Mycology
An International Journal on Fungal Biology
ISSN: 2150-1203 (Print) 2150-1211 (Online) Journal homepage: https://www.tandfonline.com/loi/tmyc20
Fungal assemblage and leaf litter decomposition
in riparian tree holes and in a coastal stream of
the south-west India
Kandikere R. Sridhar, Kishore S. Karamchand & Sahadevan Seena
To cite this article: Kandikere R. Sridhar, Kishore S. Karamchand & Sahadevan Seena (2013)
Fungal assemblage and leaf litter decomposition in riparian tree holes and in a coastal stream of
the south-west India, Mycology, 4:2, 118-124, DOI: 10.1080/21501203.2013.825657
To link to this article: https://doi.org/10.1080/21501203.2013.825657
Copyright Mycological Society of China
Published online: 12 Aug 2013.
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Mycology, 2013
Vol. 4, No. 2, 118–124, http://dx.doi.org/10.1080/21501203.2013.825657
Fungal assemblage and leaf litter decomposition in riparian tree holes and in a coastal stream of
the south-west India
Kandikere R. Sridhara*, Kishore S. Karamchanda and Sahadevan Seenab
a
Department of Biosciences, Mangalore University, Mangalagangotri 574 199, Mangalore, India; b Department of Biology, Centre of
Molecular and Environmental Biology, University of Minho, Campus de Gualtar 4710-057, Braga, Portugal
(Received 8 July 2013; final version received 12 July 2013)
Assemblage of aquatic hyphomycetes and decomposition of banyan leaf litter (Ficus benghalensis) were assessed
in riparian tree holes and in a tropical coastal stream during rainy season up to 8 weeks in south-west coast
of India. Although fungal assemblage was similar, leaf decomposition and leaf chemical changes differed between
tree holes and the stream. Out of the 18 aquatic hyphomycetes, 17 were common on leaf litter in tree holes and
the stream. Anguillospora longissima, Flagellospora curvula, Lunulospora curvula, Triscelophorus acuminatus and
T. konajensis were the top five species in tree holes and the stream. The species richness was the highest during the second week in tree holes (11 species) and the sixth week in the stream (14 species), while the conidial output was the highest
in tree holes and the stream during sixth week. The daily decay coefficient (k) was significantly faster in streams than in tree
holes. Mass loss of banyan leaves between tree holes and the stream was positively correlated with conidial output by the
aquatic hyphomycetes. The estimated half-life (t50 ) of leaf decomposition ranged between 70 days (stream) and 128 days
(tree holes). Organic carbon in leaf litter gradually decreased, while nitrogen content decreased up to 1–2 weeks and then
gradually increased over the incubation period. Ergosterol showed a gradual increase in tree hole litter, while it attained its
peak in 4 weeks and declined thereafter in the stream litter. Loss of phosphorus concentration was rapid in the stream than in
tree holes. Total phenolics concentration in leaves decreased rapidly in the stream than in tree holes. However, low dissolved
oxygen in water and the slow release of phenolics from the leaf litter showed negative impact on the fungal activity and the
rate of leaf decomposition in tree holes.
Keywords: aquatic hyphomycetes; ergosterol; leaf litter; decomposition; coastal stream; tree holes
1. Introduction
Filamentous fungi play an important role in the decomposition of organic matter and energy flow in food webs in
terrestrial and aquatic ecosystems. Aquatic hyphomycetes,
a phylogenetically heterogeneous group, dominant on dead
leaf litter and constitute an important trophic link between
leaves and the stream invertebrates (Bärlocher 1992a,
1992b). In comparison with mycological studies in aquatic
habitats, the canopies of riparian trees attracted less attention. Tree canopies represent a structurally complex and
ecologically important subsystem support the evolution of
flora, fauna and micro-organisms, which are rare or not
commonly encountered on the forest floor (Nadkarni et al.
2001; Sridhar 2009). Fungi involving in breakdown and
improvement of nutritional quality of accumulated leaf litter for the inhabiting fauna (e.g. invertebrates and tadpoles)
in tree holes (vertical or lateral cavities formed in the trunk
or branches of the trees either naturally due to stress or
damage by the attack of microbes or fauna) assumes importance in trophic structure and energy flow in tree canopies.
Additionally, tree holes might serve as source of inoculum
*Corresponding author. Email: kandikere@gmail.com
© 2013 Mycological Society of China
Published online 12 Aug 2013
of aquatic fungi that colonize organic matter in streams.
A few studies have been carried out to evaluate the waterborne fungi in tree canopies, which are mainly confined to
temperate regions (Magyar et al. 2005; Gönczöl & Révay
2006; Sridhar 2009).
Western Ghats and south-west coast of India receives
substantial rains during south-west monsoon (June–
September) and the tree canopies are wet throughout monsoon and part of post-monsoon season (October–January).
Thus, occurrence of aquatic or water-borne fungi in tree
canopies (e.g. leaf surface, through fall and stem flow)
is not surprising (Sridhar 2009). The question remains to
understand the pattern of fungal assemblage and leaf litter decomposition in tree canopies in comparison to the
stream ecosystem. There are a few detailed studies on leaf
litter decomposition in the Indian streams (Raviraja et al.
1996, 1998; Sridhar et al. 2011; Sudheep & Sridhar 2013).
The primary objective of this study was to investigate the
structure and function of aquatic hyphomycete assemblages
in riparian tree holes and to compare it with that of a
second-order stream in the south-west coast of India during
monsoon season. The secondary objective was to assess the
Mycology
relationship between descriptions of fungal colonization,
decomposition and chemical changes of banyan leaves.
2. Materials and methods
2.1. Leaf litter
As banyan leaves showed higher richness of aquatic
hyphomycetes in Konje stream (Sridhar et al. 1992), senescent and freshly fallen leaves of banyan (Ficus benghalensis L.) were collected from a single tree in Mangalore
University Campus. The lamina portions of the leaves were
punched into 1.5 cm discs using a cork borer. They were
air-dried up to 3–4 weeks (30◦ C), 75 leaf discs were placed
in each big nylon litter bags (15 × 15 cm, mesh size 1 mm),
and 25 pre-weighed leaf discs were placed in small nylon
litter bags (5 × 5 cm, mesh size 1 mm) and placed in
each big litter bag for mass-loss determination. The mass
of 25 leaf discs enclosed in mesh bags ranged between
590 and 670 mg.
2.2. Field experiments
The field experiment was conducted during monsoon season (July–August 2006). The second-order stream, Konaje
(12◦ 48′ N, 74◦ 55′ E) flowing through forest and plantation crops adjacent to the Mangalore University Campus
was selected. On 6 July 2006, the litter bags were introduced into tree holes and to the stream. The litter bags were
tied to tree roots along the margin in five different locations
at a distance of approximately 50–100 m, where there was
water flow throughout the experimental period. As banyan
trees were not close to the stream, five riparian trees were
chosen (three of Holigarna ferruginea Marchand and two
of Syzygium caryophyllatum (Linn.) Alston at a distance
of approximately 50–100 m from each other. Litter bags
were tied to the branches or the trunk and incubated in a
single hole in each tree. During the experimental period,
litter bags were fully or partially submerged in tree holes.
One litter bag from each tree hole and stream location was
sampled in five occasions (1, 2, 4, 6 and 8 weeks) and
brought to the laboratory for fungal analysis, leaf mass-loss
determination and leaf chemical analysis.
Water temperature, pH and conductivity of each tree
hole and stream location were measured at the time of sampling between 9 am and 12 noon (Water Analyzer 371,
Systronics, Gujarat, India). Water samples collected were
fixed during 9 am – 12 noon and the dissolved oxygen was
estimated by Winkler’s method (APHA 1995).
2.3.
Examination for fungi
On each sampling, five leaf discs per replicate were incubated in 150 ml distilled water in 250 ml Erlenmeyer flasks,
and aerated with Pasteur pipettes connected to aquarium
pumps, for 48 h (23 ± 2◦ C) to induce the production of
119
conidia by aquatic hyphomycetes. The conidial suspension was filtered through Millipore filter (5 µm; diameter 47 mm) and the filter was stained with 0.1% cotton
blue in lactophenol. The filter was cut into half, mounted
on a microscope slide with lactic acid and the conidia
were identified (magnification, 200–1000 ×) and counted
(1/8–1/4 of the filter area was scanned if the conidial density in the filter was high, the whole filter was scanned if
conidia were sparse). The leaf discs were dried at 100◦ C up
to 24 h and the dry mass was determined on attaining constant weight. The conidial output was expressed per gram
dry mass of leaf discs. Fungal conidia were identified using
monographs and taxonomic keys (see Sudheep & Sridhar
2011).
2.4. Ergosterol estimation
To estimate ergosterol in leaf discs, modified microwaveassisted extraction was followed (Young 1995).
Lyophilized leaf discs (50–100 mg) were churned in
liquid nitrogen, suspended in a solution of 2 ml methanol
and 0.5 ml 2 M NaOH and sealed in 15 ml screw-cap
glass tubes. Six extraction tubes with samples were placed
in a 250 ml capped plastic bottle and microwaved (50%
power for 95 s in a microwave oven) and then removed
from the plastic bottle after cooling to room temperature.
The solution was neutralized with 1 ml of 1 M HCl, and
ergosterol concentration was extracted with three consecutive hexane washes. The combined hexane fractions were
evaporated to dryness, and the residue was re-extracted
in 1 ml of methanol. The solution was injected into a
HPLC-C18 column (Sigma-Aldrich, St. Louis, MO, USA)
and eluted with methanol at 1.5 ml min−1 at 20◦ C (elution
time, 5.3 min). The ergosterol content was estimated by
comparison of peak areas with those of external standards.
2.5.
Leaf mass loss
The leaf discs in small litter bags placed within big litter bags were sampled for mass-loss assessment. This was
determined by comparing original and the remaining leaf
mass after air-drying and corrected by exposing subsamples to 100◦ C for 24 h in five replicates. Exponential decay
coefficient per day (k/d) was estimated by linear regression
on ln-transformed data (MATLAB 6.5).
2.6.
Leaf chemical analysis
Dried leaf discs were powdered and the organic carbon was
measured as outlined by Kalembasa and Jenkinson (1973).
Nitrogen estimation of dried leaf powder was carried out
according to Chale (1993). Ascorbic acid method was followed to determine the total phosphorus in leaf samples
(APHA 1995). Total phenolics of the leaf disc powder was
determined by Rosset et al. (1982).
120
K.R. Sridhar et al.
2.7. Data analysis
Water parameters, ergosterol content and leaf chemistry
between tree holes and the stream were compared by t-test
(StatSoft Inc. 2008). Sporulation by aquatic hyphomycetes
(log transformed) against incubation period were compared
among habitats using two-way ANOVA (habitats and time
as categorical variables) followed by Tukey’s test (Zar
1996). Cumulative conidial output among the habitats was
compared by ANCOVA followed by Tukey’s test.
Exponential decay coefficients per day (k/d) were estimated for leaf litter in tree holes and in the Konaje stream,
based on the negative exponential decay model (Petersen
& Cummins 1974) using MATLAB 6.5. Comparison
of slopes of habitats (tree holes and stream) was carried out by ANCOVA (habitats and time as categorical and continuous variables, respectively) followed by
Tukey’s test. Relationship between the rate of decomposition and conidial output in tree holes and the stream
was assessed by linear regression (MATLAB 6.5). The
time (days) required for the decomposition of half of the
initial leaf mass (t50 ) was determined by the following
relation:
t50 = ln(2)/k
Table 1 gives water quality in tree holes and Konaje stream
during leaf immersion and sampling period. The mean temperature in tree holes during study period ranged from
23.2 to 26.3◦ C, pH 6.5 to 6.9, conductivity 37.6 to 51.2
µS cm−1 and dissolved oxygen 3.2 to 3.9 mg l−1 , while
in the stream 23.2 to 26.3◦ C, 6.7 to 8.1, 117 to 1330 µS
cm−1 and 1.1 to 1.7 mg l−1 , respectively. Water temperature
(t-test, p < 0.001), pH (t-test, p < 0.01), conductivity (t-test, p < 0.001) and dissolved oxygen (t-test,
p < 0.01) were significantly differed between tree holes and
the stream.
Table 1. Water parameters in tree holes and Konaje stream during sampling period (n = 18, mean ± SD) (range in parenthesis).
pH
Conductivity
(µS cm−1 )
Dissolved oxygen
(mg l−1 )
Fungal colonization and ergosterol
Table 2 reveals colonization of aquatic hyphomycetes on
leaf litter submerged in tree holes and the Konaje stream
during 8 weeks of study. Altogether, 18 fungi colonized the
leaf litter in tree holes and the stream. Seventeen fungi were
found in tree holes as well as in the stream with exception of Helicomyces roseus, which was confined to tree
holes. Anguillospora longissima, Flagellospora curvula,
Lunulospora curvula, Triscelophorus acuminatus and
T. konajensis were the top five species in tree holes and
the stream. Maximum species richness was attained after
2 weeks in tree holes (11 species) and 6 weeks in the
stream (Figure 1(a)). Except for eighth week, conidial output was higher in the stream than in tree holes (Figure 1(b)).
The mean conidial output was the highest in the stream
compared to the tree holes (2437 vs. 1848 g−1 leaf litter)
(Table 2). Considering all sampling dates, the conidial output was one-third lower in tree holes than in the stream
(two-way ANOVA, p < 0.001) (Table 2). The cumulative
conidial output was higher in the stream than in tree holes
(ANCOVA, p = 0.004).
The ergosterol concentration in leaf discs was
1.4 µg g−1 before immersion. In tree holes, ergosterol
increased gradually over the study period, while it attained
a peak at 4 weeks (33.9 µg g−1 ) in the stream (Figure 1(c)).
Table 2. Water-borne fungi (conidia g−1 dry mass) on banyan
leaf litter exposed to tree holes and Konaje stream during sampling period.
3. Results
3.1. Water quality
Temperature (◦ C)
3.2.
Tree holes
Stream
p-value
25.1 ± 1.10
(23.2–26.3)
7.3 ± 0.33
(6.7–8.1)
624.1 ± 85.9
(117–1330)
2.5 ± 1.78
(1.1–1.7)
24.5 ± 1.03
(23.2–26.3)
6.7 ± 0.28
(6.5–6.9)
43.4 ± 7.0
(37.6–51.2)
3.6 ± 0.29
(3.2–5.7)
<0.001
<0.01
<0.001
<0.01
Mean conidia g−1 dry
mass
Taxon
Lunulospora curvula Ingold
Triscelophorus acuminatus Nawawi
T. konajensis K.R. Sridhar & Kaver.
Anguillospora longissima (Sacc.
& P. Syd.) Ingold
Flagellospora curvula Ingold
Arborispora palma K. Ando
T. monosporus Ingold
Clavariopsis aquatica de Wild.
Flabellospora verticillata Alas.
Campylospora chaetocladia Ranzoni
Tricladium splendens Ingold
Alatospora acumianta Ingold
Tricladium fuscum Nawawi
Phalangispora constricta Nawawi &
J. Webster
F. penicillioides Ingold
Culicidospora aquatica R.H.
Petersen
Helicomyces roseus Link
Isthmotricladia gombakiensis
Nawawi
Total species
Total conidia g−1 dry mass
Tree holes
Stream
267
1143
131
147
1632
530
102
36
67
33
6
2
22
2
8
5
5
4
34
9
23
22
2
17
11
11
5
3
2
2
4
1
2
1
−
<1
18
1849
17
2442
Mycology
121
indicating significantly higher mass loss in the stream
(ANCOVA, p = 0.020). This translated into a significantly
lower decomposition rate in tree holes (k = 0.0024 d−1 )
than in the stream (k = 0.0043 d−1 ) (Table 3). Mass loss
of banyan leaves between the habitats was positively correlated with the conidial output by aquatic hyphomycetes
(linear regression, p = 0.003, r2 = 0.25).
The estimated time for 50% litter mass loss was higher
in tree holes (128 days) than in the stream (70 days)
(Table 3). Comparison of decay dynamics between environments was performed by considering all replicates and
sampling dates. The daily decay coefficient (k) varied
between 0.0024 (tree hole) and 0.0043 (stream). Estimated
time for 50% mass loss (t50 ), was ranged between 70
(stream) and 128 (tree holes) days.
3.4.
Leaf chemical changes
Organic carbon concentration in leaf discs decreased gradually in the stream and tree holes, however, stream leaf
discs lost organic carbon faster compared to leaf discs
in tree holes (Figure 2(b)). Nitrogen concentration in leaf
discs decreased initially (1–2 weeks) and then increased
gradually until the end of the experiment in both habitats
(Figure 2(c)). The C:N ratio of leaf discs peaked after 1
(stream)–2 weeks (tree holes) and then decreased until the
end of the incubation period (Figure 2(d)). Phosphorus concentration decreased faster in the stream than in tree hole
leaf discs (Figure 2(e)). A steep decrease in total phenolics concentrations in leaf discs was found in the stream
than in tree holes; however, after 8 weeks the phenolics
concentration was similar in both habitats (Figure 2(f)).
Organic carbon, nitrogen, phosphorus and phenolics in leaf
discs were significantly differed between tree holes and the
stream (t-test, p < 0.05).
Figure 1. Fluctuation in (a) species richness, (b) conidial output and (c) ergosterol during the decomposition of banyan leaf
litter immersed in tree holes and the Konaje stream (n = 5,
mean ± SD).
The ergosterol content in leaf litter between tree holes and
the stream was significantly differed (t-test, p < 0.05).
3.3. Leaf mass loss
After 8 weeks immersion, litter had lost 11% initial mass in
tree holes and 22% initial mass in the stream (Figure 2(a))
4. Discussion
Aquatic hyphomycetes have evolved different strategies
to survive under unusual conditions (e.g. soil, tree roots
and tree canopies) (Sridhar & Kaveriappa 1987; Bärlocher
1992b; Sridhar & Bärlocher 1993; Gönczöl & Révay 2003,
2004). Their ability to survive outside the water (e.g.
leaf/wood litter in the stream margins) and tree canopies
(e.g. tree holes and epiphytes) make these environments
potential sources of aquatic hyphomycetes that can colonize the streams. In this study, we found 18 species of
aquatic hyphomycetes colonizing leaf litter in tree holes,
which is similar to the species richness associated with
decomposing litter in the Konaje stream (18–20 species;
this study; Sridhar & Kaveriappa 1984; Sridhar et al. 1992).
Based on daily decay coefficients (k), Petersen and
Cummins (1974) classified the decomposition rates of
leaf litter roughly into three groups: ‘slow’ (k < 0.005),
‘medium’ (k = 0.005–0.01) and ‘fast’ (k > 0.01).
122
K.R. Sridhar et al.
Figure 2. Leaf mass remaining (a), organic carbon (b), nitrogen (c), C:N ratio (d), phosphorus (e) and total phenolics (f) of banyan leaf
litter during decomposition in tree holes and the Konaje stream (n = 5, mean ± SD).
Table 3. Summary of regression analysis of mass losses of
banyan leaf discs in tree holes and stream (k = decay coefficient;
r2 = coefficient of determination; t50 = half-life).
Habitat
Tree holes
Stream
k (per day)
r2
t50
0.0024
0.0043
0.99
0.99
127.88
69.91
The previous studies on decay of acacia, banyan,
cashew and eucalypt leaves in Konaje streams revealed
medium decay coefficient compared to the present study
(0.0063–0.0090 vs. 0.0043 d−1 , respectively) possibly due
to environmental differences between the stream reaches
considered in this study and in the previous ones (Raviraja
et al. 1996). The decay coefficient was also higher in midaltitude of the Western Ghat stream (Sampaje) than the
present study (0.0063–0.009 vs. 0.0043), so also the polluted stretches of River Nethravathi (0.0066–0.0075 vs.
0.0043) (Raviraja et al. 1996, 1998). No studies are
available to compare the decay coefficient of leaf litter in
tree holes. The current study revealed that the decay rate
is 1–3 folds slower in tree holes compared to the Konaje
or Sampaje streams and the River Nethravathi (0.0024 vs.
0.0043–0.009).
The nitrogen concentration on decomposing leaf litter decreased initially, which can be attributed to initial leaching (Davis et al. 2003). Subsequently, nitrogen concentration increased most likely as a result from
microbial colonization and the formation of complexes
between phenolics/lignin and proteins or other nitrogenous
compounds (Graça & Bärlocher 1998; Canhoto & Graça
1999; Bärlocher 2005). The slower and more prolonged
decrease in nitrogen concentration of leaf litter in tree holes
than in the stream suggests a less intense leaching as well
as a delayed fungal colonization of leaf litter in tree holes.
Initial increase and subsequent decrease in C:N ratio of leaf
litter in the streams and tree holes in the present study also
reflected the elevation of nitrogen concentration and loss of
carbon.
Changes in phosphorus in decomposing leaf litter are
dependent on the type of leaf litter and nature of the stream.
Mycology
Phosphorus seems to reflect a balance between leaching
and immobilization by fungal activity. Alder, poplar and
willow leaf litter in temperate regions showed an initial
increase followed by a gradual decrease (Chauvet 1987).
Eucalypt in Spanish streams lost phosphorus in headwaters,
while accumulated in lower reaches (Pozo 1993). In fresh
and dried leaves of Rhizophora mucronata in a south-west
mangrove, an initial loss of phosphorus was seen up to
1 week followed by its elevation (Ananda et al. 2008). In the
present study, loss of phosphorus was rapid in the stream
than in tree holes and coincided with more conidial output.
Similar observation was made in the streams of Konaje and
Sampaje, where banyan and cashew leaves lost phosphorus more rapidly (than acacia and eucalypt) and coincided
with elevated conidial output (Raviraja et al. 1996). Sharp
decline of phosphorus in banyan leaf in River Nethravathi
also showed more conidial output of aquatic hyphomycetes
(Raviraja et al. 1998).
Greater conidial output of aquatic hyphomycetes was
associated with faster leaf colonization and rates of leaf
decomposition (Gulis & Subekropp 2003). Most likely, the
very high or too low nutrients (especially nitrogen and
phosphorus contents) in the Konaje stream during the study
period might be responsible for low rates of leaf decomposition. On the contrary, although tree holes possess more
nitrogen and phosphorus concentration, they reflect low
dissolved oxygen (0–4 weeks, 1.1–1.7 mg l−1 ) or increased
phenolic contents in water might have a negative effect on
the colonization and functions of aquatic hyphomycetes.
The low dissolved oxygen concentration (Medeiros et al.
2009) and the higher phenolics concentration in water
resulting from leaf leaching in a contained environment
(Canhoto & Graça 1999), might negatively affect fungal
colonization of and activity on litter in tree holes than in
the stream. Loss of phenolics in banyan litter in the present
study is comparable to banyan and cashew litter in the
Konaje and Sampaje streams, while it was slow in acacia
and eucalypt (Raviraja et al. 1996, 1998). Besides phenolics, perhaps higher bacterial activity depending on the
nutrient levels in tree holes might reflect in low dissolved
oxygen in our study.
Aquatic hyphomycete assemblages seem to have
adapted to harsh conditions, which might partially be due to
the presence of species-specific requirements (for instance,
L. curvula is known to be a warm water preferring species
and its dominance in tree holes and streams in the present
study is not surprising). Similar adaptation or tolerance
for low dissolved oxygen might exist in tree hole inhabitants. The species richness in tree holes is similar to that in
the stream and thus tree holes serve as a potential source
of inoculum to the streams. Nevertheless, an increased
fungal biomass and nitrogen concentration in leaf litter in
tree holes by aquatic hyphomycetes meet the nutritional
requirement of invertebrates inhabiting in tree holes.
123
5. Conclusions
Compared to temperate region, studies on leaf litter decomposition in aquatic habitats of tropical region are scarce.
The present study constitutes the first study on the leaf litter
decomposition in tree holes of tropical habitats. Future
studies need to evaluate the pattern and process of plant litter decomposition in different tropical aquatic habitats, so
also the canopies of riparian tree species. Such studies will
give insight into the basic process of energy flow and food
web in tropical ecosystem. Besides, these studies serve as
baseline data to forecast the status of the habitat (healthy or
impoverished or regenerated) to impart remedial measures
to control human interference on the ecosystem.
Acknowledgements
The authors are grateful to Mangalore University for granting permission to carry out this study at the Department of
Biosciences. One of us (KSK) acknowledges the research fellowship granted under SC/ST Cell, Mangalore University and Rajiv
Gandhi Fellowship, University Grants Commission, New Delhi,
India. Authors are thankful to Prof. F. Bärlocher for helping in the
estimation of ergosterol and Madhu S. Kandikere for his statistical analysis. We are indebted to the referees for critical comments
to improve this paper.
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