Inter- and intra stream variation of lignicolous freshwater fungi in tropical Australia

Abstract

Vijaykrishna, D. and Hyde, K.D. (2006). Inter-and intra stream variation of lignicolous freshwater fungi in tropical Australia. Fungal Diversity 21: 203-224. Freshwater ecosystems are in a constant interaction with the terrestrial environment (riparian) surrounding them. Riparian vegetation is the major source of organic input into the stream ecosystem, which includes woody debris. The impact of the type of riparian vegetation on the biodiversity of lignicolous freshwater fungi in five tropical streams of the Barron River catchment area in Atherton Tablelands, Queensland, Australia was investigated. The collection sites were broadly classified in three types based on the kind of riparian vegetation; pristine, re-growth and agricultural zones. Fifty wood samples collected from each of 12 sites yielded 162 fungal taxa. The dominant fungi were species of Annulatascus, Aquaticola (Annulatascaceae), Anthostomella (Xylariaceae), Massarina (Lophiostomataceae) and Savoryella (Sordariales incertae sedis). The highest species diversity was found in the pristine forest zone, followed by agricultural zone, while the re-growth zone was least diverse. Species overlap was seen between all streams, and between the three types of riparian vegetation. The fungal species showed little habitat recurrence, however major changes were observed in species richness and abundance, with varying degrees of human disturbance.

Full text

Fungal Diversity
203
Inter- and intra stream variation of lignicolous freshwater fungi
in tropical Australia
D. Vijaykrishna and Kevin D. Hyde*
Centre for Research in Fungal Diversity, Department of Ecology & Biodiversity, The
University of Hong Kong, Pokfulam Road, Hong Kong SAR, PR China
Vijaykrishna, D. and Hyde, K.D. (2006). Inter- and intra stream variation of lignicolous
freshwater fungi in tropical Australia. Fungal Diversity 21: 203-224.
Freshwater ecosystems are in a constant interaction with the terrestrial environment (riparian)
surrounding them. Riparian vegetation is the major source of organic input into the stream
ecosystem, which includes woody debris. The impact of the type of riparian vegetation on the
biodiversity of lignicolous freshwater fungi in five tropical streams of the Barron River
catchment area in Atherton Tablelands, Queensland, Australia was investigated. The collection
sites were broadly classified in three types based on the kind of riparian vegetation; pristine, re-
growth and agricultural zones. Fifty wood samples collected from each of 12 sites yielded 162
fungal taxa. The dominant fungi were species of Annulatascus, Aquaticola (Annulatascaceae),
Anthostomella (Xylariaceae), Massarina (Lophiostomataceae) and Savoryella (Sordariales
incertae sedis). The highest species diversity was found in the pristine forest zone, followed by
agricultural zone, while the re-growth zone was least diverse. Species overlap was seen
between all streams, and between the three types of riparian vegetation. The fungal species
showed little habitat recurrence, however major changes were observed in species richness and
abundance, with varying degrees of human disturbance.
Key words: ascomycetes, biodiversity, riparian vegetation, species abundance, submerged
wood
Introduction
The riparian zone is a dynamic habitat characterised by material
exchange between terrestrial and stream ecosystems (Wetzel, 2001). The input
of organic matter (leaves, twigs, branches, whole trees) from riparian
vegetation to streams is one of the most significant processes occurring at the
interface of terrestrial and stream ecosystems (Dolloff and Webster, 2000),
influencing stream food webs and total ecosystem functioning (Wallace et al.,
1997). Woody debris deposited into stream ecosystems affects hydraulic
conditions, defines channel morphology and provide habitats for various
aquatic organisms (Keller and Swanson, 1979; Mosley, 1981; Harmon et al.,
1986; Sedell et al., 1988). Leaves, small pieces of wood (< 2 mm) and
*Corresponding author: K.D. Hyde; e-mail: kdhyde@hkucc.hku.hk

204
reproductive structures of terrestrial plants account for more than 90% of the
overall transport of organic matter downstream in temperate as well as tropical
streams (Benson and Pearson, 1993). In contrast, larger woody substrates
entering the stream from the riparian vegetation show little or no transport and
limited breakdown (Webster et al., 1999). Even though the breakdown of wood
is slower and its inputs are lower than that of leaf litter, their contributions to
energy flux are higher. This is due to their resistance to downstream transport
resulting in a relatively high standing stock (Díez et al., 2002). However,
excessive amounts in stream ecosystems can have negative impacts, such as
reduce dissolved oxygen and build up toxic levels of wood (Hicks, 1997).
Therefore the decomposition of woody substrates into fine particulate organic
matter (FPOM) that are easily transported and recycled is essential. The
transport rates of FPOM resulting from wood breakdown (< 3 mm diam) have
been shown to be substantially larger in tropical than those in temperate
systems (Webster et al., 1995b).
Freshwater fungi play an important ecological role in tropical stream
food webs as they enzymatically degrade allochthonous matter into more
palatable forms, e.g. Ingoldian fungi are the dominant micro-organisms
associated with decomposing leaf litter in stream ecosystems (Suberkropp,
1997; Abdel-Raheem and Shearer, 2002; Bucher et al., 2004; Gönczöl and
Révay, 2004) and their biodiversity, distribution and ecological role in
decaying leaves have been well investigated (Bärlocher, 1992; Suberkropp,
1997; Pascoal et al., 2005).
Studies on fungi inhabiting submerged plant debris and wood have been
undertaken in temperate (Bärlocher, 1992; Shearer, 1993; Hyde and Goh,
1999a; Cai et al., 2002b; Van Ryckegem and Verbeken, 2005a,b) and tropical
regions (Hyde and Goh, 1997, 1998a,b; Goh and Hyde, 1999, Tsui et al., 2000;
Cai et al., 2002a, 2003a, 2005; Luo et al., 2004; Tsui and Hyde, 2004; Fryar et
al., 2004, 2005), with the tropics exhibiting a greater diversity of aquatic fungi.
More than 600 species, belonging to diverse taxonomic groups, have been
described from woody substrates in freshwater ecosystems (see Shearer, 2001;
Cai et al., 2003b, Goh and Tsui, 2003). Collection expeditions have resulted in
a number of new taxa from freshwater samples in tropical Australia (Hyde et
al., 1997; Goh and Hyde, 1999).
Fungi generally show recurrence in specific habitats and abundance of
habitat therefore limits species abundance, and likely influences fungal
communities (Lodge, 1997). Studies have shown significant correlations
between certain attributes of stream habit and the local diversity of fungi (e.g.
Tsui and Hyde, 2004). Thus, habitat modification resulting from human
activities (e.g. riparian deforestation) should strongly influence the distribution

Fungal Diversity
205
and abundance of freshwater fungi. However, habitat abundance relationships
for lignicolous freshwater have rarely been conducted. This is particularly the
case for tropical streams, where anthropogenic habitat modification has been
shown to influence biodiversity of many other stream organisms (Dudgeon,
1999, 2000).
The aim of this study was to assess the impact of riparian forest
disturbance on the diversity, richness and abundance of lignicolous freshwater
fungi in tropical Australia. We compared the diversity of wood-inhabiting
fungi from five tropical streams, with three different kinds of riparian
vegetation (Primary forest, secondary/regrowth forest and cleared/agricultural
riparian forests). The objectives are (1) to assess the biodiversity of fungi on
submerged wood in a large tropical forest area, (2) to assess the inter- and
intra- stream variation of fungi on submerged wood and (3) to characterise the
effects of riparian forest cover on the diversity of fungi on wood logs and
twigs.
Materials and methods
Site of study
This research was undertaken in the Atherton Tablelands, a high flat land
in Queensland, northeastern Australia. The tableland rises gradually to the east
and southeast, where Mount Bartle Frere, 5287 feet high, forms the highest
point. The region receives a mean annual rainfall of 1200-4000 mm, over 60%
of which falls in summer (December-March). The period of study was drier
than average.
Experimental design
To examine the effect of riparian vegetation on the diversity of
lignicolous freshwater fungi, stream sites were categorised into three types
according to their riparian vegetation: pristine forest area, which is mostly
undisturbed and lies within the World Heritage forest; secondary forest area
comprising wood lands or a buffer forest area beside the stream, which are
mostly part of National or State forests; and the agricultural riparian zone,
which is in close proximity to agricultural fields and roads (Fig. 1).
The sampling was conducted during April 2002 and March 2003 (Table
1), when the temperatures ranged from an average of 17ºC at night to 26ºC in
the day. The 12 study sites were separated by approximately 2 to 20 kms and
included 5, third and fourth order streams, the Clohesy River, Davies Creek,
Emerald, Kauri Creek and Tinaroo Creeks (Fig. 1).

206
Fig. 1. The location of the 12 collection sites, on the 5 streams, flowing into the Barron River.
Arrows indicate flow direction.
Methodology
Fifty submerged wood samples were randomly collected from a 100 m
section of each stream site. The collection was carried out starting by walking
upstream systematically in a zigzag fashion to randomise any bias associated
with near-shore or midstream observations, and also pool and riffle effects.
Utmost care was taken to collect only samples that had been submerged for a
long period, by observing the degree of degradation. The collected samples
included segments of trunks, branches and twigs from a variety of unidentified
angiosperms. The size of the wood samples ranged from 15-30 cm long and 1-
5 cm in diameter. All samples were placed in plastic bags in the field with
moistened tissue paper, and returned to the laboratory in Hong Kong within 4
days. The samples were then incubated individually in sterile plastic boxes
lined with moistened tissue paper at room temperature (22-24ºC). The woody
substrates were examined under a dissecting microscope for fruiting bodies on

Fungal Diversity
207
Table 1. Name, place and date of collection.
Day one and at 7 days intervals thereafter until 9 months after collection. For
identification, fungi were mounted in water and lactic acid. Measurements
were made from fresh material mounted in water. Attempts were made to
isolate single spore cultures of new and interesting fungi.
Number of species, and frequencies of occurrence of each species were
recorded and calculated for each sampling site. Frequency of occurrence was
calculated based on the following formula:
Number of samples of wood that a particular fungal species occurred on ×100%
Number of samples of wood examined
The number of species, their frequency and abundance was used to calculate
the biodiversity index Brillouin (HB) (Magurran, 2004):
HB =lnN!−lnni!
∑
N
where ni is the species abundance of the ith species, and N the sum of
abundance of all species in the community.
pi =ni
ni
i=1
s
∑
×100%,i=1,2,3,...,S,
where pi is percentage abundance of the ith species, ni is the number of samples
with the ith species, and S is the number of species in the community. Taxa
with more than 3% abundance were classified as dominant fungi (Ho et al.,
2001). To compare community structure, all taxa were sorted in descending
order by their abundance, and species-abundance distributions were plotted for
each sample. To emphasise which were dominant and which were rare, taxa
are presented in overall descending total abundance for the 12 sample sites.
Davies
Creek
Clohesy
River
Tinaroo
Creek
Kauri
Creek
Emerald
Creek
Primary Apr' 2002 Apr' 2002 NA Mar' 2003 NA
Secondary Apr' 2002 Apr' 2002 Mar' 2003 Mar' 2003 Mar' 2003
Agricultural Apr' 2002 Apr' 2002 Mar' 2003 NA Mar' 2003

208
Sørenson’s similarity index was used to compare the fungal assemblages
at different sites. Sørenson’s similarity index expresses the measured species
occurrences against the theoretically possible ones (Sørenson, 1948).
Sørenson’s index of similarity is calculated using the formula 2c/(a+b), where
a = total number of species in the first community, b = total number of species
in the second community, and c = number of species both communities have in
common. Communities at all sites were first compared resulting in 66
combinations. This was followed by the comparison of the sites based on
riparian vegetation resulting in 3 combinations.
Multivariate analysis (Kenkel and Booth, 1992) was performed to
summarise and reveal underlying trends of fungal community structure to
collection sites. To perform ordination of variables simultaneously (i.e. with
variables positioned in the same ordination space), correspondence analyses
were performed (Kenkel and Booth, 1992). A data matrix consisting of the
numbers of colonised wood samples from each collection site and their
frequencies of occurrence were subjected to correspondence analysis (Anon,
1995).
Results
Species diversity and richness
Fungal sporulation was observed from the first week until 9 months from
the collection date. A total of 162 fungal taxa were recorded from 600 wood
samples from the 12 sites within the Barron river catchment (Table 2),
including 101 ascomycetes and 61 anamorphic fungi (56 hyphomycetes; 5
coelomycetes). An average of 2.03 taxa per woody substrate occurred at each
site, with a range of 1.3 - 2.84 taxa per sample per site.
Most ascomycete species belonged to the families Annulatascaceae (20
species), Halosphaeriaceae (20 species) and Lophiostomataceae sensu Barr
(1979) (10 species) and also many species whose taxonomic placement are
uncertain (Table 3). Annulatascus velatisporus was most common, appearing
in all the 12 sites, followed by Canalisporium pulchrum, Massarina
australiensis and Massarina thalassioidea in 10 sites, followed by Aquaticola
hyalomura, Lophiostoma ingoldianum, Ophioceras dolichostomum and
Savoryella lignicola in 9 sites each.
Pristine and agricultural zones had higher species richness than regrowth
sites, with >100 taxa (Table 2). The pristine zone at Kauri creek (KP) had the
greatest species richness (142 taxa). In contrast, four out of five regrowth zones
had <100 species, with the Clohesy regrowth site (CR) having the lowest
species richness, 63 taxa (1.3 species per wood sample).

Fungal Diversity
209
Table 2. Diversity and species richness of freshwater fungi at various sites.
*CP = Clohesy Pristine, CR = Clohesy Regrowth, CV = Clohesy Agricultural, DP = Davies
Pristine, DR = Davies Regrowth, DV = Davies Agricultural, ER = Emerald Regrowth, EV =
Emerald Agricultural, KP = Kauri Pristine, KR = Kauri Regrowth, TR = Tinaroo Regrowth and
TV = Tinaroo Agricultural.
Dominance
Species-abundance distributions (Fig. 2) showed that only a few species
dominated at each site (percentage abundance ≥ 3%). Among the 12 sites, the
most abundant taxa were Massarina australiensis from Clohesy River
Agricultural (CV) zone (15% of the total occurrence of the fungi from the site),
followed by Canalisporium pulchrum from Clohesy Pristine (CP) zone (14%;
Table 3). There was moderate overlap of dominant species among the sites. All
sites had many rare taxa that appeared only once, except for the Tinaroo Creek
Agricultural site (TV) where every taxon was recorded at least twice. There
were also taxa with intermediate abundance (1-3% of the total occurrence of
fungi) in all sites.
Similarity
Similarity indices were calculated among all 12 sites to evaluate the
similarity of freshwater fungal communities at different sites (Table 4).
Similarity was greatest between Emerald Creek regowth zones (ER) and
Tinaroo creek Agricultural zone (TV) (36%) and lowest between Clohesy
pristine (CR) and Tinaroo regrowth (TR) (5%), Clohesy regrowth (CR) and
Tinaroo agricultural (TV) (5%). For sites on the same stream, similarity was
greatest for Clohesy River (18% to 31%) and Davies creek (25% to 27%) sites.
The least similarity between the sites on the same stream was for Tinaroo
creek, where the regrowth zone (TR) and agricultural zone (TV) had a
similarity of 16%.
Multivariate analysis
Multivariate analysis showed that samples from sites Davies Creek (DP,
DR, DV) and Emerald Creek (ER, EV) grouped together, while in Clohesy
Creek only the pristine and regrowth zones (CP, CV) grouped together.
CP* CR CV DP DR DV ER EV KP KR TR TV
Total number of species 35 31 39 61 45 51 32 29 56 41 36 25
Total number of isolates 114 63 120 107 91 100 72 110 142 89 111 102
Number of taxa per sample 2.28 1.3 2.4 2.14 1.82 2 1.44 2.2 2.84 1.78 2.22 2.04
Brillouin's index 1.25 1.21 1.29 1.47 1.34 1.4 1.22 1.21 2.03 1.35 1.27 1.26

210
Table 3. Average % abundance of lignicolous freshwater fungi in all the 12 sites, in decreasing
order. See Table 2 for abbreviation.
Species Name CP CR CV DP DR DV ER EV KP KR TR TV Average %
abundance
Massarina australiensis 3.5 3.2 15 3.7 7.7 3 - - 0.7 3.4 11.7 6.9 4.9
Annulatascus velatisporus 6.1 4.8 5.8 3.7 6.6 5 2.8 9.1 1.4 4.5 2.7 4.9 4.8
Canalisporium pulchrum 14 4.8 4.2 0.9 - 3 8.3 5.5 4.9 3.4 5.4 - 4.5
Lophiostoma ingoldianum - 6.3 0.8 2.8 7.7 1 9.7 11.8 2.8 - 5.4 - 4
Massarina thalassioidea - 11.1 0.8 0.9 2.2 1 5.6 5.5 10.6 1.1 4.5 - 3.6
Aquaticola ellipsoidea - 6.3 - 2.8 2.2 3 - - 3.5 5.6 2.7 5.9 2.7
Anthostomella aquatica 10.5 1.6 1.7 - - - 2.8 4.5 6.3 3.4 - - 2.6
Acanthophysis-like taxon 6.1 7.9 - 6.5 2.2 - 2.8 - 2.1 - - - 2.3
Savoryella aquatica 3.5 4.8 - 2.8 1.1 1 - 5.5 - 4.5 3.6 - 2.2
Ophioceras dolichostomum 4.4 4.8 4.2 1.9 1.1 1 - - 0.7 5.6 0.9 - 2
Lophiostoma bipolare 6.1 - - 2.8 3.3 - 2.8 - 2.1 - - 6.9 2
Helicomyces roseus - 1.6 - 2.8 5.5 6 4.2 - 1.4 - 1.8 - 1.9
Annulatascus biatriisporus 1.8 1.6 3.3 - 1.1 - - 4.5 - - 6.3 3.9 1.9
Clohesyomyces aquaticus - - 5.8 4.7 1.1 3 1.4 - - 2.2 - 2.9 1.8
Aquaphila albicans - - - 5.6 2.2 1 4.2 - - 3.4 4.5 - 1.7
Annulatascus triseptatus - - - 1.9 1.1 - - - 0.7 3.4 9.9 2.9 1.7
Quintaria submersa 2.6 3.2 3.3 1.9 5.5 3 - - - - - - 1.6
Sporoschisma nigroseptatum 0.9 - - 0.9 4.4 1 - 0.9 - 4.5 0.9 5.9 1.6
Aquaticola hyalomura 3.5 3.2 - 2.8 1.1 1 - - 1.4 3.4 0.9 2 1.6
Halosarpheia aquatica - 1.6 6.7 - - 2 1.4 5.5 - 1.1 - - 1.5
Kirschsteiniothelia elaterascus 4.4 4.8 4.2 0.9 - 3 - - - - - - 1.4
Quintaria sp. - - - - - - 8.3 8.2 - - - - 1.4
Mamillisphaeria dimorphospora - - - - - 7 1.4 - - - 5.4 2 1.3
Savoryella lignicola 0.9 - - 1.9 1.1 4 1.4 - 1.4 1.1 0.9 2.9 1.3
Dactylaria tunicata 0.9 - - 1.9 1.1 1 2.8 0.9 0.7 4.5 - - 1.1
Xylomyces elegans - - - - - 4 - 0.9 1.4 3.4 - 3.9 1.1
Rivulicola incrustata 0.9 - - 0.9 4.4 - - 0.9 - - 5.4 - 1
Aniptodera lignatilis - - - 0.9 1.1 1 1.4 5.5 1.4 1.1 - - 1
Jahnula bipolaris 0.9 1.6 0.8 0.9 - 2 - - - - - 5.9 1
Aquaticola longicolla 3.5 3.2 - - - - - - 0.7 4.5 - - 1
Submersisphaeria aquatica - - 4.2 - - - - - - - - 6.9 0.9
Cataractispora appendiculata 0.9 - - 0.9 - - - 2.7 3.5 1.1 1.8 - 0.9
Savoryella fusiformis - - - - - - 8.3 - 1.4 - - - 0.8
Jahnula australiensis 3.5 3.2 - 0.9 - 2 - - - - - - 0.8
Chaetosphaeriaceae sp. - - - 1.9 - 2 - - 3.5 - - 2 0.8
Aniptodera chesapeakensis - - - - - - 1.4 4.5 2.1 1.1 - - 0.8
Nais inornata - - 4.2 - - - - - 0.7 - - 3.9 0.7
Helicosporium hiospiroides - - - - - - 5.6 0.9 - 2.2 - - 0.7
Delortia palmicola - - - 1.9 1.1 1 - 0.9 - - - 2.9 0.7
Acrogenospora sphaerocephala - - - 0.9 - - - 5.5 1.4 - - - 0.6

Fungal Diversity
211
Table 3. Average % abundance of lignicolous freshwater fungi in all the 12 sites.
Species Name CP CR CV DP DR DV ER EV KP KR TR TV Average %
abundance
Halosarpheia retorquens - - - - - - - 1.8 0.7 2.2 - 2.9 0.6
Ellisembia leonense 0.9 1.6 1.7 - - - - - 3.5 - - - 0.6
Aniptodera lignicola 2.6 1.6 3.3 - - - - - - - - - 0.6
Tripterosporaceae sp. - - - 0.9 1.1 - - - - 4.5 0.9 - 0.6
Quintaria aquatica 0.9 - - 1.9 - 1 2.8 - - - 0.9 - 0.6
Coelomycete sp. 1 - - - - - - 2.8 - 1.4 2.2 0.9 - 0.6
Trichoderma sp. 2.6 3.2 - - - - 1.4 - - - - - 0.6
Aquaticola tropica - - - - - 2 1.4 3.6 - - - - 0.6
Jahnula systyla - - 0.8 - - 1 1.4 - - - 3.6 - 0.6
Xylomyces giganteus - - - - - - - - 5.6 1.1 - - 0.6
Halosarpheia lotica - 1.6 5 - - - - - - - - - 0.5
Massarina peerallyi - - - - - 3 - 0.9 0.7 - - 2 0.5
Tamsiniella labiosa 4.4 - - - - - - - 2.1 - - - 0.5
Canalisporium variabile - - - 0.9 2.2 1 - - 2.1 - - - 0.5
Ophioceras commune 0.9 1.6 0.8 - - - - - 0.7 1.1 0.9 - 0.5
Neta angliae - - - - - - - - - - - 5.9 0.5
Sporoschisma juvenile - - - 1.9 1.1 2 - - - - 0.9 - 0.5
Sporormiella sp. - - 5.8 - - - - - - - - - 0.5
Pseudohalonectria lignicola - - 4.2 - - - - 0.9 0.7 - - - 0.5
Sporoschisma saccardoi - - - - 1.1 - - - - 3.4 0.9 - 0.4
Lophiostoma lunisporum 1.8 1.6 - 0.9 - 1 - - - - - - 0.4
Bactrodesmium longisporum - - - 0.9 3.3 - - - - - 0.9 - 0.4
Coniochaeta sp. - - - - - 5 - - - - - - 0.4
Dictyosporium elegans - - - - - - - - - - - 4.9 0.4
Quintaria lignatilis - - - 1.9 - 3 - - - - - - 0.4
Aquasphaeria dimorphospora - - - 0.9 2.2 1 - - 0.7 - - - 0.4
Coelomycete sp. 2 - - - - - 2 - 2.7 - - - - 0.4
Jahnula aquatica - - 0.8 0.9 - - 2.8 - - - - - 0.4
Annulatascus fusiformis - - - 0.9 3.3 - - - - - - - 0.4
Dictyosporium sp. - 1.6 1.7 - - - - - 0.7 - - - 0.3
Trematosphaeria confusa - - 0.8 - 2.2 - - 0.9 - - - - 0.3
Dactylaria uniseptata - - - - - - - - - - - 3.9 0.3
Hyphomycete sp. - - - - 2.2 1 - - 0.7 - - - 0.3
Ellisembia opaca - - - - - - - - - 1.1 2.7 - 0.3
Micropeltopsis quinquecladiopsis - - - - - - - - - 1.1 2.7 - 0.3
Clohesyomyces sp. - - - - - 1 - 2.7 - - - - 0.3
Digitodesmium recurvum - - - - - - - - - - 3.6 - 0.3
Cryptophiale multiseptata - - 0.8 - 1.1 - - 0.9 0.7 - - - 0.3
Nectria sp. - - - - - - - - 3.5 - - - 0.3
Helicosporium gigaspora - - - - 1.1 1 1.4 - - - - - 0.3
Helicosporium griseum - 1.6 0.8 0.9 - - - - - - - - 0.3

212
Table 3. Average % abundance of lignicolous freshwater fungi in all the 12 sites.
Species Name CP CR CV DP DR DV ER EV KP KR TR TV Average %
abundance
Rivulicola aquatica - - 3.3 - - - - - - - - - 0.3
Savoryella longispora 0.9 1.6 0.8 - - - - - - - - - 0.3
Helicosporium decumbens - - - - - - - - 2.1 1.1 - - 0.3
Sporoschisma uniseptatum 0.9 - - - 1.1 1 - - - - - - 0.2
Ellisembia opaca - - - - - - - - - - - 2.9 0.2
Tiarosporella paludosa - - - - - - - - - - - 2.9 0.2
Lophiosphaeria sp. - - - 0.9 - 2 - - - - - - 0.2
Ophioceras arcuatisporum - - - 1.9 - 1 - - - - - - 0.2
Melanomma sp. - - - - - - 2.8 - - - - - 0.2
Cancellidium applanatum 0.9 - - 1.9 - - - - - - - - 0.2
Herpotrichia sp. - - - 0.9 1.1 - - - 0.7 - - - 0.2
Stilbella fusca - 1.6 - - - - - - - 1.1 - - 0.2
Lophiostoma frondisubmersum 0.9 - 0.8 0.9 - - - - - - - - 0.2
Paraniesslia tuberculata - - - 0.9 - - - - 0.7 - 0.9 - 0.2
Massarina corticola 0.9 1.6 - - - - - - - - - - 0.2
Nais aquatica - - - - - - - - 1.4 - 0.9 - 0.2
Vaginatispora aquatica - - - - - - 1.4 0.9 - - - - 0.2
Annulatascus lamtuensis - - - - - - - - - 2.2 - - 0.2
Phaeosphaeria culmorum - - - - - - - - - 2.2 - - 0.2
Chloridium sp. - - - - 2.2 - - - - - - - 0.2
Ophioceras venezuelense - - - - - - - - 2.1 - - - 0.2
Helicomyces torquatus - - - - 1.1 1 - - - - - - 0.2
Paecilomyces sp. - - - 0.9 - - - - - 1.1 - - 0.2
Torrentispora fibrosa - - - 0.9 1.1 - - - - - - - 0.2
Aquaticola rhomboidea - - - - - - - - - 1.1 0.9 - 0.2
Glomerella sp. - - - - - - - - - 1.1 0.9 - 0.2
Nectria cinnabarina - - - - - - - - - 1.1 0.9 - 0.2
Hymenoscyphus varicosporoides - - - - - - - - - - - 2 0.2
Aquaticola lignicola - - - 0.9 - 1 - - - - - - 0.2
Annulatascus aquaticus - - - 1.9 - - - - - - - - 0.2
Aquaticola lutea - - - 1.9 - - - - - - - - 0.2
Chalara sp. 0.9 - - 0.9 - - - - - - - - 0.2
Halosarpheia viscosa - - - - - 1 - - 0.7 - - - 0.1
Cataractispora aquatica - - - 0.9 - - - - 0.7 - - - 0.1
Immersiella immersa - - - - - - - - 0.7 - 0.9 - 0.1
Sporidesmium hyalospermum - 1.6 - - - - - - - - - - 0.1
Monotosporella sphaerocephala - - 0.8 - - - - - 0.7 - - - 0.1
Ascomycete sp. - - - - - - - - 1.4 - - - 0.1
Astrosphaeriella maquilingiana - - - - - - - - 1.4 - - - 0.1
Cataractispora bipolaris - - - - - - - - 1.4 - - - 0.1
Lollipopaia sp. - - - - - - - - 1.4 - - - 0.1

Fungal Diversity
213
Table 3. Average % abundance of lignicolous freshwater fungi in all the 12 sites.
Species Name CP CR CV DP DR DV ER EV KP KR TR TV Average %
abundance
Astrosphaeriella trochus - - - - - - 1.4 - - - - - 0.1
Clohesia aquatica - - - - - - 1.4 - - - - - 0.1
Phaeonectriella appendiculata - - - - - - 1.4 - - - - - 0.1
Pseudoproboscispora caudae-suis - - - - - - 1.4 - - - - - 0.1
Halosarpheia aquadulcus - - - - - - - - - 1.1 - - 0.1
Helicoon gigantisporum - - - - - - - - - 1.1 - - 0.1
Tiarosporella sp. - - - - - - - - - 1.1 - - 0.1
Annulatascus tropicalis - - - - 1.1 - - - - - - - 0.1
Dictyosporium giganticum - - - - 1.1 - - - - - - - 0.1
Diplodia sp. - - - - 1.1 - - - - - - - 0.1
Glonium sp. - - - - 1.1 - - - - - - - 0.1
Iodosphaeria aquatica - - - - 1.1 - - - - - - - 0.1
Melanomma australiense - - - - 1.1 - - - - - - - 0.1
Saccardoella minuta - - - - 1.1 - - - - - - - 0.1
Ascotaiwania pallida - - - - - 1 - - - - - - 0.1
Massarina aquatica - - - - - 1 - - - - - - 0.1
Oxydothis sp. - - - - - 1 - - - - - - 0.1
Sporidesmium sp. - - - - - 1 - - - - - - 0.1
Sporormiella minima - - - - - 1 - - - - - - 0.1
Sporoschisma parcicuneatum - - - - - 1 - - - - - - 0.1
Aqualignicola hyalina - - - 0.9 - - - - - - - - 0.1
Aquaticola trisepata - - - 0.9 - - - - - - - - 0.1
Candelosynnema ranunculosporum - - - 0.9 - - - - - - - - 0.1
Cataractispora viscosa - - - 0.9 - - - - - - - - 0.1
Dictyochaeta daphnioides - - - 0.9 - - - - - - - - 0.1
Gonytrichum caesium - - - 0.9 - - - - - - - - 0.1
Massarina purpurascens - - - 0.9 - - - - - - - - 0.1
Sporoschismopsis australiensis - - - 0.9 - - - - - - - - 0.1
Dictyosporium digitatum - - - - - - - 0.9 - - - - 0.1
Monodictys putredinis - - - - - - - - - - 0.9 - 0.1
Digitodesmium elegans 0.9 - - - - - - - - - - - 0.1
Spadicoides sp. 0.9 - - - - - - - - - - - 0.1
Janetia curviapices - - 0.8 - - - - - - - - - 0.1
Phaeosphaeria sylvatica - - 0.8 - - - - - - - - - 0.1
Phomatospora aquatica - - 0.8 - - - - - - - - - 0.1
Ascitendus austriaca - - - - - - - - 0.7 - - - 0.1
Eutypa sp. - - - - - - - - 0.7 - - - 0.1
Lasiosphaeria sp. - - - - - - - - 0.7 - - - 0.1
Phomatospora berkeleyi - - - - - - - - 0.7 - - - 0.1
Pleurophragmium malayense - - - - - - - 0.7 - - - 0.1

214
Clohesy creek agricultural zone (CV), Kauri pristine (KP) and Tinaroo
agricultural (TV) were quite far removed from the rest of the collection (Fig.
3). In ecological terms, fungal assemblages on submerged wood in the sites
within the same stream, had more similarity and the type of riparian zone had
lesser effect on the species assemblages.
Discussion
Number of species
This is the first large-scale study (12 sites spread across 5 streams) of
lignicolous freshwater fungi in the wet heritage tropical forests of Australia.
An average of 40 species was collected from a total of 50 woody substrates at
each site. Within a given stream, higher numbers of species were observed at
pristine zones, with 35, 61 and 56 at the Clohesy (CP), Davies (DP) and Kauri
(KP) sites respectively (Table 2). Earlier, studies at Mt. Lewis and Lake
Barrine, Queensland, Australia (Hyde and Goh, 1997, 1998a) produced similar
results. Several studies of freshwater fungi, have been carried out on naturally
occurring submerged woody substrates around the world, and the numbers of
taxa identified from single collections varies from as low as 28 taxa from River
Coln, Britain (Hyde and Goh, 1998a) to more than 100 fungal taxa from the
USA (Shearer and Crane, 1986) and Hong Kong (Ho et al., 2001, Tsui et al.,
2000). Differences in the number of taxa recorded have been attributed to
geographical variations such as temperate/tropical conditions or isolation of
islands, physical attributes such as temperature, concentration of dissolved
oxygen, perennial state of the water body, size of the stream, degree of shading
by riparian vegetation (Ho et al., 2001, 2002), and period of incubation of
collected substrates (Tsui et al., 2000). The lower biodiversity in tropical
Australia streams, in comparison to other sub-tropical regions, may be because
sub-tropical regions can harbor both temperate and tropical species (Ho et al.,
2001).
Number of fungi per sample
In general, species richness declines with increase in habitat
modification. In this study, there was a remarkable difference in species
richness at different sites on the same stream and a general trend was observed
in all streams: species richness was highest in the pristine zone (average taxa
per woody substrate, 2.42), where shading by riparian vegetation and organic
input was highest, whereas regrowth zones, which had moderate riparian cover
and moderate organic input, had the lowest species richness (average taxa per

Fungal Diversity
215
Fig. 2. Species-abundance distributions of lignicolous freshwater fungi collected from various
sites. See Table 2 for abbreviation.
woody substrate, 1.71). The reduction of species richness is positively
correlated with the reduction of organic input. However, the agricultural zone
with the least riparian cover had a higher species richness (average taxa per
woody substrate, 2.16) than the regrowth zone. The increase in species richness
at the agricultural zones may be attributed to the replacement of organic input
from allochthonous to autochthonous sources, as there is more light, due to the
reduced riparian cover, which enables growth of plants in the stream
(Bärlocher, 1992). Alternatively, due to the presence of herbaceous plants and
the reduced speed of water currents, there is an increased trapping of organic
material, therefore increasing fungal species richness. These results should be
interpreted with caution, due to the lack of information on water quality at the
time of collection. Given that the data collected were at the same geographical
location in the same season, the likelihood that geographical separation,
isolation, and seasonal changes may have effect on the species richness of
freshwater fungi, is minimal.

216
Table 4. Sorenson’s similarity index showing similarity indices within 12 sites.
Shaded values represent similarities greater than 25%.
CP CR CV DP DR DV ER EV KP KR TR TV
CP 0.31 0.18 0.19 0.15 0.17 0.10 0.14 0.15 0.14 0.05 0.10
CR
0.26 0.17 0.15 0.19 0.13 0.13 0.16 0.17 0.15 0.05
CV 0.13 0.10 0.14 0.10 0.14 0.14 0.11 0.08 0.10
DP
0.25 0.27 0.13 0.10 0.17 0.18 0.21 0.11
DR
0.25 0.14 0.14 0.18 0.17 0.22 0.13
DV 0.18 0.15 0.15 0.15 0.18 0.14
ER 0.19 0.14 0.18 0.13
0.36
EV 0.16 0.17 0.09 0.09
KP 0.19 0.13 0.08
KR 0.23 0.12
TR 0.16
TV
Species abundance
The relative abundance distribution describes how the individual in a
community are partitioned among rare and common species (Fig. 2). A
community containing many rare species and relatively few common species is
associated with a community in equilibrium. Likewise, an equitable share of
individuals amongst species groups is associated with non-equilibrium
behaviour resulting from perturbation due to disturbance, pollution or
immigration (Pachepsky et al., 2001). A total of 29 taxa were found dominant
in more than 2 sites. All sites with the exception of Tinaroo Creek agricultural
zone (TV) had a majority of rare taxa (Fig. 2). Higher evenness was recorded
at TV (Fig. 2). At TV, a majority of the wood substrates collected were black,
covered with carbon, which might be due to a wildfire near that region
(Vijaykrishna pers. obs), which can be attributed to the non-equilibrium or high
evenness of the fungal communities at the site.
Diversity
Shannon’s index of diversity is the most popular choice of diversity
index for various biodiversity studies, and has been mainly used for studying
freshwater fungal communities. However, when the randomness of the sample
cannot be guaranteed (Southwood and Henderson, 2000), or if the community
is completely censured and every individual accounted for, the Brillouin index,
is the appropriate form of the information index (Pielou, 1969, 1975) and has
often been used (Lo et al., 1998; Dans et al., 1999; Ito and Imai, 2000). In this

Fungal Diversity
217
study, at the Emerald creek regrowth (ER) site, the number of wood pieces was
comparatively lesser than the other sites. This might have been due to rapid
flow of water from the waterfall at its immediate vicinity, and therefore
random selection of samples was impossible. Brillouin’s index (Magurran,
2004) was therefore deemed appropriate for this study. Brillouin’s index of
diversity shows the diversity of the actual collection, therefore statistical tests
are not necessary (Magurran, 2004), and is mathematically superior (Laxton,
1978).
The uniqueness of Kauri Creek pristine zone (KP), is shown by a higher
diversity, which is directly proportional to the highest species richness. In
comparison to other sites, KP was at the highest altitude, with comparatively
higher riparian vegetation and shade. The extent of shade provided by the
riparian vegetation has shown to affect invertebrate community (Dudgeon,
1989; Dudgeon and Corlett, 1994) and also fungi (Tsui et al. 2000).
Correspondence analysis, involving fungal species and their frequencies
show higher similarity of taxa along the same stream (Fig. 3a,b), especially in
the Clohesy River, Davies Creek and Emerald Creek. This is also due to high
species similarity between the sites on the same stream (Table 4). Due to the
unidirectional movement of water, the upstream communities (pristine) should
have an influence on species composition of the downstream communities
(regrowth, agricultural).
Composition of fungal groups
One of the most striking results of this study when compared to earlier
studies of lignicolous freshwater fungi in both tropical and temperate regions is
the teleomorph anamorph ratio. A surprisingly high number of ascomycetes
(teleomorphic fruiting bodies) as compared to anamorphic spores (101: 59)
were recorded from submerged samples in this study. In a comparative study of
fungal communities on submerged wood in tropical regions and sub tropical
regions, Ho et al., (2001) showed that, except in Malaysia, the number of
ascomycetes were always lower than the anamorphic taxa in submerged
samples. However the reason for the ratio of more teleomorphic stages in this
study as compared to other studies in freshwater habitats is unknown. Shearer
(1992) commented that wood may play an important role as a site for genetic
recombination in aquatic hyphomycetes, and a number of sexual states for
these fungi have been found on wood (Webster, 1992).
Among the teleomorphs, pyrenomycetes and loculoascomycetes
dominate the ascomycete assemblage, with representatives in the families
Annulatascaceae, Halosphaeriaceae and Lophiostomataceae. It is not
surprising, that in this study only one discomycete (Hymenoscyphus

218
Fig. 3. Three dimentional correspondence
ordination of taxa and fungal communities
recorded from submerged wood from
various sites. For site codes see Fig. 2.
varicosporoides) was recorded from Tinaroo Agricultural. Out of a total 111
discomycetes recorded from submerged wood worldwide, only
Hymenoscyphus malawiensis, Pezoloma rhodocarpus, Cudoniella indica,
Saccobolus beckii (Fisher and Spooner, 1987; Udaiyan, 1989; Webster et al.,
1995a) and more recently Hymenoscyphus varicosporoides (Sivichai et al.,
2003) have been reported from the tropics. Hyde et al., (1997) commented that
the lack of discomycetes in tropical streams is most striking. It has been argued

Fungal Diversity
219
that most of the discomycetes from temperate regions occur on graminaceous
substrata (Shearer, 1993), and those substrates have been poorly studied in the
tropics. However recently, studies on grasses and sedges in freshwater habitats
of Hong Kong yielded no discomycetes (Wong and Hyde, 2001), indicating
scarcity of this group of fungi in the tropics (Cai et al., 2003b).
The interpretation of lack of discomycetes in freshwater habitats should
be noted with caution, as studies on freshwater fungi in the tropics have largely
been conducted on naturally occurring submerged wood and wood baits
immersed in streams. Sivichai and Jones (2003) noted that the majority of
Ingoldian fungi (hyphomycetes actively growing and sporulating under water)
have ascomycete connections, and with the highest species number in the
genus Hymenoscyphus (discomycete). Furthermore, it has been shown that
after several months of incubation discomycetes, produce their teleomorphic
fruiting structures on wood (Sivichai et al., 2003). A large number of Ingoldian
fungi are found on submerged leaves and spores trapped in air bubbles in
streams (Bärlocher, 1992), and many have been recorded from tropical and
subtropical regions (Bhat and Chien, 1990; Au et al., 1992a,b; Chan et al.,
2000a,b). Sivichai et al., (2003) suggested the use of different techniques to
survey for tropical freshwater discomycetes might yield a wide range of
species.
Impact of riparian vegetation
Statistical comparison of species from different sites using the
Multivariate Correspondence analyses (Figs. 4a,b) does not provide any clear
evidence of impact of riparian vegetation on the species composition.
However, the similarity in species composition within streams is recognised,
which is also shown in the Sørenson’s similarity index (Table 4), with high
similarity between species isolated from different sites from the same stream.
This shows that fungal species of the higher reaches of the stream, obviously
have an effect on the establishment of fungal communities down stream.
Furthermore, the frequent identification of apparently same fungal taxa from
different riparian zones suggests that riparian habitat preference of individual
taxa is minimum. Major changes in overall abundance, and species richness
was observed. A general trend with decrease of species abundance and richness
with increased anthropogenic changes on the riparian zone is observed.
Conclusions
To examine the effects of riparian vegetation on taxonomic composition
and distribution of lignicolous freshwater fungi, we studied fungal

220
communities in three different kinds of riparian vegetation. Results indicate
that the type of riparian vegetation is a factor that determines species richness
of lignicolous freshwater fungi. A clear trend was observed in all streams.
There was a decline in species richness with increased habitat exploitation.
Decomposition of organic matter has shown to decrease with decline in species
richness, which might indirectly affect the overall biodiversity of the
ecosystem (McGrady-Steed et al., 1997). Therefore, these results provide
additional reasons for conserving species richness in relatively intact
ecosystems and restoring diversity in degraded systems. An extension of the
results obtained through this study would be to understand the effects of
decline in fungal species richness to other communities, especially organisms
that feed on woody debris and other organic matter (e.g. invertebrates). Also to
further understand the reasons for variation in species richness manipulative
experiments involving the terrestrial detritus input would provide valuable
information regarding stream ecology.
In this study we also found that the type of riparian vegetation has little
effect on the taxonomic composition of lignicolous freshwater fungi. Similarity
between different sites of the same stream demonstrate that the fungal
communities from the upper reaches of the stream have obvious effects in
determining the taxonomic composition of downstream communities. It is
intriguing, how the upper reaches of the stream continue to maintain fungal
populations, with a constant drainage of spores and hyphae downstream, due to
the unidirectional rapid flow of water in small streams (Bärlocher, 1992).
Comparative studies of fungal assemblages from both submerged wood within
the stream and decaying wood beside the stream would probably provide
evidence of a vessel for the continued occurrence of fungal assemblages in the
upper reaches of the stream.
Most dominant fungal taxa identified in this study belonged to the
families Annulatascaceae and Lophiostomataceae. The diversity estimates of
the present study relied on the production of fruiting structures under
incubation, which might, however, underestimate the actual species
composition present on submerged wood. This is because certain fungi may
not sporulate when removed from their natural habitat. PCR based
identification methods, which have primarily been used for bacteria, has gained
popularity in fungal ecology (Nikolcheva et al., 2003). Future studies involving
the combination of traditional methods and PCR based methods (e.g. DGGE)
should be used in order to provide additional insight in to the ecology of fungi
on submerged wood.

Fungal Diversity
221
Acknowledgements
We thank Environment Australia (Queensland, Australia) for providing permits to
collect at the State and National parks. Dr. C. Pearce is thanked for her invaluable help for
obtaining collection permits and sampling. Dr. C.K.M. Tsui and L. Cai for advice on the
identification of ascomycetes. Dr. W.H. Ho, A.M.C. Tang are thanked for the help with
statistics. D.V acknowledges The University of Hong Kong for a postgraduate studentship.
References
Abdel-Raheem, A. and Shearer, C.A. (2002). Extracellular enzyme production by freshwater
ascomycetes. Fungal Diversity 11: 1-19.
Anon. (1995). JMP® Statistics and Graphics Guide. Version 3.1. SAS Institute, Cary NC.
Au, D.W.T., Hodgkiss, I.J., and Vrimoed, L.L.P. (1992a). Decomposition of Bauhinia
purpurea leaf litter in a polluted and unpolluted Hong Kong waterway. Canadian
Journal of Botany 70: 1061-1070.
Au, D.W.T., Hodgkiss, I.J., and Vrimoed, L.L.P. (1992b). Fungi and cellulolytic activity
associated with decomposition of Bauhinia purpurea leaf litter in a polluted and
unpolluted waterway. Canadian Journal of Botany 70: 1071-1079.
Bärlocher, F. (1992). The Ecology of Aquatic Hyphomycetes. Berlin: Springer-Verlag.
Barr, M. E. (1979). A classification of Loculoascomycetes. Mycologia 71: 935-957.
Benson, L.J. and Pearson, R.G. (1993). Litter inputs to a tropical Australian rainforest stream.
Australian Journal of Ecology 18: 377-383.
Bhat, D.J. and Chien, C.Y. (1990). Water-Borne hyphomycetes found in Ethiopia. Transactions
of the Mycological society of Japan 31: 147-158.
Bucher, V.V.C., Hyde, K.D., Pointing, S.B., and Reddy, C.A. (2004). Production of wood
decay enzymes mass loss and lignin solubilization in wood by marine ascomycetes and
their anamorphs. Fungal Diversity 15: 1-14.
Cai, L., Lumyong, P., Zhang, K.Q. and Hyde, K.D. (2002a). New species of Annulatascus and
Saccardoella from the Philippines. Mycotaxon 84: 255-263.
Cai, L., Tsui, C.K.M., Zhang, K.Q., and Hyde, K.D. (2002b). Aquatic fungi from Lake Fuxian,
Yunnan, China. Fungal Diversity 9: 57-70.
Cai, L., Zhang, K.Q., McKenzie, E.H.C. and Hyde, K.D. (2003a). Freshwater fungi from
bamboo and wood submerged in the Liput River in the Philippines. Fungal Diversity
13: 1-12.
Cai, L., Zhang, K.Q. and Hyde, K.D. (2003b). Freshwater Ascomycetes. Fungal Diversity
Research Series 10: 275-324.
Cai, L., Zhang, K.Q. and Hyde, K.D. (2005). Ascoyunnania chameleonia gen. et sp. nov. a
freshwater fungus collected from China and its microcyclic conidiation. Fungal
Diversity 18: 1-8
Chan, S.Y., Goh, T.K, and Hyde, K.D. (2000a). Ingoldian fungi in Hong Kong. Fungal
Diversity 5: 89-107.
Chan, S.Y., Goh, T.K, and Hyde, K.D. (2000b). Ingoldian fungi in Lam Tsuen River and Tai
Po Kau Forest Stream, Hong Kong. Fungal Diversity 5: 109-118.
Dans, S.L., Reyes, L.M., Pedraza, S.N., Raga, J.A. and Crespo, E.A. (1999). Gastrointestinal
helminths of the dusky dolphin, Lagenorhynchus obscurus (Gray, 1928), off Patagonia
in the southwestern Atlantic. Marine Mammal Science 15: 649-660.
Díez, J., Elosegi, A., Chauvet, E. and Pozo, J. (2002). Breakdown of wood in the Agera stream.

222
Freshwater Biology 47: 2205-2215.
Dolloff, C.A. and Webster, J.R. (2000). Particulate organic contributions from forests to
streams: debris isn't so bad. In: Riparian Management in Forests of the Continental
eastern United States (eds. E.S. Verry, J.W. Hombeck and C.A. Dolloff). Lewis
Publishers, Boca Raton, Florida: 125-138.
Dudgeon, D. (1989). The influence of riparian vegetation on the functional organization of four
Hong Kong stream communities. Hydrobiologia 179:183-194.
Dudgeon, D. (1999). Tropical Asian Streams. Hong Kong University Press, Hong Kong.
Dudgeon, D. (2000). The ecology of tropical Asian rivers and streams in relation to
biodiversity conservation. Annual Review of Ecology and Systematics 31: 239-263.
Dudgeon, D. and Corlett, R. (1994). Hills and Streams: an ecology of Hong Kong. Hong Kong
University Press, Hong Kong.
Fisher, P.J. and Spooner, B. (1987). Two new ascomycetes from Malawi. Transactions of the
British Mycological Society 75: 201-206.
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.
Fryar, S.C., Booth, W., Davies, J., Hodgkiss, I.J. and Hyde, K.D. (2005). Evidence of in situ
competition between fungi in freshwater. Fungal Diversity 18: 59-71.
Goh, T.K. and Hyde, K.D. (1999). Fungi on submerged wood and bamboo in the Plover Cove
Reservoir, Hong Kong. Fungal Diversity 3:57-85.
Goh, T.K. and Tsui, C.K.M. (2003). Key to common dematiaceous hyphomycetes from
freshwater. Fungal Diversity Research Series 10: 325-343.
Gönczöl, J. and Révay, Á (2004). Fungal spores in rainwater: stemflow, throughfall and gutter
conidial assemblages. Fungal Diversity 16: 67- 86.
Harmon, M.E., Franklin, J.F., Swanson, F.J., Sollins, O., Gregory, S.V., Lattin, J.D., Anderson,
N.H., Cline, S.P., Aumen, N.J., Sedell, J.R., Lienkaemper, G.W., Cromack, K. and
Cummins, K.W. (1986). Ecology of coarse woody debris in temperate ecosystems.
Advances in Ecological Research 15: 133-302.
Hicks, B.J. (1997). Food webs in forest and pasture streams in the Waikato region: a study
based on analyses of stable isotopes of carbon and nitrogen, and fish gut contents. New
Zealand Journal of Marine and Freshwater Research 31:651-664.
Ho, W.H., Hyde, K.D., Hodgkiss, I.J. and Yanna (2001). Fungal communities on submerged
wood from streams in Brunei, Hong Kong and Malaysia. Mycological Research 105:
1492-1501.
Ho, W.H., Yanna, Hyde, K.D. and Hodgkiss, I.J. (2002). Seasonality and sequencial
occurrence of fungi on wood submerged in Tai Po forest stream, Hong Kong. Fungal
Diversity 10:21-43.
Hyde, K.D. and Goh, T.K. (1997). Fungi on submerged wood in a small stream on Mt Lewis,
North Queensland, Australia. Muerllera 10: 145-157.
Hyde, K.D. and Goh, T.K. (1998a). Fungi on submerged wood in Lake Barrine, North
Queensland, Australia. Mycological Research 102: 739-749.
Hyde, K.D. and Goh, T.K. (1998b). Fungi on submerged wood in the Riviere St Marie-Louis,
The Seychelles. South African Journal of Botany 64: 330-336.
Hyde, K.D. and Goh, T.K. (1999a). Fungi on submerged wood from River Coln, England.
Mycological Research 103:1561-1574.
Hyde, K.D., Wong, S.W. and Jones, E.B.G. (1997). Freshwater ascomycetes. In: Biodiversity
of Tropical Microfungi (ed. K.D. Hyde). Hong Kong University Press, Hong Kong:
179-189.
Ito, A. and Imai, S. (2000). Ciliates from the cecum of capybara (Hydrocheorus hydrochaeris)

Fungal Diversity
223
in Bolivia 2. The family Cycloposthiidae. Europian Journal of Protistology 36: 169-200.
Keller, E.A. and Swanson, F.J. (1979). Effects of large organic material on channel form and
fluvial processes. Earth Surface Processes 4: 361-380.
Kenkel, N.C. and Booth, T. (1992). Multivariate analysis in fungal ecology. In: The fungal
Community: its organization and role in the ecosystem. 2nd edn. (eds. G.C. Carroll and
D.T. Wicklow). Marcel Dekker, New York: 209-227
Laxton, R.R. (1978). The measure of diversity. Journal of Theoretical Biology 70: 51-67.
Lo, C.M., Morand, S. and Galzin, R. (1998). Parasite diversity and size relationship in three
coral-reef fishes from French Polynesia. International Journal of Parasitology 28: 1695-
1708.
Lodge, D.J. (1997). Factors related to diversity of decomposer fungi in tropical forests.
Biodiversity and Conservation 6: 681-688.
Ludwig, J.A. and Reynolds, F.F. (1998). Statistical Ecology: a primer on methods and
computing. John Wiley and Sons, Inc. New York.
Luo, J., Yin, J.F., Cai, L., Zhang, K. and Hyde, K.D. (2004). Freshwater fungi in Lake Dianchi,
a heavily polluted lake in Yunnan, China. Fungal Diversity 16: 93-112.
Magurran, A.E. (2004). Measuring Biological Diversity. Blackwell Science, Oxford.
McGrady-Steed, J., Harris, P.M. and Morin, P.J. (1997). Biodiversity regulates ecosystem
predictability. Nature 390: 162-165.
Mosley, P.M. (1981). The influence of organic debris on channel morphology and bedload
transport in a New Zealand forest stream. Earth surface Processes and Landforms 6:
571-579.
Nikolcheva, L.G., Cockshutt, A.M. and Bärlocher, F. (2003). Determining diversity of
freshwater fungi on decaying leaves: comparison of traditional and molecular
approaches. Applied and Environmental Microbiology 69: 2548-2554.
Pachepsky, E., Crawford, J.W., Brown, J.L. and Squire, G. (2001). Towards a general theory of
biodiversity. Nature 410: 923-926.
Pascoal, C., Marvanová, L. and Cássio, F. (2005). Aquatic hyphomycete diversity in streams of
Northwest Portugal. Fungal Diversity 19: 109-128.
Pielou, E.C. (1969). An Introduction to Mathematical Ecology. Wiley, New York.
Pielou, E.C. (1975). Ecological Diversity. Wiley InterScience, New York.
Sedell, J.R., Bisson, P.A., Swanson, F.J. and Gregory, S.V. (1988). What we know about large
trees that fall in streams and rivers. In: From the Forest to the Sea: a story of fallen
trees (eds. C. Maser, R.F. Tarrant, J.M. Trappe, and J.F. Franklin). General Technical
Report PNW-GTR-229, USDA Forest Service, Pacific northwest Research Station; U.S.
Department of the Interior Bureau of Land Management, Portland, Oregon: 47-81.
Sivichai, S. and Jones, E.B.G. (2003). Teleomorphic-anamorphic connections of freshwater
fungi. Freshwater mycology. Fungal Diversity Research Series 10: 259-274.
Sivichai, S., Jones, E.B.G. and Hewyl-Jones, N.L. (2003). Lignicolous freshwater Ascomycota
from Thailand: Hymenoscyphus varicosporoides and its Tricladium anamorph.
Mycological Research 95: 340-346.
Shearer, C.A. (1992). The role of woody debris. In: The Ecology of Aquatic Hyphomycetes (ed.
F. Bärlocher). Springer Verlag, Berlin: 1-77.
Shearer, C.A. (1993). The freshwater ascomycetes. Nova Hedwigia 56: 1-33.
Shearer, C.A. (2001). The distribution of freshwater filamentous ascomycetes. In:
Trichomycetes and other Fungal Groups: Robert W. Lichtwardt Commemoration (eds.
J.K. Misra and B.W. Horn). Science Publishers, Inc. Enfield, New Hampshire: 225-292.
Shearer, C.A. and Crane, J.L. (1986). Illinios fungi XII Fungi and Myxomycetes from wood
and leaves submerged in a freshwater swamp. Mycotaxon 25: 527-538.

224
Southwood, R. and Henderson, P.A. (2000). Ecological Methods. Blackwell Science, Oxford.
Sørenson, T. (1948). A method of establishing groups of equal amplitude in plant sociology
based on similarity of species content and its application to analyses of the vegetation
on Danish commons. Biologiske Skrifter Kongelige Danske Videnskabernes Selskab 5:
1-34.
Suberkropp, K. (1997). Annual production of leaf-decaying fungi in a woodland stream.
Freshwater Biology 38: 169-178.
Tsui, C.K.M. and Hyde, K.D. (2004). Biodiversity of fungi on submerged wood in a stream
and estuary in the Tai Ho Bay, Hong Kong. Fungal Diversity 15: 205-220.
Tsui, C.K.M., Hyde, K.D, and Hodgkiss, I.J. (2000). Biodiversity of fungi on submerged wood
in Hong Kong streams. Aquatic Microbial Ecology 21: 289-298.
Udaiyan, K. (1989). Some interesting ascomycetes from water-cooling towers. Kavaka 17: 11-
16.
Van Ryckegem, G. and Verbeken, A. (2005). Fungal ecology and succession on Phragmites
australis in a blackish tidal marsh. I Leaf sheaths. Fungal Diversity 19: 157-187.
Van Ryckegem, G. and Verbeken, A. (2005). Fungal ecology and succession on Phragmites
australis in a blackish tidal marsh. II Stems. Fungal Diversity 20: 209-233.
Wallace, J.B., Eggert, S.L., Meyer, J.L. and Webster, J.R. (1997). Multiple trophic levels of a
forest stream linked to terrestrial litter inputs. Science 277: 102-104.
Webster, J. (1992). Anamorph-teleomorph relationships. In: The Ecology of Aquatic
Hyphomycetes (ed. F. Bärlocher). Springer Verlag, Berlin Press, Berlin: 99-117.
Webster, J., Eicher, A. and Spooner, B.M. (1995a). Cudoniella indica sp. nov. (Ascomycetes,
Leotiales), the teleomorph of Tricladium indicum, an aquatic fungus isolated from a
South African river. Nova Hedwigia 60: 493-498.
Webster, J.R., Wallace, J.B. and Benfield, E.F. (1995b). Organic processes in streams of the
eastern United States. In: Ecosystems of the World, Vol. 22. River and Stream
Ecosystems (eds. C.E. Cushing, K.W. Cummins and G.W. Minshall). Elsevier,
Amsterdam: 117-187.
Webster, J.R., Benfield, E.F., Ehrman, T.P., Schaeffer, M.A., Tank, J.L., Hutchens, J.J. and
DíAngelo, D.J. (1999). What happens to allochthonous material that falls into streams?
A synthesis of new and published information from Coweeta. Freshwater Biology 41:
687-705.
Wetzel, R.G. (2001). Limnology, Lake and River Ecosystems. Academic Press, San Diego, CA.
Wong, S.W. and Hyde, K.D. (2001). Diversity of fungi on six species of Gramineae and one
species of Cyperaceae in Hong Kong. Mycological Research 105: 1485-1491.
(Received 15 December 2005; accepted 25 January 2006)