2 1n3 1n3 0n7
Pyriculariopsis parasitica 0n7 2n6 6n7 2n6 8n7
Stachybotrys sp. 2 1n3 0n7 4n7 7n3 2n6 0n7
Sterile mycelium sp. 1 2 1n3 5n3 5n3 6 4n7 6n7 12
Sterile mycelium sp. 2 1n3 1n3 1n3
Sterile mycelium sp. 3 1n3 0n7 0n7 2 18n3 8 6 8
Sterile mycelium sp. 4 2
Sterile mycelium sp. 5 2 0n7 2 5n3 10n6 3n3 8 11n3 1n3
Sterile mycelium sp. 6 6n7 3n3 0n7 7n3 2
Sterile mycelium sp. 7 4
Sterile mycelium sp. 8 4n7 0n7
Sterile mycelium sp. 19 1n3 1n3
Verticillium sp. 1n3 0n7
Xylariaceous taxa 2 4 0n7 22n7 24 50n7 18n7 38n7

Rare isolates (less than 1 % relative frequency) in each site :

Site I (Young) : Deightoniella torulosa, Nigrospora oryzae and Constantiniella sp.
Site I (old) : Arthrinium sp., Drechslera sp., Glomerella sp., Dactylaria sp., Hyphomycete sp. 12, and 14
Site II (Young) : Colletotrichum gloeosporioides, Nigrospora oryzae, Hyphomycete sp. 1, and Sterile mycelium sp. 5
Site II (old) : Fusarium sp. 2, Drechslera sp., Pestalotiopsis sp., Phoma sp. 2, Verticillium sp., Sterile mycelium sp. 14, and 18
Site III (Young) : Fusarium sp. 1, Phoma sp. 1, Hyphomycete sp. 1, Cordana musae, and Verticillium sp.
Site III (old) : Fusarium sp. 2, Drechslera sp., Glomerella sp., Periconia digitata, Periconiella musae, Pestalotiopsis sp., Phoma sp. 2, Phomopsis
sp. 3, Sterile mycelium sp. 6, 9, 16, and 17, and Hyphomycete sp. 1
Site IV (Young) : Cladosporium sp., Curvularia sp., Fusarium sp. 3, xylariaceous spp., and Sterile mycelium sp. 3
Site IV (old) : Phoma sp. 2, Scytalidium sp., Sterile mycelium sp. 3, and Constantiniella sp.
Site V (Young) : Pyriculariopsis parasitica, Stachybotrys sp., Sterile mycelium sp. 3 and Constantiniella sp.
Site V (old) : Fusarium sp. 1, Glomerella sp., Nigrospora oryzae, Phoma sp. 2, Sagenoma sp., Stachybotrys sp., Sterile mycelium sp. 8, Hyphomycete sp. 5, and 6

environmental conditions under which the banana plants
grew. Differences in environmental factors included humidity,
temperature, rainfall and potential inoculum sources. For
example, Musa plants at sites I-IV are natural stands
underneath mixed deciduous forest, while site V is a wild
banana plantation in Queen Sirikit Botanic Garden. Site I is
located near to a village where there may be an effect due to
human activity. The canopy above sites II and IV is less dense
than at other sites and humidity and thus potential inoculum
is probably lower. Site III is deep in the forest where the
humidity is high (80–90 %) all year round, and is likely to have
a higher inoculum potential.
Age effect
There were significantly more endophyte isolates obtained
from old than from young banana tissues (P  0n05) (Fig. 2).
This result is in agreement with previous studies (Brown et al.
1998, Espinosa–Garcia & Langenheim 1990, Fisher et al. 1995,
Hata & Futai 1993, Rodrigues 1994, Taylor et al. 1999). With
 60 the exception of Ascomycete sp. 1, Chaetomium sp. and
Constantiniella sp. found in young samples, all fungal taxa
50
isolated were present on old samples. Most endophytes are
Colonization Rate

40 believed to enter a plant when a spore lands on a leaf surface

and grows into the plant through the stoma, or penetrates the
30 host directly (Fro$ hlich et al. 2000). The increased incidence of
20
endophyte taxa on the older leaves provide indirect evidence
for this because the older leaves would have had more time to
10 accumulate endophytes from the environment. Older leaves
would also have more time to accumulate vertically transmitted
0 colonisers that enter from the petiole (Fro$ hlich et al. 2000).
Site I Site II Site III Site IV Site V

Fig. 1. Overall colonization rates of endophytic fungi of Musa

acuminata in Doi Suthep Pui National Park.
Effect of the tissue type
Young With the exception of site III, the results of the Kruskal–Wallis
Old test indicate that there were no significant differences between
60 the numbers of isolates recovered from vein, intervein, midrib,
50
petiole and pseudostem tissues. In site III, the number of
isolates from the pseudostems were significantly lower (P l
Colonization Rate

40 0n032) than in other tissues. Bar graphs were used to compare

30
the colonisation rates of taxa against tissue types (Figs 3) and
were : petiole (60n8 %), pseudostem (57n6 %), midrib (49n5 %),
20 intervein (39 %), and vein (36n7 %), respectively.
There were, however, differences in the colonization rates
10
of taxa isolated from tissue types, which indicates that some
0 taxa have an affinity for different tissue types. Xylariaceous
Site I Site II Site III Site IV Site V
taxa and Guignardia cocoicola were the most frequently
Fig. 2. Colonization rates of endophytic fungi on young and old isolated from leaves. Pyriculariopsis parasitica and Dactylaria sp.
samples of Musa acuminata. were most common in the pseudostem. Colletotrichum musae
and C. gloeosporioides were most common in the midribs and
70
petioles (Table 2). These results are in agreement with many
studies that have found differences in species abundance
60 between plant tissues, such as bark, stems and leaves (Fisher
et al. 1994), between midrib and laminar tissue (Rodrigues
Colonization Rate

50
1994, Fisher et al. 1995, Brown et al. 1998).
40
Previous studies have indicated that endophytes may
30 exhibit tissue specificity (Luginbuhl & Mu$ ller, 1980 ; Rodrigues
20 & Samuels, 1990 ; Clay, 1992 ; Rodrigues, 1994 ; Fro$ hlich et al.
2000). Differences in endophyte assemblages in different
10
tissue types might be a reflection of tissue preferences of
0 individual dominating taxa (Luginbuhl & Mu$ ller, 1980 ;
Intervein Vein Midrib Petiole Pseudostem
Widler & Mu$ ller, 1984 ; Rodrigues & Samuels, 1990 ; von
Fig. 3. Colonization rates of endophytic fungi from Musa acuminata Halmschlager, Butin & Donaubauer, 1993), and might reflect
tissues. their capacity for utilizing or surviving within a specific
substrate (Rodrigues, 1994). The factors that may be important
Table 2. The most frequency isolated fungal endophytes from different in this respect, include the weathering of the leaf cuticle, tissue
tissues of Musa acuminata. texture and changes in the tissue physiology and chemistry
(Petrini & Carroll, 1981 ; Stone, 1987).
Tissue

Taxa Internal vein vein midrib petiole pseudostem

Xylariaceous fungi 23n6 21n4 11n4 10n9 6 Taxonomic composition of the endophytes
Deightoniella torulosa 6n3 2n3 3n6 10n2 6n6
Guignardia cocoicola 22n4 16n5 3n2 3n8 0n3 In this study we identified 61 taxa of endophytes from Musa
Cordana musae 5n9 9n4 11n4 4n7 0n8 acuminata in Doi Suthep Pui National Park. This comprised
Colletotrichum musae 6n7 10n7 4n3 10n7 8n6 eight identified species, 34 taxa identified to the generic level
C. gloeosporioides 6n3 10 12n5 13n2 6n1 and separated based on spore and colony morphology, five
Pyriculariopsis parasitica 0n4 0n3 0 0n3 8n8
xylariaceous taxa based on stromatal structure and colony
Dactylaria sp. 0n8 0 1n1 2 14n9
morphology and 14 sterile mycelia based on colony
morphology. The numbers of endophytic taxa found in this
study are similar to those found on other tropical hosts, e.g.
Livistona palm (Guo, Hyde & Liew, 1998), Licuala palm
(Fro$ hlich et al. 2000), bamboo (Umali et al. 1999 ; Lumyong et
al. 2000). However, as most endophytic species cannot be
identified to species level and often are only labeled as sterile
mycelia, it is hard to speculate whether many endophytes are
specific to an individual host.
Five of the species identified in this study, Deightoniella
torulosa, Cordana musae, Colletotrichum musae, Guignardia musae
and Pyriculariopsis parasitica, are only known from Musa sp.
(Photita et al. 2001a). Their presence as endophytes in banana
is interesting as they may be latent pathogens. There are 37
species of Musa and seven species of Ensete (Price, 1995) and
most of these banana species have yet to be studied for
saprobes or endophytes. Some of the taxa we identified from
M. acuminata are also likely to be specific at the family level.
Some of the endophytes isolated from healthy tissue in this
study are established pathogens of banana. Cordana musae
causes leaf blotch, while Deightoniella torulosa causes leaf spots
(Ellis, 1976). Guignardia musae causes freckle, Colletotrichum
gloeosporioides and C. musae also cause anthracnose of fruits
(Holliday, 1980). Brown et al. (1998) alluded to the possibility
that banana pathogens may spend part of their life as
endophytes. The results from this study are certainly
supportive of this, although it is not apparent whether all
strains isolated as endophytes are capable of producing
disease. Collections from diseased tissues of the same plants as
examined for endophytes in this study have revealed the
presence of several pathogens such as C. gloeosporioides, C.
musae and Deightoniella torulosa and saprobes such as Nigrospora
oryzae, Periconiella musae and Pyriculariopsis parasitica (Holliday,
1980). We have also artificially infected detached leaves of
Musa acuminata with endophytic strains of Deightoniella
torulosa in moist chambers and disease symptoms were
produced.
The percentage of colonies that did not sporulate (sterile
mycelia) identified is typical of other endophyte studies in the
tropics (Lodge, Fisher & Sutton, 1996 ; Brown et al. 1998 ;
Umali et al. 1998 ; Lumyong et al. 2000). Guo et al. (1998)
isolated mycelia from palm petioles in a conical flask and
obtained better sporulation, and were able to identify two
species that were saprobes of Livistona chinensis. The evidence
of Guo et al. (1998) suggest that some of the sterile mycelia
isolated in endophyte studies may be specific to that host or
host family. Guo, Hyde & Liew (2000) have also sequenced
representative sterile mycelia and identified some to genus,
some to family, and most to order. One group belonged in the
Xylariales, but were not xylariaceous species. There are many
xylariaceous-like species that occur on palms (e.g. Arecophila,
Capsulospora, Oxydothis), that are members of the Xylariales. It
would be interesting to find such taxa as endophytic sterile
mycelia on palms, especially if they are host specific. Although
we used the same incubation method (Guo et al. 2000), we
were only able to get xylariaceous taxa to produce stromata
in culture.
Brown et al. (1998) list 46 pathogens that are probably
unique to Musa species, while in this study five endophytes
were identified that are probably unique to Musa species.
Photita et al. (2001a) list 46 saprobes from Musa species in
Hong Kong and 6 of these are most probably unique to Musa
species. In a list of fungi on plants in the USA (Farr et al. 1989),
there are 47 fungi that occur on Musa species. As a
conservative estimate there are probably more than 200 fungi
that are presently known only from Musa species (Farr et al.
1989 ; Brown et al. 1998 ; Photita et al. 2001a, b). The ratio of
6 fungi to each plant which is used amongst other things to
estimate global fungal species numbers (Hawksworth, 1991),
therefore appears to hold for Musa species, with a ratio of
200 : 37 or 5n4 : 1.
A C K N O W L E D G E M E N TS
This research was supported by The Royal Golden Jubilee PhD Program
(4BCM41D1) and the Biodiversity Research and Training Program (BRT
142006). A. Nuangmek is thanked for help with collecting samples. The
Multiple Cropping Center, Faculty of Agriculture, Chiang Mai University, is
thanked for laboratory facilities.
REFERENCES
Brown, K. B., Hyde, K. D. & Guest, D. I. (1998) Preliminary studies on
endophytic fungal communities of Musa acuminata species complex in
Hong Kong and Australia. Fungal Diversity 1 : 27–51.
Carroll, G. C. (1995) Forest endophytes : pattern and process. Canadian Journal
of Botany 73(Suppl.) : S1316–S1324.
Clay, K. (1992) Fungal endophytes of plants : biological and chemical
diversity. Natural toxins 1 : 147–149.
Dreyfuss, M. & Petrini, O. (1984) Further investigations on the occurrence
and distribution of endophytic fungi in tropical plants. Botanica Helvetica
94 : 33–40.
Ellis, M. B. (1976) More Dematiaceous Hyphomycetes. Commonwealth My-
cological Institute, Kew.
Espinosa-Garcia, F. J. & Langenheim, J. H. (1990) The leaf fungal endophytic
community of a coastal redwood population–diversity and spatial patterns.
New Phythologist 116 : 89–98.
Farr, D. F., Bills, G. F., Chamuris, G. P. & Rossman, A. Y. (1989) Fungi on Plants
and Plant Products in the United States. American Phytopathological Society
Press, St Paul, MN.
Fisher, P. J., Petrini, O., Petrini, L. E. & Sutton, B. C. (1994) Fungal endophytes
from the leaves and twigs of Quercus ilex L. from England, Majorca and
Switzerland. New Phytologist 127 : 133–137.
Fisher, P. J., Petrini, L. E., Sutton, B. C. & Petrini, O. (1995) A study of fungal
endophytes in leaves, stems and root of Gynoxis oleifolia Muchler
(Compositae) from Ecuador. Nova Hedwigia 60 : 589–594.
Fro$ hlich, J., Hyde, K. D. & Petrini, O. (2000) Endophytic fungi associated with
palms. Mycological Research 104 : 1202–1212.
Guo, L. D., Hyde, K. D. & Liew, E. C. Y. (1998) A method to promote
sporulation in palm endophytic fungi. Fungal Diversity 1 : 109–113.
Guo, L. D., Hyde, K. D. & Liew, E. C. Y. (2000) Identification of endophytic
fungi from Livistona chinensis based on morphology and rDNA sequences.
New Phytologist 147 : 617–630.
Hata, K. & Futai, K. (1993) Effect of the needle aging on the total colonization
rates of Pinus thunbergii and Pinus densiflora needles. Journal of Japanese Forest
Society 75 : 338–341.
Hawksworth, D. L. (1991) The fungal dimension of biodiversity : magnitude,
significance, and conservation. Mycological Research 95 : 641–655.
Holliday, P. (1980) Fungus Disease of Tropical Crops. Cambridge, UK :
Cambridge University Press.
Hyde, K. D. (2001) Increasing the likelihood of novel compound discovery
from fungi. In Bio-Exploitation of Filamentous Fungi (S. B. Pointing & K. D.
Hyde, eds) : 77–91. [Fungal Diversity Research Series No. 6] Fungal
Diversity Press, Hong Kong.
Lodge, D. J., Fisher, P. J. & Sutton, B. C. (1996) Endophytic fungi of Manilkara
bidentata leaves in Puerto Rico. Mycologia 88 : 733–738.
Luginbuhl, M. & Mu$ ller, E. (1980) Endophytische Pilze den oberirdischen
rganen von 4 gemeinsam an glechen Standorten achsenden Pflanzen
(Buxus, Hedera, Ilex, Ruscus). Sydowia 33 : 185–209.
Lumyong, S., Thongantha, S., Lumyong, P. & Tomita, F. (2000) Endophytic
fungi from 13 bamboo species in Thailand. Biotechnology for Sustainable
Utilization of Biological Resources in the Tropics 14 : 96–101.
Pereira, J. O., Azevedo, J. L. & Petrini, O. (1993) Endophytic fungi of
Stylosanthes ; a first report. Mycologia 85 : 362–364.
Petrini, O. & Carroll, G. C. (1981) Endophytic fungi in foliage of some
Cupressaceae in Oregon. Canadian Journal of Botany 59 : 629–636.
Petrini, O., Stone, J. & Carroll, F. E. (1982) Endophytic fungi in evergreen
shrubs in western Oregon : A preliminary study. Canadian Journal of Botany
60 : 789–796.
Photita, W., Lumyong, S., Lumyong, P., Hyde, K. D. (1999). Surface
sterilization techniques for the isolation of endophytic fungi from Musa
acuminata (Banana). The 5th Asia Pacific Biochemical Engineering Conference
15–18 November 1999, Phuket, Thailand P-PB14 : 1–7.
Photita, W., Lumyong, S., Lumyong, P. & Hyde, K. D. (2001a) Fungi on Musa
acuminata in Hong Kong. Fungal Diversity 6 : 99–106.
Photita, W., Lumyong, S., Lumyong, P., Hyde, K. D. & McKenzie, E. H. C.
(2001b). Index of fungi described from Musaceae. Mycotaxon 80 : in press.
Price, N. S. (1995) The origin and development of banana and plantain
cultivation. In Bananas and Plantains (S. Gowen, ed.) : 317–381. Chapman
& Hall, London.
Rodrigues, K. F. (1994) The foliar fungal endophytes of the Amazonian palm
Euterpe oleracea. Mycologia 86 : 376–385.
Rodrigues, K. F. & Samuels, G. J. (1990) Preliminary study of endophytic
fungi in tropical palm. Mycological Research 94 : 827–830.
Southcott, K. A. & Johnson, J. A. (1997) Isolation of endophytes from two
species of palm from Bermuda. Canadian Journal of Microbiology 43 :
789–792.
Stone, J. K. (1987) Initiation and development of latent infection by Rhabdocline
parkeri on Douglas fir. Canadian Journal of Botany 65 : 2614–2621.
Taylor, J. E., Hyde, K. D. & Jones, E. B. G. (1999) Endophytic fungi associated
with the temperate palm, Trachycarpus fortunei, within and outside its
natural geographic range. New Phytologist 142 : 335–346.
Umali, T. E., Quimio, T. H. & Hyde, K. D. (1999) Endophytic fungi in leaves
of Bambusa tuldoides. Fungal Science 14 : 11–18.
von Halmschlager, E., Butin, H. & Donaubauer, E. (1993) Endophytische pilze
in bla$ ttern und zweigen von Quercus pertraea. European Journal of Forest
Pathology 23 : 51–63.
Widler, B. & Mu$ ller, E. (1984) Untersuchungen u$ ber endophytische Pilze von
Arctostaphylos uva-ursi (L.) Sprengel (Ericaceae). Botanica Helvetica 94 :
307–337.
Corresponding Editor : K. D. Hyde