RESEARCH ARTICLE
Diversity of filamentous fungi on decomposing
leaf and woody litter of mangrove forests in the
southwest coast of India
K. Ananda and K. R. Sridhar*
Department of Biosciences, Mangalore University, Mangalagangotri, Mangalore 574 199, India
Assemblage and diversity of filamentous fungi on leaf
and woody litter accumulated on the floor of two man-
grove forests (Nethravathi and Udyavara) in the south-
west coast of India have been studied. Leaf and woody
litter collected during summer and monsoon seasons
incubated in the laboratory (2, 8, 16 and 32 weeks)
yielded 78 taxa belonging to 32 ascomycetes and 46 mito-
sporic fungi. Significant difference (ANOVA) was seen
in the frequency of occurrence of fungal taxa in diffe-
rent incubation periods of leaf litter (P = 0.179 × 10–5 to
0.24 × 10–5) and woody litter (P = 0.887 × 10–8 to 0.0326).
The number of fungal taxa per leaf litter ranged from
1 to 5, while for woody litter it was 1 to 8. Mean number
of fungal taxa per wood ranged between 2.5 and 3.6,
which is higher than that in the mangroves of the Indian
Ocean and South East Asia. Seven ascomycetes (Anip-
todera chesapeakensis, Aniptodera sp., Arenariomyces
parvulus, Lulworthia grandispora, Savoryella lignicola,
MANGROVES are highly productive ecosystems next to the
coral reefs and provide energy to marine habitats through
production and decomposition of plant detritus1,2. Litter
biomass in mangrove forests3,4 ranges between 0.011 and
23.69 t h–1 y–1. Studies have been carried out on fungal
colonization and the pattern of decomposition of mangrove
leaf litter5–10 and woody litter10–14. However, a few quan-
titative studies are available on fungal association with
mangrove litter in the Indian coast (west coast15–18; east
coast19,20). Raghukumar et al.16 studied the sequence of col-
onization, densities and biomass of fungi, thraustochytrids
and bacteria on leaf litter of Rhizophora apiculata at Caorao
mangrove, Goa. Several studies on mangroves in India re-
ported either marine fungi19,20 or terrestrial fungi21,22. It is
evident that both marine and terrestrial fungi are involved
in mangrove litter decomposition23. Terrestrial fungi in man-
grove habitats are ignored as transient mycota24,25. There-
fore, the current study aims at understanding (i) the richness
and diversity of filamentous fungi on randomly collected
leaf and woody litter of two mangrove forests in the south-
west coast of India during summer and monsoon seasons;
(ii) the succession of fungal sporulation on mangrove litter
at different intervals of incubation.
*For correspondence. (e-mail: sirikr@yahoo.com)
CURRENT SCIENCE, VOL. 87, NO. 10, 25 NOVEMBER 2004
Kallichroma tethys and Verruculina enalia) and seven
anamorphic taxa (Arthrinium sp., Aspergillus sp. 1, Cir-
renalia pygmea, Cirrenalia tropicalis, Penicillium sp.,
Periconia prolifica and Trichocladium alopallonellum)
belonging to core-group fungi (frequency of occurrence,
10%) were identified. Woody litter collected during
summer season showed highest fungal diversity than
during monsoon. Rarefaction estimates of expected
number of fungal taxa out of 150 isolates were higher
in woody litter than leaf litter. Sporulation of terrestrial
fungi, marine fungi and arenicolous fungi (sand-inhabit-
ing) was successional on leaf and woody litter. Terres-
trial fungi sporulated within eight weeks and declined
thereafter, sporulating marine fungi were highest dur-
ing 16 weeks, while arenicolous fungi attained a peak
after 32 weeks of incubation. Substrate preference, suc-
cession, competition and techniques of study of mangrove
filamentous fungi have been discussed.
Materials and methods
Study area and sampling
Nethravathi and Udyavara mangroves in the southwest
coast of India were chosen for the study. Nethravathi is situa-
ted south of Mangalore (12°50′00″N, 74°50′00″E) and its
major part has been denuded due to human interference
(tile and wood factories, railways, agriculture and aqua-
culture). Recently, part of this mangrove area has been reve-
getated. Common mangrove plant species include Rhizophora
mucronata and Avicennia officinalis. Udyavara is located
north of Mangalore (13°17′50″N, 74°44′32″E) with large
mangrove vegetation. Acanthus ilicifolius, A. officinalis
and R. mucronata are the major mangrove species. Dry
season (summer) starts during February, while wet season
(monsoon) during June. Dead leaf and woody litter accu-
mulated on the mangrove floor (about 0.25 km2 area) were
randomly sampled three months after initiation of summer
(May 2000) and three months after initiation of monsoon
(September 2000). They were pooled in sterile polythene
bags and brought to the laboratory for fungal assessment.
Incubation, observation and data analyses
Each leaf and wood was incubated (25 ± 2°C) separately
in polythene bags containing sterile sand wetted with sterile
1431
RESEARCH ARTICLE
50% sea water. They were screened under a stereomicro-
scope for the fungal structures at different intervals of incu-
bation (2, 8, 16 and 32 weeks) and the sporulated fungi were
identified.
Per cent frequency of occurrence of each fungus and
mean number of fungi per litter sample were calculated:
Frequency of occurrence (%) =
Total number of leaf or wood colonized by a specific fungus
Total number of leaf or wood supporting sporulating fungi
× 100.
Mean number of fungi per sample =
Total number of fungal isolations
. were dominant on woody litter. Based on the frequency of
Total number of leaf or wood sporulating fungi
One-way ANOVA was employed to determine the differ-
ences in the frequency of occurrence of fungal taxa based
on sporulation at different incubation periods26.
Diversity of fungi on litter during monsoon and summer
seasons was assessed based on the diversity indices27:
Simpson diversity, D′ = 1/Σ(pi)2 and Shannon diversity,
H′ = –Σ(pilnpi), where pi is the proportion of individuals
that taxa i contributes to the total.
The evenness was expressed by: J′ = H′/H′max, where
H′max is the maximum value of diversity for the number of
taxa present28.
To compare species richness among the isolations on
substrata during two seasons, rarefaction indices were
calculated29. The expected number of fungal taxa, E(t), in
a random sample of n isolations taken from a total popu-
lation of N isolations was calculated using the formula:
s   N − ni   N   
E(t) = ∑ 1 −      ,
i =1   n   n   
where ni is the number of isolations of the ith taxa.
Results
Altogether 78 fungal taxa were recovered from leaf and
woody litter. Out of 473 isolations from leaf litter, 45 taxa
and among 599 isolations from woody litter, 65 taxa were
obtained (Tables 1 and 2). Samples collected during summer
showed maximum fungi (68 taxa) than those collected
during monsoon (55 taxa). Per cent fungal taxa and cumu-
lative fungal taxa sporulated on litter at different periods
of incubation are given in Figure 1. Sporulating fungi on
leaf litter attained a peak during 8 or 16 weeks and decli-
ned thereafter, while on woody litter such drastic decline
was not seen. Cumulative taxa between 16 and 32 weeks
showed slight increase. The frequency of occurrence was
highest on leaves at eight weeks incubation, while on
wood, it was at 16 weeks incubation. One-way ANOVA
1432
revealed significant difference in the frequency of occurrence
of fungal taxa during different incubation periods of leaf litter
(Nethravathi, P = 0.24 × 10–5; Udyavara P = 0.179 × 10–5) as
well as woody litter (Nethravathi, P = 0.887 × 10–8; Udya-
vara, P = 0.0326).
Among 13 core-group fungi (>10%) on mangrove litter,
Arenariomyces parvulus and Cirrenalia pygmea were
dominant on leaf as well as woody litter (Table 1). Arthri-
nium sp. was restricted to leaf litter, while Kallichroma
tethys and Verruculina enalia were confined to woody litter.
Aspergillus sp. 1, Cirrenalia tropicalis, Lulworthia gran-
dispora, Penicillium sp. and Trichocladium alopallonellum
were dominant on leaf litter, while Aniptodera chesa-
peakensis, Periconia prolifica and Savoryella lignicola
occurrence in summer and monsoon, 12 fungi were consi-
dered as core-group fungi. Aniptodera chesapeakensis, Are-
nariomyces parvulus, Arthrinium sp., C. pygmea, L. grandi-
spora and T. alopallonellum were dominant during summer
as well as monsoon seasons. Aniptodera sp. was dominant in
summer, while Aspergillus sp. 1, C. tropicalis, Penicillium
sp., P. prolifica and V. enalia were dominant during monsoon.
The mean number of fungal taxa per sample of leaf and
wood was 2.4 (range 2.1–2.6) and 3 (range: 2.5–3.6) res-
pectively (Table 2). The number of taxa per leaf litter
ranged from 1 to 5, while for woody litter it was 1 to 8
(Figure 2). Maximum leaf and woody litter (33.8 and
26%) was colonized by three taxa. Leaf litter of Nethra-
vathi mangrove showed the highest fungal diversity during
summer, while for Udyavara it was during monsoon. On
woody litter of both locations, summer samples showed
the highest diversity. The rarefaction estimates of expected
number of species out of 150 isolations were consistently
higher on woody litter irrespective of substrata and season
(Table 2, Figure 3).
Fungal taxa on leaves and woody litter comprised ter-
restrial, marine and arenicolous (sand-inhabiting) fungi.
The pattern of sporulation of these fungi at different dura-
tions of incubation is given in Figure 4. More mitosporic
fungi sporulated at initial phases of incubation followed by
ascomycetes and arenicolous fungi. By and large, maxi-
mum terrestrial fungi sporulated during eight weeks and
declined thereafter. Sporulating marine fungi peaked dur-
ing 16 weeks and declined subsequently, except on woody
litter of Udyavara. Sporulating arenicolous fungi reached
a peak at 32 weeks of incubation.
Discussion
The occurrence of core-group fungi in our study corrobo-
rates with earlier investigations17,18. It is predicted that
among fungal species in a community, core-group fungi
exert major influence on turnover of litter in mangrove
ecosystem. Perhaps fungi colonized on leaf and woody
litter may have different vegetative and reproductive stra-
CURRENT SCIENCE, VOL. 87, NO. 10, 25 NOVEMBER 2004
 RESEARCH ARTICLE
Table 1. Frequency of occurrence (%) of filamentous fungi on leaf and woody litter of Nethravathi and Udyavara mangroves in the west coast of India

Leaf litter Woody litter

Mean % Mean %
Nethravathi Udyavara Nethravathi Udyavara frequency frequency
Taxon S, M S, M S, M S, M L, W S, M

Ascomycetes
Arenariomyces parvulus Koch 11.1, 15.8 36.4, 10.5 15.1, 7.8 5.7, 15.9 18.5, 11.1 17.1, 12.5
Lulworthia grandispora Meyers 35.6, 18.4 0, 15.8 9.1, 7.8 3.8, 4.6 17.5, 6.3 12.1, 11.7
Aniptodera chesapeakensis Shearer et Miller 11.1, 7.9 6.1, 10.5 18.2, 20.3 5.7, 11.4 8.9, 13.9 10.3, 12.5
Verruculina enalia (Kohlm.) Kohlm. et Volkm.-Kohlm. 0, 0 0, 0 19.7, 28.1 3.8, 22.7 0, 18.6 5.9, 12.7
Sovoryella lignicola Eaton et Jones 0, 0 6.1, 2.6 15.1, 23.4 7.6, 6.8 2.2, 13.2 7.2, 8.2
Aniptodera sp. 6.7, 0 15.1, 0 10.6, 0 9.4, 18.2 5.5, 9.6 10.5, 4.6
Kallichroma tethys (Kohlm. et Kohlm.) 0, 0 0, 0 10.6, 7.8 5.7, 22.7 0, 11.7 4.1, 7.6
Kohlm. et Volkm.-Kohlm.
Savoryella paucispora (Cribb et Cribb) Koch 0, 0 7.9, 0 4.6, 7.8 9.4, 11.4 2, 8.3 5.5, 4.8
Corollospora maritima Werdermann 2.2, 0 3, 0 10.6, 4.7 1.9, 11.4 1.3, 7.2 4.4, 4
Leptosphaeria australiensis (Cribb et Cribb) Hughes 11.1, 0 3, 0 4.6, 0 9.4, 0 3.3, 3.5 7, 0
Corollospora pulchella Kohlm., Schmidt et Nair 0, 0 3, 0 6.1, 0 3.8, 13.6 0.8, 5.9 1.3.4
Corollospora angusta Nakagiri et Tokura 0, 10.5 0, 0 6.1, 4.7 0, 0 2.6, 2.7 1.5, 3.8
Ascomycete sp. 1 4.4, 0 9.1, 0 0, 0 7.6, 0 3.4, 1.9 5.3, 0
Buergenerula sp. 4.4, 10.5 0, 0 0, 0 0, 0 3.7, 0 1.1, 2.6
Lignincola laevis Höhnk 0, 0 0, 0 0, 3.1 9.4, 0 0, 3.1 2.4, 0.8
Lulworthia sp. 0, 0 0, 0 3, 0 9.4, 0 0, 3.1 3.1, 0
Torpedospora radiata Meyers 0, 0 0, 0 4.6, 0 0, 6.8 0, 2.9 1.2, 1.7
Mycosporella salicorniae (Auerswald) Petrak 0, 0 0, 7.9 0, 0 1.9, 0 2, 0.5 0.5, 2
Ascosalsum cincinnatula (Shearer et Crane) Campb., 0, 0 0, 0 0, 0 1.9, 6.8 0, 2.2 0.5, 1.7
Anderson et Shearer
Dactylospora haliotrepha (Kohlm. et Kohlm) Hafellner 0, 0 3, 0 1.5, 0 3.8, 0 0.8, 1.3 2.1, 0
Didymosphaera lignomaris Strongman et Miller 0, 0 0, 0 0, 0 7.6, 0 0, 1.9 1.9, 0
Etheirophora unijubata Kohlm. et Volkm.-Kohlm. 0, 0 0, 0 0, 0 3.8, 2.3 0, 1.5 1, 0.6
Mycosphaerella apophlaeae Kohlm. 0, 0 6.1, 0 0, 0 0, 0 1.5, 0 1.5, 0
Panorbis viscosa (Schmidt) Campb., Anderson et Shearer 0, 0 0, 0 0, 0 5.7, 0 0, 1.4 1.4, 0
Arenariomyces trifurcatus Höhnk 0, 0 0, 0 1.5, 0 3.8, 0 0, 1.3 1.3, 0
Biflua physasca Koch et Jones 0, 0 0, 5.3 0, 0 0, 0 1.3, 0 0, 1.3
Mycosporella staticicola (Patouillard) Dias 0, 0 3, 0 0, 1.5 0, 0 0.8, 0.4 0.8, 0.4
Lindra marinera Meyers 4.4, 0 0, 0 0, 0 0, 0 1.1, 0 1.1, 0
Corollospora sp. 0, 0 0, 0 3, 0 0, 0 0, 0.8 0.8, 0
Corollospora cinnamomea Koch 0, 0 0, 2.6 0, 0 0, 0 0.7, 0 0, 0.7
Littospora ratnagiriensis (Patil et Borse) Campb., 0, 0 0, 0 0, 0 1.9, 0 0, 0.5 0, 5, 0
Anderson et Shearer
Natiantispora retorquens (Shearer et Crane) Campb., 0, 0 0, 0 0, 0 1.9, 0 0, 0.5 0.5, 0
Anderson et Shearer
Halographis runica Kohlm. et Volkm.-Kohlm. 0, 0 0, 0 0, 1.6 0, 0 0, 0.4 0, 0.4
Halosarpheia sp. 0, 0 0, 0 1.5, 0 0, 0 0, 0.4 0.4, 0
Leptosphaeria oraemaris Linder 0, 0 0, 0 0, 1.6 0, 0 0, 0.4 0, 0.4
Ascomycete sp. 2 0, 0 0, 0 1.5, 0 0, 0 0, 0.4 0.4, 0
Mitosporic fungi
Cirrenalia pygmea Kohlm. 13.3, 18.4 6.1, 10.5 27.3, 21.9 20.8, 13.6 12.1, 20.9 16.9, 16.1
Arthrinium sp. 37.8, 36.8 24.2, 29 0, 0 0, 0 32, 0 15.5, 16.4
Aspergillus sp. 1 20, 44.7 0, 18.4 1.5, 0 5.7, 15.9 20.8, 5.8 6.8, 19.8
Trichocladium alopallonellum (Kohlm. et Kohlm.) 11.1, 10.5 30.3, 15.8 4.6, 12.5 7.6, 13.6 16.9, 9.6 13.4, 13.1
Kohlm. et Volkm.-Kohlm.
Periconia prolifica Anastasiou 8.9, 0 3, 18.4 9.1, 17.2 3.8, 11.4 7.6, 10.1 6.2, 11.8
Cirrenalia tropicalis Kohlm. 2.2, 0 6.1, 7.9 13.6, 34.4 0, 0 4.1, 12 5.5, 10.6
Penicillium sp. 0, 29 0, 13.2 4.6, 1.6 0, 13.6 10.6, 5 1.2, 14.4
Zalerion varium Anastasiou 0, 0 0, 15.8 12.1, 1.6 5.7, 18.2 4, 9.4 4.5, 8.9
Alternaria sp. 1 2.2, 13.2 6.1, 2.6 4.6, 6.1 5.7, 0 6, 4.1 4.7, 5.5
Mitosporic fungus 15.6, 0 12.1, 5.3 0, 3.1 3.8, 0 8.3, 1.7 7.9, 2.1
Trichocladium sp. 1 2.2, 7.9 0, 0 4.6, 4.7 1.9, 18.2 2.5, 7.4 2.2, 7.7
Zalerion maritimum (Linder) Anastasiou 0, 0 6.1, 2.6 10.6, 7.8 9.4, 2.4 2.2, 7.6 6.5, 3.2
Aspergillus sp. 2 2.2, 18.4 0, 13.2 0, 0 0, 4.6 8.5, 1.2 0.6, 9.1
Trichocladium achrasporum (Meyers et Moore) Dixon 6.7, 10.5 6.1, 10.5 3, 0 0, 0 8.5, 0.8 3.4, 5.3
Alternaria sp. 2 8.9, 7.9 0, 2.6 6.1, 3.1 0, 0 4.9, 2.3 3.4, 3.4
(Contd…)

CURRENT SCIENCE, VOL. 87, NO. 10, 25 NOVEMBER 2004 1433

RESEARCH ARTICLE
Table 1. (Continued)

Leaf litter Woody litter

Mean % Mean %
Nethravathi Udyavara Nethravathi Udyavara frequency frequency
Taxon S, M S, M S, M S, M L, W S, M

Clavatospora bulbosa (Anast.) Nakagiri et Tubaki 0, 0 6.1, 5.3 6.1, 4.7 3.8, 0 2.9, 3.7 4, 2.5
Alternaria sp. 3 0, 0 0, 0 0, 0 17, 9.1 0, 6.5 4.3, 2.3
Zygosporium masonii 0, 7.9 0, 6.1 1.5, 3.1 5.7, 0 3.5, 2.6 1.8, 4.3
Fusarium oxysporum Schl. et Fries 4.4, 15.8 0, 0 0, 0 0, 0 5.1, 0 1.1, 4
Nigrospora oryzae (Berk. et Br.) Petch 17.8, 0 0, 0 0, 1.6 0, 0 4.5, 0.4 4.4, 0.4
Monodictys pelagica (Johnson) Jones 0, 2.6 0, 2.6 6.1, 3.1 3.8, 0 1.3, 3.3 2.5, 2.1
Pestalotiopsis sp. 4.4, 10.5 0, 0 0, 0 0, 0 3.7, 0 1.1, 2.6
Phoma sp. 1 0, 0 0, 0 3, 3.1 3.8, 4.6 0, 3.6 1.7, 1.9
Cladosporium sp. 3 11.1, 0 0, 0 0, 0 0, 0 2.8, 0 2.8, 0
Cladosporium oxysporum Berk. et Curt. 4.4, 0 0, 0 0, 3.1 0, 2.3 1.1, 1.4 1.1, 1.4
Periconia sp. 0, 0 0, 0 0, 0 9.4, 0 0, 2.4 2.4, 0
Fusarium sp. 1 0, 2.6 0, 0 1.5, 4.7 0, 0 0.7, 1.6 0.4, 1.8
Fusarium sp. 2 8.9, 0 0, 0 0, 0 0, 0 2.2, 0 2.2, 0
Cirrenalia basiminuta Raghukumar et Jainal 0, 0 6.1, 0 0, 0 0, 2.3 1.5, 0.6 1.5, 0.6
Cladosporium sp. 2 0, 7.9 0, 0 0, 0 0, 0 2, 0 0, 2
Sporidesmium sp. 0, 0 0, 0 0, 7.8 0, 0 0, 2 0, 1.2
Cladosporium sp. 1 0, 0 0, 0 1.5, 0 0, 4.6 0, 1.5 0.4, 1.2
Cytospora sp. 4.4, 0 0, 0 0, 1.6 0, 0 1.1, 0.4 1.1, 0.4
Alternaria chartarum Preuss 0, 0 0, 0 0, 0 0, 4.6 0, 1.2 0, 1.2
Camarosporium sp. 0, 0 0, 0 3, 0 0, 0 0, 0.8 0.8, 0
Curvularia lunata (Wakker) Boedijn 0, 0 3, 0 0, 0 0, 0 0.8, 0 0.8, 0
Phomopsis sp. 0, 0 3, 0 0, 0 0, 0 0.8, 0 0.8, 0
Coelomycete sp. 2 0, 0 0, 0 0, 0 1.9, 0 0, 0.5 0.5, 0
Dicyma pulvinata (Berk. et Curt.) v. Arx 0, 0 0, 0 0, 1.6 0, 0 0, 0.4 0, 0.4
Phoma sp. 2 0, 0 0, 0 1.5, 0 0, 0 0, 0.4 0.4, 0
Trichocladium sp. 2 0, 0 0, 0 0, 1.6 0, 0 0, 0.4 0, 0.4
Coelomycete sp. 1 0, 0 0, 0 1.5, 0 0, 0 0, 0.4 0.4, 0
Total ascomycetes 9, 5 12, 7 19, 13 23, 13 17, 31 32, 22
Total mitosporic fungi 19, 18 13, 17 21, 22 16, 15 28, 34 36, 33
Total taxa 28, 23 25, 25 40, 35 39, 28 45, 65 68, 55

S, Summer; M, Monsoon; L, Leaf litter; W, Woody litter; arranged in decreasing frequency of occurrence (%) in each group.

Figure 1. Per cent fungal taxa and cumulative fungal taxa sporulated on mangrove litter collected from
Nethravathi and Udyavara mangroves during summer and monsoon seasons on incubation in the labora-
tory for different durations.

1434 CURRENT SCIENCE, VOL. 87, NO. 10, 25 NOVEMBER 2004

 RESEARCH ARTICLE
Table 2. Fungal assemblage, diversity and species richness on mangrove leaf and woody litter of Nethravathi and Udyavara mangroves
during summer and monsoon seasons

Assemblage* Diversity Evenness Species

richness
Litter/mangrove Season SA FI FPS Simpson Shannon Simpson Shannon E(t150)**

Leaf litter
Nethravathi Summer 50 125 2.5 0.939 4.349 0.971 0.905 23
Nethravathi Monsoon 50 116 2.6 0.932 4.082 0.975 0.929 19
Udyavara Summer 50 71 2.1 0.925 4.134 0.959 0.890 23
Udyavara Monsoon 50 94 2.2 0.946 4.333 0.981 0.933 23
Woody litter

Nethravathi Summer 50 181 3.6 0.962 4.900 0.982 0.915 28

Nethravathi Monsoon 50 168 3.4 0.942 4.459 0.967 0.876 24
Udyavara Summer 50 124 2.5 0.967 5.020 0.989 0.950 31
Udyavara Monsoon 50 126 2.5 0.955 4.540 0.987 0.944 23

*SA, Number of samples assessed; FI, Number of fungal isolations; FPS, Number of fungi per sample.
**E(t150), Expected number of taxa out of 150 isolations.

Figure 2. Number of fungal taxa sporulated on leaf and woody litter.

tegies, as pointed out by Newell30. Due to the fast decom-

position of leaf litter in mangrove habitat, colonized fungi
might reproduce quickly in order to escape from the
competitors (e.g. fungi, protozoa, bacteria and detritus
feeders) and capture another substrate for colonization.
Due to persistent nature of the woody litter, colonized fungi
might prefer biomass accumulation strategy than repro-
ductive strategy. However, vegetative and reproductive
phases might overlap among the terrestrial, marine and
arenicolous fungi, which coexist on leaf and woody litter
in the current study. These fungi showed a clear-cut suc-
cession of sporulation. About 47% out of 78 fungi was
typical terrestrial fungi than marine fungi, of which most
belong to mitosporic fungi. Occurrence of terrestrial fungi
on mangrove litter in our study corroborates with earlier
observations18,20,23,31. Newell31 found terrestrial fungi dur-
Figure 3. Rarefaction curves on the occurrence of fungal taxa [num-
ber of isolates vs expected number of taxa, E(t)] on leaf and woody lit-
ter collected from Nethravathi and Udyavara mangroves during
different seasons.
ing first and second seral stages on baiting the seedlings of
Rhizophora mangle, but the third seral stage was domi-
nated by L. grandispora and Zalerion varium. Dominance
of terrestrial fungi on mangrove litter is dependent on the
duration and severity of rains during southwest monsoon
in the southwest coast of India. Usually heavy rains occur
during June–November, which decrease the salinity (0–
1.05‰) of mangrove waters, carry more sediments from

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RESEARCH ARTICLE
Figure 4. Per cent terrestrial, marine and arenicolous fungal taxa sporulated on mangrove litter col-
lected from Nethravathi and Udyavara mangroves on incubation in the laboratory for different durations.
upstream and result in colonization of substrata by terres-
trial fungi. Preference of some fungi to the kind of sub-
strata and season was seen in our study. Mitosporic fungi
were dominant on leaf and woody litter than ascomycetes;
so also during summer and monsoon seasons. Terrestrial
fungi, Arthrinium sp., Aspergillus sp. 1 and Penicillium
sp. were competitive against marine fungi on leaves and
they were dominant during monsoon than summer. Areni-
colous fungi usually reached a peak after 32 weeks of in-
cubation. Similarly, marine fungi, including arenicolous
fungi were seen on the intertidal wood collected from the
North Island (St. Mary’s Isles) in the southwest coast of
India32. A few taxa of arenicolous fungi were recorded on
the mangrove substrata11,33, possibly due to short-term in-
cubation in the laboratory.
Mangrove woody litter has been considered the second
important host for marine fungi after driftwood34. Greater
fungal diversity and species richness were seen on woody
litter than on leaf litter (65 vs 49 taxa), so also the fungal
taxa per sample (1–8 vs 1–5) in the present study. Such
diversity and richness of fungi might be due to the persis-
tence of woody litter than leaf litter in mangrove habitats.
However, mean fungal taxa per woody litter was 3 (range:
2.5–3.6). This is higher than the mangrove woody litter
studied at Seychelles, Brunei, Malaysia, Sumatra, Thailand,
Singapore, Hong Kong and Macau (1–2.7 per wood)35.
Tan et al.36 showed that V. enalia exhibited the highest
antagonism against associated fungi followed by Aigialus
parvus and Lignincola laevis. V. enalia became the most
‘aggressive’ taxon than A. parvus and L. laevis on coculturing
on mangrove wood36. Corollospora maritima and Lulwor-
thia sp. were also considered as ‘aggressive’ taxa by Miller
et al.37. But V. enalia in the present study coexisted with
many core-group fungi (A. chesapeakensis, A. parvulus,
C. pygmea, K. tethys, L. grandispora, P. prolifica and S.
1436
lignicola) on wood. It is likely that such coexistence prevails
due to positive rather than negative interaction on the
woody litter. It may be possible that such aggressive behaviour
of fungi is dependent on the kind of substrata. Halogra-
phis runica was recovered on the woody litter of Nethrava-
thi mangrove (1.6%) after long incubation (32 weeks). It has
been reported as an endolithic lichenoid from the Belize
and the Great Barrier Reef38,39. It was associated with
coral rock, shells of snails and worm tubes. Its phyloge-
nesis links to the terrestrial rather than marine ascomyce-
tes.
Mangrove fungi are known for their niche and substrate
specificity40. Based on the kind of mangrove substrata
(leaf, wood and bark), fungal preferences and succession
may vary. Assessment of fungal colonization on mangrove
litter will be more effective and informative, if different
methods are employed. Fungal colonization of mangrove
and intertidal woody litter was assessed by plating22,41 and
damp incubation42–44 techniques. Plating technique usually
results in the isolation of terrestrial fungi, while using the
damp incubation technique, besides terrestrial fungi, marine
fungi can also be recovered. Similarly, submerged aeration
technique facilitates isolation of freshwater hyphomycetes
from mangrove or estuarine substrata45,46. Future studies may
address substrate specificity, seasonal impacts, interference
competition and biotechnological potential of fungi in the
mangrove ecosystem.
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ACKNOWLEDGEMENTS. We are grateful to Mangalore University
for granting permission to carry out this study at the Department of
Biosciences and award of junior research fellowship to K.A. We thank
Dr N. S. Raviraja for part of the statistical assessment.
Received 23 April 2004; revised accepted 23 July 2004
1437