Distribution of fungi on wood in the Tutong River, Brunei

Abstract

Five sites along the Tutong river and its tributary, the Sungai Kelakas, were sampled for fungi on decaying wood. Sixty-six taxa were found including two new genera and two new species. Nearly half of these (48%) are new records for Brunei. Although common taxa were generally not unique to any particular site, species distributions were correlated to the salinity gradient. These findings are consistent with previous studies showing that some marine fungi are tolerant to less saline conditions and that some freshwater species are tolerant to more saline conditions. Some species could be potentially classified as brackish water species. The ratio of ascomycetes to anamorphic fungi was higher in marine than freshwater sites. A checklist of fungi found in aquatic habitats in Brunei is included.

Full text

Fungal Diversity
17
Distribution of fungi on wood in the Tutong River, Brunei
Sally C. Fryar1*, Webber Booth2, Johnathon Davies2, I. John Hodgkiss1
and Kevin D. Hyde1
1Centre for Research in Fungal Diversity, Department of Ecology & Biodiversity, The
University of Hong Kong, Pokfulam Road, Hong Kong SAR, PR China
2School of Biological Sciences, Universiti Brunei Darussalam, Brunei Darussalam
Fryar, S.C., Booth, W., Davies, J., Hodgkiss, I.J. and Hyde, K.D. (2004). Distribution of fungi
on wood in the Tutong River, Brunei. Fungal Diversity 17: 17-38.
Five sites along the Tutong river and its tributary, the Sungai Kelakas, were sampled for fungi
on decaying wood. Sixty-six taxa were found including two new genera and two new species.
Nearly half of these (48%) are new records for Brunei. Although common taxa were generally
not unique to any particular site, species distributions were correlated to the salinity gradient.
These findings are consistent with previous studies showing that some marine fungi are
tolerant to less saline conditions and that some freshwater species are tolerant to more saline
conditions. Some species could be potentially classified as brackish water species. The ratio of
ascomycetes to anamorphic fungi was higher in marine than freshwater sites. A checklist of
fungi found in aquatic habitats in Brunei is included.
Key words: brackish, freshwater fungi, fungal diversity, marine fungi, salinity, wood.
Introduction
Biological diversity, especially in the tropics is under threat (Wilson,
1988). It is essential that the range of species present in different habitats be
recorded rapidly for several reasons: (i) destruction of habitats is so rapid that
species may be either locally or globally extinct before we discover them; (ii)
many species contain medicinally important chemicals vital for humanity
(Wilson, 1988); (iii) records of species within geographical areas is essential
for environmental decision making (Hyde, 2003). Fungi are particularly poorly
studied throughout the tropics and substantial funding, training and study are
required to address this imbalance (Hawksworth, 2003; Hyde, 2003).
The diversity of fungi has been estimated to be 1.5 million (Hawksworth,
1991, 2001; Hyde, 2001). Although this estimate has been contested (May,
1994), there are approximately 72,000 fungal species currently described and
*Corresponding author: Sally Fryar: e-mail: scfryar@bigpond.com - present address, 23 Onyx
Rd, Artarmon NSW 2064, Australia

18
about 1600 are currently being described each year as new (Hawksworth,
2001). This high species richness of fungi means that they are an important
part of the biodiversity of an area.
The diversity of fungi in aquatic environments has been shown to be high
(Goh and Hyde, 1996; Tsui et al., 2000; Cai et al., 2002, 2003; Kane et al.,
2002; Luo et al., 2004), with more than 1000 species already discovered, most
of which are either ascomycetes or anamorphic fungi (e.g. Goh, 1997; Hyde et
al., 1997; Tsui and Hyde, 2003). The proportion of these species that actually
grow and reproduce within the aquatic habitat is, however, unknown (Shearer,
1993).
Fungi utilise many different substrates within the aquatic environment
including wood (Shearer and Von Bodman, 1983; Abdel-Raheem and Shearer,
2002; Bucher et al., 2004), leaves (Premdas, 1991), algae (Haythorn et al.,
1980), coral (Morrison-Gardiner, 2002), soil (Mer et al., 1980), insects
(Williams and Lichtwardt, 1990; Cafaro, 2002), and various other substrates
(Czeczuga, 1996; Czeczuga and Muszyńska, 2004). Their spores even occur in
tree holes and gutters (Gönczöl and Révay, 2003, 2004). There is often a
different assemblage of fungi on each substrate, although some generalists
utilise more than one substrate.
The distribution of wood-inhabiting fungi in aquatic habitats is affected
by salinity and temperature (Shearer, 1972; Hyde and Lee, 1995; Jones, 2000;
Tsui and Hyde, 2004). Jones and Oliver (1964) found that the assemblage of
fungi in seawater was very different to that found in brackish and fresh water.
However, several species were common to both fresh and brackish water.
There have been many studies recording the distribution and abundance of
fungi found in aquatic habitats (Hyde et al., 2000). Besides the study of Tsui
and Hyde (2004) the distribution of fungi from fresh through to seawater in one
system has not been studied.
In Brunei, nine studies on fungal diversity in aquatic habitats have
revealed distinct assemblages of fungi in freshwater and marine habitats (Hyde,
1988a,b, 1989, 1990a,b, 1991, 1992; Wong, 1996; Ho et al., 2002). However,
none of these studies examined fungi along a gradient from freshwater to
marine habitats. It was our intention in this study to determine the pattern of
fungal species abundance and distribution along a salinity gradient. The aim
was to determine if there was a distinct assemblage of fungi in brackish sites or
if there was simply a mix of freshwater and marine species.

Fungal Diversity
19
Fig. 1. Position of Brunei. a. Map of the study area; b. showing the Tutong river and its
tributaries.
Materials and methods
Study area
The Tutong River in Brunei feeds into the South China Sea on the
northwest side of Borneo (Fig. 1). Samples were collected from five sites along
the Tutong River and one of its tributaries, the Sungai Kelakas. Data on the
physico-chemical conditions of the sites were collected during field sampling
(Table 1). There was generally a gradient of salinity from 0 to 32‰, but at
times of high river flow, all sites had salinity that was effectively 0‰.
5km
South China Sea
Marine
Brackish 1
Brackish 2
Fresh 1
Fresh 2
N
= major road

20
Table 1. Physical properties of the water at each of the sites.
Marine Brackish A Brackish B Fresh water Peat swamp
Salinity (‰) 0-31 0-11.9 0-10 0-3 0
pH 4.8-5.6 3.1-6.0 6.3 5.4 3.1-4.3
Temperature 26.1-30 26.4-30 29 26.6-29 29
DO2 (%) 16-77 18-92 38 78 8-85
Marine site
The Tutong River at this site was approximately 50 m wide and 8.5 m
deep at the mid-point. The riparian vegetation at this point was dominated by
Sonneratia caseolaris, Avicennia alba and Nypa fruticans. The tidal range was
approximately 2 m.
Brackish site A
Sungai Kelakas at this point was approximately 20 m wide and 2 m deep
in the deepest part. The water was also tidal. A riparian strip approximately
50 m in width, bordered the river. The strip was inundated in times of high
river flow (when the water was effectively fresh) and during high tides. The
vegetation was a mixture of Barringtonia cf. racemosa, Bruguiera
gymnorrhiza, Cerbera odallam, Ficus cf. microcarpa, Gluta velutina, Heritiera
globosa, Hibiscus tiliaceus, Nypa fruticans and Sonneratia caseolaris.
Brackish site B
This site was 100 m upstream of Brackish site A, and had very similar
vegetation. However, the topography of the site was more that of a floodplain.
The channel at this point was less distinct and not as deep as at Brackish site A
(approximately 1 m maximum). Plant growth was not restricted to the
riverbanks, and some plants grew in the middle of the stream. Water at this site
was also tidal.
Freshwater site
This area was a floodplain with no defined channel. Maximum depth was
approximately 1 m. The riverine vegetation was dominated by species
associated with disturbance such as Dillenia suffruticosa, Ischaemum
barbatum, Scleria sp. and Vitex pubescens together with some regenerating
peat swamp species.

Fungal Diversity
21
Peat swamp forest (PSF)
Slightly further upstream from the freshwater site was a floodplain with a
peat swamp forest. The forest was on shallow peat dominated by
Campnosperma coriaceum, Lophopetalum multinervium and Syzygium sp. In
times of high rainfall, the area was inundated with water up to approximately
1.5 m. During drier periods, the ground was exposed, but there was a series of
pools throughout the forest. This site was referred to as a freshwater site.
Collection of samples
Thirty wood samples (approximately 40 × 5 × 3 cm) were randomly
collected from the edge of the river (up to 50 cm deep) at each site. Samples
were not collected if they were buried in the mud (and were therefore
anaerobic), or had recently fallen into the water. Abiotic measurements, pH,
salinity, temperature and dissolved oxygen (DO2), of the water at each site
were taken over two years. Samples were briefly washed in the river to remove
excess mud and algae and placed individually into zip-lock bags and incubated
at 27ºC in 24 hour light/dark cycles. After two weeks, the samples were
observed at 200× magnification with a dissecting microscope for fungal
fruiting bodies on the surface of the wood. Samples were observed again after
3 months of incubation. Fruiting bodies were identified at 400× magnification
using a compound microscope, and the fungus isolated onto artificial agar
media. Abundance was measured as presence/absence on a sample. That is, if a
fruiting body of one species was seen more than once on a sample, it was
counted only once.
Statistical analysis
Species abundance patterns
Shannon-Weiner diversity and evenness was calculated for each site
along with Margalef species richness. Calculations were carried out according
to Magurran (1988).
Multivariate analysis
Detrended Correspondence Analysis (DCA) was used to detect
differences in species abundance patterns between sites. MultiVariate
Statistical Package (MVSP) (www.kovcomp.com/mvsp) was used for this
analysis.

22
Results
Sixty-six taxa were found during the survey (Table 2), 43 were
ascomycetes and 23 anamorphic fungi. The most common species were
Cancellidium applanatum (N = 93), Sungaiicola brachydesmiella (N = 38),
Aniptodera chesapeakensis (N = 19), Papulospora sp. 2 (N = 18) and
Papulospora sp. 1 (N = 17). Cancellidium applanatum was common at all
sites, but was slightly more common in fresh and brackish water sites.
Sungaiicola brachydesmiella occurred in all habitats, but was more common in
brackish and marine sites. Aniptodera chesapeakensis occurred in brackish and
marine sites, but not in fresh water. Both Papulospora sp. 1 and sp. 2 occurred
only in brackish and marine sites. In fresh water no particular species were
very common (apart from Cancellidium applanatum which was common in all
sites).
Alysidium sp., Annulatascus triseptatus, Aquaticola ellipsoidea,
Ascomycete sp. 1, Ascotaiwania sp., Beverwykella pulmonaria,
Brachysporiella gayana, Cataractispora applanatum, Cataractispora viscosa,
Chaetosphaeria sp., Exserticlava vasiformis, Glomerella sp., Massarina sp.,
and Sporidesmium crassisporum were all unique to freshwater. However, apart
from Beverwykella pulmonaria and Brachysporiella gayana many of these
species were rare (N = 1, 2 or 3) and so may also be rare in the other sites, but
not observed during this study.
Eleven species were unique to brackish water sites: Aniptodera sp. 1,
Aniptodera cf. haispora, Annulatascus lacicola, Annulatascus palmietensis,
Aquaticola sp., Dactylospora mangrovei, Mangrovispora pemphii, Monodictys
sp. 1, Monodicyts sp. 2, Savoryella verrucosa and Trichocladium
achrasporum.
Eleven species were unique to the marine site: Aniptodera megalospora,
Ceratosphaeria sp. 1, Chaetosphaeriaceae sp., Lasiosphaeria sp. 3,
Lophiostoma bipolare, L. frondisubmersum, Phomatospora sp., Savoryella
lignicola, S. paucispora, Swampomyces sp. and Chalara sp.
Only seven species occurred in all habitats: Annulatascus velatisporus,
Aquaticola longicolla, Fluviatispora reticulata, Lasiosphaeria sp. 1,
Sungaiicola brachydesmiella, Cancellidium applanatum and Sporidesmium cf.
anglicum.
With the exception of Beverwykella pulmonaria and Brachysporiella
gayana, no species that occurred more than three times were unique to any one
habitat. Species that were common in any one habitat were either common in
the other habitats, or at least occurred in them.

Fungal Diversity
23
Table 2. Fungal species found at each of five sites. Numbers refer to the number of wood
pieces (maximum 30) on which each fungus occurred.
Site Fresh PSF Brackish
A
Brackish
B
Marine
Collection period 1 2 1 2 1 1 2
Ascomycetes
Unidentified Ascomycete sp. 1
Aniptodera sp. 1
Aniptodera sp. 2
Aniptodera chesapeakensis Shearer & M.A. Mill.
Aniptodera cf. haispora Vrijmoed, K.D.Hyde &
E.B.G. Jones
Aniptodera megalospora K.D. Hyde, W.H. Ho &
K.M. Tsui
Annulatascus sp.
Annulatascus palmietensis K.D. Hyde, Goh & T.D.
Steinke
Annulatascus triseptatus S.W. Wong, K.D. Hyde &
E.B.G. Jones
Annulatascus velatisporus K.D. Hyde
Aquaticola sp.
Aquaticola ellipsoidea W.H. Ho, K.M. Tsui,
Hodgkiss & K.D. Hyde
Aquaticola longicolla W.H. Ho, K.M. Tsui,
Hodgkiss & K.D. Hyde
Ayria appendiculata Fryar & K.D. Hyde
Ascotaiwania sp.
Cataractispora appendiculata K.D. Hyde, S.W.
Wong & E.B.G. Jones
Cataractispora viscosa K.D. Hyde, S.W. Wong &
E.B.G. Jones
Ceratosphaeria sp. 1
Ceratosphaeria sp. 2
Chaetosphaericeae sp.
Chaetosphaeria sp.
Dactylospora mangrovei E.B.G. Jones, Alias,
Abdel-Wahab & S.Y. Hsieh
Fluviatispora reticulata K.D. Hyde
Glomerella sp.
Lasiosphaeria sp. 1
Lasiosphaeria sp. 2
Lasiosphaeria sp. 3
Lophiostoma bipolare (K.D. Hyde) Liew, Aptroot &
K.D. Hyde
Lophiostoma frondisubmersum (K.D. Hyde) Liew,
Aptroot & K.D. Hyde
1
-
-
-
-
-
-
-
-
2
-
1
-
-
1
-
-
-
-
1
-
3
-
1
-
-
-
-
-
-
-
-
-
-
-
-
3
-
-
1
3
-
-
1
1
-
-
-
-
2
1
-
-
-
-
-
-
-
2
1
1
-
1
-
-
1
1
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
-
-
-
-
1
-
-
-
-
-
-
-
1
1
-
2
1
-
-
-
-
1
-
1
-
-
-
-
-
4
-
-
-
3
-
-
-
-
-
-
-
1
-
-
1
-
-
-
-
-
1
14
-
-
-
-
-
-
-
-
-
1
-
-
-
-
2
-
-
-
-
1
-
-
-
-
-
-
-
3
-
4
-
-
-
3
-
-
1
-
-
-
-
1
-
-
-
1
-
-
1
1
1
1
N = total number of individuals, S = total number of species.

24
Table 2 continued. Fungal species found at each of five sites. Numbers refer to the number of
wood pieces (maximum 30) on which each fungus occurred.
Site Fresh PSF Brackish
A
Brackish
B
Marine
Collection period 1 2 1 2 1 1 2
Lulworthia spp.
Mangrovispora pemphii K.D. Hyde & Nakagiri
Massarina sp.
Orbilia sp.
Phaeosphaeria capensis T.D. Steinke & K.D. Hyde
Phomatospora sp.
Savoryella lignicola E.B.G Jones & R.A. Eaton
Savoryella paucispora (Cribb & Cribb) Jørgen
Koch
Savoryella verrucosa Minoura & Muroi
Sungaiicola brachydesmiella Fryar & K.D. Hyde
Swampomyces sp.
Torrentispora crassiparietis Fryar & K.D. Hyde
Torrentispora fibrosa K.D. Hyde, W.H. Ho, E.B.G.
Jones, K.M. Tsui & S.W. Wong
Torrentispora fusiformis Fryar & K.D. Hyde
Number of individuals of ascomycetes
Number of species of ascomycetes
Anamorphic fungi
Alysidium sp.
Berkleasmium sp.
Beverwykella pulmonaria (Beverw.) Tubaki
Brachysporiella gayana Bat.
Cancellidium applanatum Tubaki
Candelabrum sp.
Chalara sp.
Dactylella sp.
Exserticlava vasiformis (Matsush.) S. Hughes
Monodictys pelagica (T.W. Johnson) E.B.G. Jones
Monodictys sp. 1
Monodictys sp. 2
Papulospora sp. 1
Papulospora sp. 2
Phaeoisaria clematidis (Fuckel) Hughes
Pleurophragmium-like
Sporidesmium cf. anglicum (Grove) M.B. Ellis
Sporidesmium crassisporum M.B. Ellis
Trichocladium achrasporum (Meyers & Moore)
Dixon
Tritirachium sp.
-
-
1
-
1
-
-
-
-
1
-
-
-
3
16
11
1
-
7
7
22
1
-
6
1
1
-
-
-
-
1
6
1
2
-
2
-
-
-
3
-
-
-
-
-
1
-
2
3
-
21
11
-
2
-
-
18
-
-
-
-
2
-
-
-
-
5
-
-
-
-
-
1
1
-
-
-
-
-
-
15
-
-
-
-
25
10
-
-
-
-
20
-
-
-
-
1
1
-
-
7
1
-
1
-
-
-
1
-
-
-
-
-
-
-
-
-
-
-
1
1
10
9
-
3
-
-
6
-
-
-
-
3
-
1
6
3
1
-
1
-
1
-
-
-
-
1
1
-
-
-
1
2
-
1
-
-
15
11
-
-
-
-
15
1
-
1
-
-
-
-
1
-
1
1
-
-
-
1
3
-
-
-
-
-
3
1
-
18
1
-
-
-
47
10
-
-
-
-
6
-
1
-
-
-
-
-
7
7
-
-
-
-
-
-
-
-
-
-
-
1
-
-
-
1
-
-
-
-
19
12
-
-
-
-
6
-
-
-
-
-
-
-
3
1
-
-
3
-
-
-

Fungal Diversity
25
Table 2 continued. Fungal species found at each of five sites. Numbers refer to the number of
wood pieces (maximum 30) on which each fungus occurred.
Site Fresh PSF Brackish
A
Brackish
B
Marine
Collection time 1 2 1 2 1 1 2
Xylomyces sp.
Xylomyces chlamydosporis Goos, R.D. Brooks &
Lamore
Xylomyces giganteus Goh, W.H. Ho, K.D. Hyde &
K.M. Tsui
Number of individuals of anamorphic fungi
Number of species of anamorphic fungi
-
1
-
59
14
-
-
-
27
4
2
1
4
38
9
-
-
-
21
9
-
-
-
21
7
1
-
1
23
6
-
-
-
13
4
Total no. of individuals 75 48 63 31 36 70 32
Total no. of species 25 15 19 18 18 16 16
Several species demonstrated a gradient in abundance between habitats.
During the first sampling Aniptodera chesapeakensis was common in the
marine site, rare in the brackish sites and did not occur in the freshwater site.
Similarly, during the first sampling Sungaiicola brachydesmiella was common
at the marine site and the brackish site closest to the sea and rare in the other
brackish site and freshwater site. This pattern was however not observed in the
second sampling.
No basidiomycete fruiting bodies were seen on the samples. However
some samples had thick white hyphal cords that may have been
basidiomycetous.
The ratio of ascomycetes to anamorphic fungi increased with increasing
salinity in the first set of samples (both number of species and number of
individuals) (Table 3). However, in the second sample, the ratio was lowest for
brackish water, and highest for sea water.
New species and genera
During this study two new genera (Ayria and Sungaiicola) and two new
species (Torrentispora crassiparietis and T. fusiformis) were discovered and
are presently being described. Several other species were also new (e.g.
Aniptodera sp. 1, and Aquaticola sp.) However, there was insufficient material
to describe these as new. The material has been lodged in the HKU(M).

26
Table 3. Descriptive statistics for each of the samples.
Measure Fresh PSF Brackish A Brac B Marine
Sampling time 1 2 1 2 1 1 2
No. of species (S)
No. of individuals (N)
Shannon-Weiner index (H’)
H’ evenness
Species richness (Margalef)
No. species per branch
No. branches with no fungi
No. ascomycetes/No. anamorphic
No. of species of ascomycetes/
anamorphic
25
60
1.29 ±
0.05
0.925
12.8
2.5 ±
1.5
2
0.3
0.8
15
46
0.99 ±
0.07
0.818
8.92
1.6 ±
1.2
6
0.8
1.8
19
63
0.90 ±
0.06
0.717
9.76
2.1 ±
1.5
5
0.7
1.1
18
32
1.14 ±
0.06
0.906
11
1.2 ±
0.9
8
0.5
1
18
36
1.01 ±
0.09
0.789
10.8
1.2 ±
1.1
7
0.7
1.6
16
70
0.96 ±
0.05
0.799
7.93
2.3 ±
1.6
6
2
1.7
16
32
1.11 ±
0.05
0.918
9.97
1.1 ±
1.3
14
1.5
3
New records for Brunei
Thirty-two species found during this survey are new records for Brunei.
A checklist of fungi found in aquatic habitats in Brunei in presented in Table 4.
The 246 fungal taxa consists of 158 ascomycetes, 3 basidiomycetes and 85
anamorphic taxa.
Species diversity
Generally more species and individuals were observed in the first set of
samples (Table 3) and overall, species diversity (H’) was highest at the
freshwater site during the first sampling (individual t-tests p<0.05). Although
the species richness and evenness of the samples canot be compared
statistically, the higher species diversity in the freshwater site appeared to be
due to both higher species richness and evenness at the site (Table 3).
The high species richness of the freshwater site was not as distinctive,
however, when the number of species per samples was observed. The
freshwater sample along with the Brackish A (1) and Marine (1) samples all
had more species than the other samples (ANOVA, F6,203=5.96, p<0.001; LSD,
p<0.05).
An average of 1.7 species were found on each branch in this survey.
However this number was highly variable (s.d. = 1.4). Many samples (23%)
had no fruiting bodies and some were found with up to 6 taxa.

Fungal Diversity
27
Table 4. Checklist of fungi from aquatic habitats in Brunei.
Species Habitat Reference
Ascomycetes
Acrocordia-like sp.
Acrocordiopsis patilii Borse & K.D. Hyde
Aigialus grandis Kohlm & Schatz
Aigialus mangrovei Borse
Aigialus parvus Schatz & Kohlm.
Aniptodera sp.
*Aniptodera new species
Aniptodera sp. (spores <15 µm)
Aniptodera chesapeakensis Shearer & M.A. Mill.
*A. cf. haispora Vrijmoed, K.D. Hyde & E.B.G. Jones
A. inflatiascigera K.M. Tsui, K.D. Hyde & Hodgkiss
A. lignicola K.D. Hyde, W.H. Ho & K.M. Tsui
A. longispora K.D. Hyde
A. mangrovii K.D. Hyde
*A. megalospora K.D. Hyde, W.H. Ho & K.M. Tsui
*Annulatascus sp.
Annulatascus bipolaris K.D. Hyde
A. triseptatus S.W. Wong, K.D. Hyde & E.B.G. Jones
A. velatisporus K.D. Hyde
Antennospora quadricornuta (Cribb & J. Cribb) T.W. Johnson
*Anthostomella sp.
Aquaticola sp. 1
Aquaticola sp. 2
*Aquaticola ellipsoidea W.H. Ho, K.M. Tsui, Hodgkiss & K.D.
Hyde
*A. longicolla W.H. Ho, K.M. Tsui, Hodgkiss & K.D. Hyde
Arenariomyces trifurcatus Höhnk
*Ayria appendiculata Fryar & K.D. Hyde
Ascocratera manglicola Kohlm.
*Ascotaiwania sp.
+Ascotaiwania pallida K.D. Hyde & Goh.
Bathyascus grandisporus K.D. Hyde & Jones
Belizeana tuberculata Kohlm. & Volkmann-Kohlm.
Biatriospora marina K.D. Hyde & Borse
Biconiosporella corniculata Schaumann
Bionectria sp.
+Bruneiapiospora sp.
Capillataspora corticola K.D. Hyde
Carbosphaerella leptosphaerioides I. Schmidt
Caryospora mangrovei K.D. Hyde
Caryosporella rhizophorae Kohlm.
Cataractispora appendiculata K.D. Hyde, S.W. Wong & E.B.G.
Jones
*C. viscosa K.D. Hyde, S.W. Wong & E.B.G. Jones
M
M
M
M
M
M
B
M
F,B,M
B
F
F
M
M
F,M
B
F
F
F,B,M
M
B
F
F
F
F,B,M,R
M
M
M
F
R
M
M
M
M
F
R
M
M
M
M
F
F
1,2,3
4,5
1,2,3,4,5
6,5
1,2,3,4,5,7
1,2,3,4
10
1
1,2,3,5,6,9,10
10
9
9
5
1,2,3,4,5,6,7
10
10
9
8,10
8,9,10,11
1,2,3,4
11
9
9
10
10,11
2,3
10
5
10
11
1,2,4,5
2,3,4,5
2,5
2,3
9
11
6
2,3
5
1,2,4,5
8,10
10

28
Table 4 continued. Checklist of fungi from aquatic habitats in Brunei.
Species Habitat Reference
*Ceratosphaeria sp.
+Chaetosphaeria sp.
Chaetosphaeria anglica P.J. Fisher & Petrini
Clohiesia corticola K.D. Hyde
Corollospora colossa Nakagiri & Tokura
C. maritima Werdermann
C. pulchella Kohlm., I. Schmidt & Nair
Crinigera mangrovei K.D. Hyde & E.B.G. Jones
C. maritima I.Schmidt
Cucullospora mangrovei K.D. Hyde & E.B.G. Jones
Cytospora rhizophorae Kohlm. & E. Kohlm.
Dactylospora haliotrepha (Kohlm. & Kohlm.) Hafellner
*D. mangrovei E.B.G. Jones, Alias, Abdel-Wahab & S.Y. Hsieh
Didymosphaeria sp.
Didymosphaeria enalia Kohlm.
Didymella avicenniae Patil & Borse
Diluviocola capensis K.D. Hyde, S.W. Wong & E.B.G. Jones
Etheirophora blepharospora (Kohlm. & E. Kohlm.) Kohlm. &
Volkm.-Kohlm.
Eutypa sp.
*Fluviatispora boothii Fryar & K.D. Hyde
*F. reticulata K.D. Hyde
*Glomerella sp.
Gnomoniella rubicola Pass.
Haligena salina Farrant & E.B.G. Jones
Halocyphina villosa Kohlm. & E. Kohlm.
Halosarpheia abonnis Kohlm.
H. cincinnatula Shearer & Crane
H. fibrosa Kohlm. & E. Kohlm.
H. heteroguttulata K.D. Hyde, S.W. Wong & E.B.G. Jones
H. marina (Cribb & J. Cribb) Kohlm.
H. minuta Leong
H. ratnagiriensis Patil & Borse
H. retorquens Shearer & Crane
H. viscosa I. Schmidt
Halosphaeria appendiculata-like
H. cucullata (Kohlm.) Kohlm.
H. salina (Meyers) Kohlm.
Helicascus kanaloanus Kohlm.
Hydronectria tethys Kohlm. & E. Kohlm.
Hypophloeda rhizospora K.D. Hyde & E.B.G. Jones
Hypoxylon oceanicum Schatz
Hypoxylon kretzschmarioides Y.M. Ju & J.D. Rogers
Jahnula australiensis K.D. Hyde
Kirschsteiniothelia elaterascus Shearer
B,R
R
F
F
M
M
M
M
M
M
M
M
B
M
M
M
F
M
M
B
B
F
F
M
M
M
M
M
F
M
M
M
M
M
M
M
M
M
M
M
M
F
F
F
11
11
9
9
2,3
3
2,3
2
1
2,3,4,5
1
1,2,3,4,5
10
2
1,2,3,4,5
2,5
8
4
5
11
11
10
9
2,3
1,4
2,3,5,7
2,3
2,4,5
9
1,2,3,4,5,6
5,6
2,3,4,5,6
2,5
2,3,4,5
2,3
2,3,5
2,3
2,3,5,7
1,2,3,4,5
4,5,6
4,5,6,7
9
9
9

Fungal Diversity
29
Table 4 continued. Checklist of fungi from aquatic habitats in Brunei.
Species Habitat Reference
*Lasiosphaeria sp. 1
*Lasiosphaeria sp. 2
Lasiosphaeria breviseta P. Karst.
*L. immersa P. Karst.
Lautospora gigantea K.D. Hyde & E.B.G. Jones
Leiosphaerella sp.
Leptosphaeria sp.
Leptosphaeria australiensis (Cribb & J. Cribb) G.C. Hughes
L. cf. avicenniae Kohlm. & E. Kohlm.
Lignincola laevis Höhnk
L. tropica Kohlm.
Lindra marinera Meyers
L. thalassiae Orpurt, Meyers, Boral & Simms
Linocarpon pandani (Syd. & P. Syd.) Syd. & P. Syd.
Linocarpon sp.
Lophiostoma aquaticum (Webster) Aptroot & K.D. Hyde
L. bipolare (K.D. Hyde) Liew, Aptroot & K.D. Hyde
*L. frondisubmersum (K.D. Hyde) Liew, Aptroot & K.D. Hyde
*L. tetraploa (Scheuer) Aptroot & K.D. Hyde
Lulworthia sp. (spores 136-195 µm)
Lulworthia sp. (spores 220-335 µm)
Lulworthia sp. (spores 340-490 µm)
Lulworthia grandispora Meyers
Mamillisphaeria dimorphospora K.D. Hyde, S.W. Wong &
E.B.G. Jones
Manglicola guatemalensis Kohlm. & E. Kohlm.
*Mangrovispora pemphii K.D. Hyde & Nakagiri
Marinosphaera mangrovei K.D. Hyde
Massarina sp. 1
Massarina sp. 2
Massarina sp. 3
+M. rubi (Fuckel) Sacc.
M. thalassiae Kohlm. & E. Kohlm.
M. velataspora K.D. Hyde & Borse
Mycosphaerella pneumatophora Kohlm.
Nais glitra Crane & Shearer
Nectria cf. byssicola
+N. haematococca Berk. & Broome
Nimbospora bipolaris K.D. Hyde & E.B.G. Jones
N. octonae Kohlm.
Ophioceras dolichostomum (Berk. & Curtis) Sacc.
Ophiodeira monosemeia Kohlm. & Volkm.-Kohlm.
*Orbilia sp.
Orcadia cf. ascophylli
Oxydothis sp.
B
B,R
F
B
M
M
M
M,R
M
M
M
M
M
M
M
F
F,M
F,M
B
M
M
B,M
M
F
M
B
M
F
M
M
R
M
M
M
M
F
R
M
M
F
M
F,B,R
M
M
11
11
9
11
5
2
1,2,3,4,5
1,2,3,4,5,6,11
1,2,3,4,5,7
1,2,3,4,5,7
1,2,3,4,5
2
2
2,3
2,3
9
9,10
8,10
11
2,3,4
1,2,3,4,5,6
1,2,3,4,5,7,10,
11
1,2,3,4,5,6,7,
8,9
1,2,4,6
10
1,4,5
8
2
2
11
1,2,5
1,2,3,4,5
5
5
9
11
2,3
2
9
4,5
10,11
1,2,3
3

30
Table 4 continued. Checklist of fungi from aquatic habitats in Brunei.
Species Habitat Reference
Passeriniella savoryellopsis K.D. Hyde & Mouzouras
+Phaeosphaeria sp.
*Phaeosphaeria capensis T.D. Steinke & K.D. Hyde
*Phomatospora sp.
Phomatospora kandeliae K.D. Hyde
Pleospora sp.
Pleospora state of Stemphylium majusculum
Quintaria sp.
Remispora crispa Kohlm.
R. galerita Tubaki
Rhizophila marina K.D. Hyde & E.B.G. Jones
Rosellinia sp.
Salsuginea ramicola K.D. Hyde
Savoryella sp.
Savoryella aquatica K.D. Hyde
S. fusiformis W.H. Ho, K.D. Hyde & Hodgkiss
S. lignicola E.B.G. Jones & R.A. Eaton
S. paucispora (Cribb & J. Cribb) Koch
S. verrucosa Minoura & Muroi
Sphaerulina cf. oraemaris
*Sungaiicola brachydesmiella Fryar & K.D. Hyde
Swampomyces cf. armeniacus Kohlm. & Volkm.
+S. triseptatus K.D. Hyde & Nakagiri
Thalassogena sphaerica Kohlm. & Volkm.-Kohlm.
Torpedospora radiata Meyers
*Torrentispora crassiparietis Fryar & K.D. Hyde
*T. fibrosa K.D. Hyde, W.H. Ho, E.B.G. Jones, K.M. Tsui & S.W.
Wong
*T. fusiformis Fryar & K.D. Hyde
Trematosphaeria lignatilis Kohlm.
T. striataspora K.D. Hyde
Trematosphaeria sp.
Tubeufia palmarum (Torrend) Samuels, Rossman & E. Müll.
Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm.
Basidiomycetes
Calathella mangrovei Jones & Kohlm.
Halocyphina villosa Kohlm. & E. Kohlm.
Nia vibrissa Moore & Meyers
Anamorphic fungi
Acrogenospora sphaerocephala (Berk. & Broome) Ellis
*Alysidium sp.
Aquaphila albicans Goh, K.D. Hyde & W.H. Ho
*Arthrobotrys oligospora Fresen.
M
R
F,B
M
M
M
F
F
M
M
M
M
M
M
F
F
M,F
M
F,B
M
F,B,M
M
R
M
M
F,B
F,B
F,B
M
M
M
F
M
M
M
M
F
F
F
B,R
2,4,5
11
10
10
7
1,2,3
9
9
2,3
2,3
3,4,5
2,3
7
2,3
9
9
1,2,3,4,5,7,9,10
1,2,5,10
9,10
2,5
10,11
4,5
11
4,5
2,3,4,5
10
10
10
1,2,4,5
4,10
2,3
9
7
7
2,3,5,7
2,3
9
10
9
11

Fungal Diversity
31
Table 4 continued. Checklist of fungi from aquatic habitats in Brunei.
Species Habitat Reference
Bactrodesmium sp.
B. linderi (Crane & Shearer) Palm & Stewart
*Berkleasmium sp.
*Beverwykella pulmonaria (Beverw.) Tubaki
*Brachysporiella gayana Bat.
Canalisporium caribense (Hol.-Jech. & Mercado) Nawawi &
Kuthub.
C. elegans Nawawi & Kuthub.
C. pulchrum (Hol.-Jech. & Mercado) Nawawi & Kuthub.
*Cancellidium applanatum Tubaki
Candelabrum brocchiatum Tubaki
Ceuthospora gaeumannii Nag Raj
*Chalara sp.
Chloridium lignicola (Mangenot) W. Gams & Hol.-Jech.
Chrysosporium sp.
Cirrenalia pseudomacrocephala Kohlm.
C. pygmea Kohlm.
C. tropicalis Kohlm.
Cladorrhinum samala (Subram. & Lodha) W. Gams &
Mouchacca
Clavariopsis bulbosa Anastasiou
+Coleodictyospora cubensis Charles
+Conioscypha sp.
Cordana abramovii var. seychellensis K.D. Hyde and Goh
Cytospora rhizophorae Kohlm. & Kohlm.
Dactylaria triseptata (Matsush.) R.F. Castañeda & W.B. Kendr.
*Dactylella sp.
Denticularia limoniformis de Hoog
Dictyochaeta fertilis (Hughes & Kendrick) Holubova-Jechova
Dictyosporium alatum van Emden
D. pelagicum (Linder) G.C. Hughes
Diplodia sp.
*Exserticlava vasiformis (Matsush.) S. Hughes
+Graphium putredinis (Corda) Hughes
Haplochalara angulospora Linder
*Helicomyces sp.
Helicomyces roseus Link
H. torquatus L.C. Lane & Shearer
Humicola alopallonella Meyers & Moore
*Intercalarispora nigra J.L. Crane & Schokn.
+Lasiodiplodia sp.
Megaloseptoria mirabilis N.A. Naumov
Microsphaeropsis olivacea (Bonord.) Höhn.
Monodictys capensis R.C. Sinclair, Boshoff & Eicker
M. pelagica (T.W. Johnson) Jones
M
M
F,B
F
F,B
F
F
F
F,B,M
F
F
M
F
M
M
M
M
F
M
R
R
F
M
F
F,B
F
F
F
M
M
F,B
R
F
B
F
F
M
B
R
F
F
F
F,B,M,R
2
4
10,11
10,11
10,11
9
9
9
10,11
9
9
10
9
1,2,3
2,4,5,6
1,2,3,4,5, 6,7
1,2,3,4,5,6
9
2,3
11
11
9
5
9
10
9
9
9
2,3,4,5
5,6
10,11
11
9
11
9
9
1,2,3,4,5
11
11
9
9
9
1,2,3,10,11

32
Table 4 continued. Checklist of fungi from aquatic habitats in Brunei.
Species Habitat Reference
Monotosporella setosa var. macrospora G.C. Hughes
Nawawia filiformis (Nawawi) Marvanová
*Papulospora sp. 1
*Papulospora sp. 2
Periconia prolifica Anastasiou
Phaeoisaria clematidis (Fuckel) Hughes
P. sparsa Sutton
Phialocephala xalepensis Maggi & Persiani
Phialogeniculata africana Goh, K.D. Hyde & T.D. Steinke
Phialophorophoma litoralis Linder
Phoma sp.
Phomopsis sp.
Phragmospathula sp.
Pleurothecium recurvatum (Morgan) Höhn.
Polystigmina sp.
Pseudospiropes cubensis Hol.-Jech.
Robillarda rhizphorae Kohlm.
Scolecobasidium dendroides Pirozynski & Hodges
Sibrina orthospora W. Gams
*Spacidoides sp.
*Sporidesmium cf. anglicum (Grove) M.B. Ellis
*S. crassisporum M.B. Ellis
S. ellipticum Moore
S. macrurum (Sacc.) M.B. Ellis
Sporidesmium sp.
Sporoschisma saccardoi E.W. Mason & S. Hughes
S. uniseptatum Bhat. & W.B. Kendr.
Stilbella holubovae Seifert, S.J. Stanley & K.D. Hyde
Tiarosporella paludosa (Sacc. & Fiori) Höhn.
Topospora sp.
Trichocladium sp.
Trichocladium achrasporum (Meyers & Meyers) Dixon
Trichocladium cf. opacum (Corda & Shearer)
Varicosporina ramulosa Meyers & Kohlm.
Verticillium sp. 1
Verticillium sp. 2
Virgariella atra S. Hughes
Xylomyces sp.
Xylomyces chlamydosporis Goos, R.D Brooks & Lamore
*X. giganteus Goh, W.H. Ho, K.D. Hyde & K.M. Tsui
Zalerion varium Anastasiou
F
F
B,M
B,M
M
F,B,R
F
F
F
M
M
M
M
F
M
F
M
F
F
B
F,B,M,R
F,R
M
F
M
F
F
F
F
M
M
M
M
M
F
F
F
B,M
F,B
B
M
9
9
10,11
10,11
1,2,3,4,5,6
9,10,11
9
9
9
2,4,5,6
2,3,4,5
4,7
2,5
9
5
9
2
9
9
11
10,11
10,11
2
9
2
9
9
9
9
1
3,4,5,7
2,3,4,5
1,2,4,5
2
9
9
9
2,3,4,5,6,11
9,11
11
2,3
*indicates species which are new records for Brunei in either this study or Fryar et al., 2004b,
+ indicates species which have been found only in the riparian vegetation i.e. not from an
aquatic habitat. F = freshwater, B = brackish, M = marine, R = in riparian vegetation. 1 = Hyde,
1988a, 2 = Hyde, 1988b, 3 = Hyde, 1989, 4 = Hyde, 1990a, 5 = Hyde, 1990b, 6 = Hyde, 1991,
7 = Hyde, 1992, 8 = Wong, 1996, 9 = Ho et al., 2001, 10 = This study, 11 = Fryar et al., 2004a.

Fungal Diversity
33
Multivariate community analysis
Detrended Correspondence Analysis (DCA) showed a trend of marine
sites on the left through to the freshwater sites on the right (Fig. 2). Axes 1 and
2 explained 35% of the variation. The species that explained the most variation
in axis 1 were Alysidium sp., Ascomycete sp. 1, Ascotaiwania sp.,
Beverwykella pulmonaria, Brachysporiella gayana, Chaetosphaeria sp.,
Dactylella sp., Exserticlava vasiformis, Massarina sp., Pleurophragmium-like,
Sporidesmium crassisporum, Torrentispora fusiformis and Tritirachium sp.
Axis 2 was mostly explained by Aniptodera megalospora, Ceratosphaeria sp.
1, Lasiosphaeria sp. 3, Lophiostoma bipolare, Lophiostoma frondisubmersum
and Phomatospora sp.
Discussion
Salinity is one of the most important factors influencing the distribution
of fungi in aquatic habitats (Shearer, 1972). Studies consistently find different
assemblages of fungi in fresh and marine habitats (e.g. Shearer and Von
Bodman, 1983; Shearer, 1993; Czeczuga, 1996; Alias and Jones, 2000; Sarma
and Vittal, 2000, 2001; Sivichai et al., 2002). However, the distribution of
fungi in intermediate habitats (brackish water) is less well known. Shearer
(1972) found that significant changes occurred to the species composition at
points where salt and freshwater mixed. Although several species were able to
tolerate a wide range of salinities, many only occurred at brackish water sites
(7.8-17.9‰) (Shearer, 1972). Similarly, in this study several species were
found at all sites, and were common, whereas others were unique to the
brackish water sites.
Previous studies in Brunei had found distinct assemblages of fungi in
fresh and sea water (Hyde, 1988a,b, 1989, 1990a,b, 1991, 1992; Wong, 1996;
Ho et al., 2002). However, as salinity was generally not recorded it was
unknown whether there is a mixture of ‘freshwater’ and ‘marine’ species or a
completely different assemblage of fungi i.e. ‘brackish water’ species. The
current data suggests that there are both. Some freshwater species also
occurred in brackish habitats, but not in marine habitats (e.g. Torrentispora
crassiparietis, T. fibrosa, T. fusiformis and Dactylella sp.). Similarly some
marine species occurred in brackish habitats but not in freshwater (e.g.
Aniptodera chesapeakensis and Lulworthia spp.). This suggests that brackish
water supports a mixture of freshwater and marine species. There were,
however, several rare, unique species in the brackish habitats (e.g.
Annulatascus hongkongensis, A. palmietensis and Savoryella verrucosa) that

34
Fig. 2. Plot of axis1 vs. axis 2 scores from Detrended Correspondence Analysis.
indicate brackish habitats support an assemblage of fungi distinct from
freshwater and marine assemblages.
The Detrended Correspondence Analysis indicated that the assemblages
of fungi in the brackish sites were intermediate between marine and freshwater
sites, i.e. that there is a continuum of species rather than distinct communities
at each site. Research in other parts of the world has found that some ‘marine’
species have a tolerance for brackish water and can even grow in freshwater,
but not as vigorously (Byrne and Jones, 1975; Curran, 1980). Likewise,
‘freshwater’ species (and even some terrestrial species) have been found to
have some tolerance for salinity (Curran, 1980). This was also observed in this
study with Aniptodera chesapeakensis and Sungaiicola brachydesmiella
showing a gradient of abundance from marine to freshwater sites. However,
some species (Annulatascus velatisporus, Exserticlava vasiformis,
Papulospora sp. 1, Papulospora sp. 2, Xylomyces sp., and Xylomyces
Axis 2
Axis 1
Brackish 1(1)
Marine (1)
Brackish 2(1) Fresh (1)
PSF (2)
Brackish 1(2)
Marine (2)
0.0
0.6
1.3
1.9
2.6
3.2
0.0 0.6 1.3 1.9 2.6 3.2

Fungal Diversity
35
giganteus) were more abundant in brackish water habitats than freshwater or
marine suggesting these are ‘brackish water’ species that can also live in either
freshwater or marine habitats. A good example is Monodictys pelagica that
was found in brackish and freshwater habitats. Monodictys pelagica has
previously been noted as being a brackish water species (Tubaki, 1969) based
on physiological evidence. Optimum salinity for this fungus is 20-60% sea
water (Jones and Jennings, 1964; Tubaki, 1969).
Shearer (1972) found that the ratio of ascomycetes to hyphomycetes
increased with increasing salinity. This trend was also seen in the first set of
samples of this study. The reason for this is unknown, but perhaps there are
different stimuli for sporulation in marine and freshwater habitats, or the
osmotic potential of salt water is too great for many conidia.
This is only the second time that Fluviatispora reticulata has been found.
It was described from fruiting bodies found on rachides of Livistona sp. in the
Bensbach River, Papua New Guinea (Hyde, 1994).
Cancellidium applanatum was common at all sites. This aero-aquatic
hyphomycete was originally described from balsa-wood test blocks in a lake in
Japan (Tubaki, 1975). However, it has since been recovered from submerged
decaying leaves in Malaysia (Webster and Davey, 1980) and Queensland,
Australia (Shaw, 1994). It has not been previously recorded in Brunei.
The significant finding of two new genera and two new species is
indicative of the lack of knowledge of fungi in aquatic habitats, and the general
lack of funded fungal taxonomists (Miller, 1995; Hyde, 2003).
Acknowledgements
We would like to thank Department of Forestry, Brunei for allowing collection of wood
samples. This work was completed under a University of Hong Kong Postdoctoral Fellowship
and funding was provided by a CRGC grant (Hong Kong University).
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(Received 15 March 2004; accepted 8 June 2004)