Biodiversity and Conservation 9: 393–402, 2000.
© 2000 Kluwer Academic Publishers. Printed in the Netherlands.
Biodiversity and distribution of fungi associated with
decomposing Nypa fruticans
KEVIN D. HYDE1 and SITI A. ALIAS2
1 Centre for Research in Fungal Diversity, Department of Ecology and Biodiversity, The University of
Hong Kong, Pokfulam Road, Hong Kong; 2 Institute of Biological Sciences, University of Malaya,
Kuala Lumpur, 50603, Malaysia
Received 30 April 1999; accepted in revised form 20 July 1999
Abstract. Fungi associated with the decomposition of Nypa fruticans in Malaysia are under investigation.
Forty-one fungi have been identified including 35 ascomycetes, four mitosporic fungi and two basidiomycetes. The distribution of intertidal fungi on palm structures including leaves, leaf veins, rachides, petiole
bases, and inflorescences, and fungi on terrestrial parts have also been examined. No fungi were found
on the leaf material, although several fungi were found on the leaf midribs, and possible reasons for this
are given. Very few taxa developed on the inflorescences, but those that were present were abundant. The
greatest density of fruiting structures occurred on the rachides, and the greatest diversity of fungal species
occurred on the petiole bases. The terrestrial fungi differed from the intertidal fungi, although Linocarpon
nipae occurred in both habitats. Reasons for the differences in fungal numbers and diversity on the various
palm parts are discussed. The diversity of fungi at Morib mangrove was low when compared to previous
studies on fungi on Nypa palm at Kampong Api Api in Brunei and in this study at Kuala Selangor mangrove
in Malaysia.
Key words: host specificity, marine fungi, palms, saprobes
Introduction
There have been relatively few studies on the fungi associated with Nypa palm (Nypa
fruticans), which is a common mangrove species in South East Asia where it can form
extensive stands (Tomlinson 1986). Tirisporella beccariana (as Sphaeria beccariana)
was the first fungus reported from Nypa fruticans (Cesati 1880) and, although not
reported as so, was almost certainly intertidal in origin. The second taxon reported
from Nypa palm was Astrosphaeriella nipicola (as Melanopsamma nipaecola), and
this is a terrestrial non-marine species (Hyde and Fröhlich 1998). Subsequently, as
few as four other taxa have been reported from aerial parts of this host (Hyde and
Alias 1999).
The first report of fungi from intertidal fronds of Nypa palm was that of Hyde
(1988) who reported Linocarpon appendiculatum, L. livistonae and L. nipae. Two
further intertidal species of Linocarpon were described in a monograph on the genus
by Hyde (1992c). Hyde and Nakagiri (1989) also reported Oxydothis nypae from
394
ageing or cut fronds of Nypa palm in the intertidal region. Oxydothis has since been
monographed (Hyde 1994b), and O. nypae remains the only intertidal species of
the 41 species accepted in the genus. Subsequent papers by Hyde and co-workers
have described or listed several new species from intertidal Nypa palm (Hyde 1988,
1991a,b, 1992a,b,c, 1993; Hyde and Sutton 1992; Jones et al. 1996) and these are
listed in Table 1, with their distribution and appropriate references. A relatively large
number of fungi (59) are now known from Nypa palm, and many of the species may
be specific to this host.
In this study we have investigated the fungi associated with the decomposition
of Nypa fruticans in Kuala Selangor and Morib mangroves, in peninsular Malaysia.
Both aerial and intertidal parts of the palm have been examined and a quantitative
analysis of the fungi occurring on various parts of the mangrove palm has been made.
Percentage occurrence is calculated as in Hyde (1992a).
Materials and methods
Visits were made to Morib mangrove and Kuala Selangor mangroves on 8 July 1997
and 12 July 1997 respectively. A suitable well-developed stand of Nypa fruticans was
selected and decomposing samples were collected randomly from intertidal and aerial
parts of the palm. Care was taken to select decomposing parts in each habitat. Intertidal inflorescences, leaves, leaf veins, rachides, and leaf base material were collected
from or close to the mangrove floor, while aerial rachides were collected well above
the high tide mark. All samples were returned to the laboratory in snap lock plastic
bags. Palm parts were incubated separately in sterile plastic boxes on moist sterile
tissue paper and incubated for up to 7 days during which time they were examined
for higher fungi. The fungi were identified and voucher slides of the specimens are
held at the Institute of Biological Sciences, at the Universiti of Malaya, with some
replicates at HKU(M).
Results
Forty-one fungi were identified in this study associated with decomposing parts of the
mangrove palm Nypa fruticans. Thirty-eight species were intertidal species, while a
further 3 species are probably confined to aerial parts. The intertidal fungi included
32 ascomycetes, 4 mitosporic fungi and 2 basidiomycetes (Tables 2 and 3). There
were low numbers of aerial fungi (as it was very dry at the time of collecting) and the
3 species encountered were ascomycetes. The inflorescences were woody and were
the only samples that were colonised by marine borers.
The greatest diversity of fungi, which comprised mostly ascomycetes, occurred
on the intertidal leaf bases, followed by the intertidal rachides (and the upper petiole)
Table 1. Fungi associated with Nypa fruticans.
Taxa
Aerial or intertidal
Palm part
Known distribution on Nypa
References
Aniptodera chesapeakensis Shearer & M.A. Mill.
Aniptodera intermedia K.D. Hyde & Alias
Aniptodera nypae K.D. Hyde
Anthostomella nypae K.D. Hyde, B.S. Lu & Alias
Anthostomella nypensis K.D. Hyde, B.S. Lu & Alias
Anthostomella nypicola K.D. Hyde, B.S. Lu & Alias
Apioclypea nypicola K.D. Hyde
Arecophila nypae K.D. Hyde
Astrosphaeriella nipicola (Cooke & Massee)
K.D. Hyde & J. Fröhl.
Astrosphaeriella nypae K.D. Hyde
Astrosphaeriella striatispora (K.D. Hyde) K.D. Hyde
Carinispora nypae K.D. Hyde
Cucullosporella mangrovei (K.D. Hyde &
E.B.G. Jones) K.D. Hyde & E.B.G. Jones
Fasciatispora nypae K.D. Hyde
Fasciatispora petrakii (Mhaskar & V.G. Rao) K.D. Hyde
Frondicola tunitricuspis K.D. Hyde
Halosarpheia abonnis Kohlm.
Halosarpheia marina (Cribb & J.W. Cribb) Kohlm.
Halosarpheia ratnagiriensis S.D. Patil & Borse
Halosarpheia retorquens Shearer & J.L. Crane
Halosarpheia viscosa (I. Schmidt) Shearer & J.L. Crane
Helicascus nypae K.D. Hyde
Herpotrichia nypicola K.D. Hyde & Alias
Leptosphaeria australiensis
(Cribb & J.W. Cribb) G.C. Hughes
I
I
I
I
I
I
I
I
R
B
B, In, R
B, R
B
B
M
M
Brunei, Indonesia
Malaysia
Malaysia
Brunei, Malaysia
Malaysia
Malaysia
Malaysia
Malaysia
Hyde (1989, 1992a)
Hyde et al. (1999)
Hyde (1994a); this paper
Hyde et al. (1999)
Hyde et al. (1999)
Hyde et al. (1999)
Hyde et al. (1998)
Hyde (1996)
A
I
I
I
R
R, B
L, R, M, B
R,B
Brunei, Indonesia, Malaysia
Brunei, Malaysia
Brunei, Indonesia, Malaysia
Brunei, Malaysia
Hyde & Fröhlich (1998)
Hyde (1992a); this paper
Hyde (1989) 1992a, 1993; this paper
Hyde (1992a, 1993); this paper
I
I
A
I
I
I
I
I
I
I
I
R
R
R
R
M
B, In, R
R
R
R
B,R
B
Brunei
Brunei, Malaysia
Malaysia
Brunei
Malaysia
Brunei, Malaysia
Brunei
Brunei
Brunei
Brunei; Malaysia
Malaysia
Hyde (1992a)
Hyde (1991a, 1992a, 1993)
Hyde and Alias (1999)
Hyde (1992a)
This paper
Hyde (1992a); this paper
Hyde (1992a)
Hyde (1992a)
Hyde (1992a)
Hyde (1991b, 1992a); this paper
Hyde et al. 1999
I
R
Brunei
Hyde (1992a); Hyde and Jones (1988)
395
396
Table 1. Continued.
Taxa
Aerial or intertidal
Palm part
Known distribution on Nypa
References
Leptosphaeria nypicola K.D. Hyde & Alias
Lignincola laevis Höhnk
Lignincola longirostris (Cribb & J. Cribb) Kohlm.
Lignincola nypae K.D. Hyde & Alias
Linocarpon appendiculatum K.D. Hyde
I
I
I
I
I
B, L, M
B, R, M
B
B
B, R
Hyde et al. (1999)
Hyde (1989, 1992a); this paper
Hyde (1992a, 1993); this paper
Hyde and Alias (1999)
Hyde (1988, 1989, 1992a,c, 1993); this paper
Linocarpon angustatum Alias & K.D. Hyde
Linocarpon bipolaris K.D. Hyde
Linocarpon livistonae (Henn.) K.D. Hyde
Linocarpon longisporum K.D. Hyde
Linocarpon nipae (Henn.) K.D. Hyde
Lulworthia grandispora Meyers
Lulworthia sp. (300–400 lm)
Neolinocarpon globosicarpa K.D. Hyde
Neolinocarpon nypicola K.D. Hyde & Alias
Nipicola carbonispora K.D. Hyde
Nipicola selangorensis K.D. Hyde
Oxydothis nypae K.D. Hyde & Nakagiri
I
I
A, I
I
A, I
I
I
I
A
I
I
I
B
R
B
R
R
In, R
R
B, R
R
R
R
B, R
Malaysia
Brunei, Indonesia, Malaysia
Brunei, Malaysia
Malaysia
Brunei, Malaysia,
Indonesia, Papua New Guinea
Malaysia
Brunei
Malaysia
Brunei
Brunei, Malaysia, Philippines, Thailand
Brunei, Indonesia, Malaysia
Brunei, Malaysia
Brunei, Malaysia
Malaysia
Brunei
Malaysia
Brunei, Malaysia, Thailand
Oxydothis nypicola K.D. Hyde
Phomatospora nypae K.D. Hyde
Phomatospora nypicola K.D. Hyde & Alias
Rhipidocarpon javanicum (Pat.) Theiss. & Syd.
Rosellinia sp.
Savoryella lignicola R.A. Eaton & E.B.G. Jones
Swampomyces sp.
Tirisporella beccariana (Ces.)
E.B.G. Jones, K.D. Hyde & Alias
Verruculina enalia (Kohlm.) Kohlm. & Volkm.-Kohlm.
A
I
I
A
I
I
I
R
B, M
B
L
B, In, R
In, R
B
Brunei, Malaysia
Malaysia
Malaysia
Indonesia, Philippines
Brunei, Malaysia
Malaysia
Malaysia
Hyde and Alias (1999)
Hyde (1992a,c)
This paper
Hyde (1992a,c)
Hyde (1988, 1992a,c, 1993); this paper
Hyde (1989, 1992a); this paper
Hyde (1992a)
Hyde (1992a, 1993); this paper
Hyde and Alias (1999)
Hyde (1992a,b)
Hyde (1994a)
Hyde (1992a, 1994b); Hyde and Nakagiri
(1989); this paper
Hyde (1994b); this paper
Hyde (1993); this paper
Hyde and Alias (1999)
Patouillard (1897)
Hyde (1992a); this paper
This paper
This paper
I
I
B
R
Malaysia, Philippines
Brunei
Jones et al. (1996); this paper
Hyde (1992a)
Vibrissea nypicola K.D. Hyde & Alias
I
B, R
Brunei, Malaysia
Hyde (1992a); Hyde and Alias (1999); this paper
Basidiomycota
Halocyphina villosa Kohlm. & E. Kohlm.
I
In, R
Brunei, Indonesia, Malaysia
Hyde (1989, 1992a); Hyde and
Jones (1988); this paper
I
I
I
I
I
I
I
I
I
In
R
R
R
B
R
M
M
R
Malaysia
Brunei
Brunei
Brunei
Malaysia
Brunei
Brunei, Malaysia
Brunei, Malaysia
Brunei
This paper
Hyde (1992a)
Hyde (1992a)
Hyde (1992a)
Hyde et al. (1999)
Hyde (1992a)
Hyde and Sutton (1992); this paper
Hyde and Sutton (1992); this paper
Hyde and Sutton (1992)
I
I
R
B, R
Brunei
Brunei, Malaysia
Hyde (1992a)
Goh and Hyde (1999); Hyde (1992a);
Hyde et al. (1999)
Mitosporic fungi
Agerita sp.
Cirrenalia pygmea Kohlm.
Cirrenalia tropicalis Kohlm.
Dictyosporium pelagicum (Linder) G.C. Hughes.
Helicorhoidion nypicola K.D. Hyde & Goh
Monodictys pelagica (T.W. Johnson) E.B.G. Jones
Nypaella frondicola K.D. Hyde & B. Sutton
Plectophomella nypae K.D. Hyde & B. Sutton
Pleurophomopsis nypae K.D. Hyde & B. Sutton
Trichocladium alopallonellum (Meyers & R.T. Moore)
Kohlm. & Volkm.-Kohlm.
Trichocladium nypae K.D. Hyde & Goh
A – aerial, I – intertidal, B – Base of petiole, In – Inflorescence, R – rachid, M – midrib of leaf, L – leaf.
397
Taxa
Inflorescence (15)
Leaf (20)
Leaf midrib (20)
Rachis (20)
Petiole base (20)
Aerial parts (11)
Aegerita sp.
Aniptodera intermedia
Arecophila nypae
Astrosphaeriella striatispora
Carinispora nypae
Halosarpheia abonnis
Helicascus nypae
Helicorhoidion nypicola
Herpotrichia nypicola
Leptosphaeria nypicola
Lignincola longirostris
Lignincola nypae
Linocarpon appendiculatum
Linocarpon angustatum
Linocarpon nipae
Lulworthia grandispora
Neolinocarpon nypicolum
Nypaella frondicola
Oxydothis nypae
Oxydothis nypicola
Phomatospora nypae
Phomatospora nypicola
Plectophomella nypae
Rosellinia sp.
Savoryella lignicola
Swampomyces sp.
Trichocladium nypae
Vibrissea nypicola
Total no. species
3
–
–
–
–
–
–
–
–
–
–
–
–
–
–
1
–
–
–
–
–
–
–
12
1
–
–
–
4
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
0
–
–
1
16
–
1
–
–
–
2
–
–
–
–
–
–
–
3
–
–
–
–
5
–
–
–
–
–
6
–
–
–
2
1
–
–
–
–
–
–
–
14
–
8
–
–
–
13
–
–
–
–
–
1
–
3
1
8
–
1
–
1
–
–
1
2
6
1
6
1
1
2
–
–
–
–
–
–
1
1
–
3
–
1
–
1
15
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
20
–
–
4
–
–
–
–
–
–
–
–
2
398
Table 2. Frequency of occurrence of fungi associated with decomposing Nypa fruticans at Kuala Selangor mangrove. Number of samples
examined in brackets.
Table 3. Frequency of occurrence of fungi associated with decomposing Nypa fruticans at Morib mangrove. Number of samples examined in
brackets.
Taxa
Inflorescence (15)
Leaf (20)
Leaf midrib (20)
Rachis (20)
Petiole base (20)
Aerial parts (11)
Aniptodera nypae
Anthostomella nypae
Anthostomella nypensis
Anthostomella nypicola
Astrosphaeriella nypae
Astrosphaeriella striatispora
Carinispora nypae
Fasciatispora petrakii
Halocyphina villosa
Halosarpheia marina
Lignincola longirostris
Lignincola laevis
Linocarpon appendiculatum
Linocarpon angustatum
Linocarpon livistonae
Linocarpon nipae
Lulworthia sp.
Oxydothis nypae
Neolinocarpon globosicarpa
Neolinocarpon nypicola
Tirisporella baccariana
Total no. species
2
–
–
–
–
–
–
–
1
1
–
–
–
–
–
–
–
–
–
–
–
3
–
–
–
–
–
1
–
–
–
–
–
–
–
–
–
–
–
–
–
–
–
1
–
–
–
–
–
10
–
–
–
–
–
9
–
–
–
–
–
–
–
–
–
2
5
–
–
–
1
2
–
–
–
–
–
–
1
–
–
2
1
–
–
–
–
6
9
1
1
1
1
–
1
–
–
1
1
3
–
2
1
–
–
1
1
–
3
14
–
–
–
–
–
–
–
3
–
–
–
–
–
–
–
3
–
–
–
5
–
3
399
400
(Tables 2 and 3). Very few fungi occurred on the leaf material and those that did were
mainly confined to the midrib. Although the diversity of fungi on the inflorescences
was relatively low, the fungi that were present were abundant.
Discussion
This study compares the fungi occurring on the intertidal and terrestrial (non-marine)
fronds of Nypa palm, but does not investigate vertical distribution patterns within the
tidal range. The only other study that has looked at the mycota of intertidal Nypa
palm is that of Hyde (1992a), who reported 43 intertidal fungi from decaying fronds
in Brunei. In this study 41 intertidal fungi are reported.
The most common intertidal species reported by Hyde (1992a), were Linocarpon
appendiculatum, Oxydothis nypae, Lignincola laevis, Linocarpon nipae and
Astrosphaeriella nypae. In this study the intertidal species Aniptodera nypae, Astrosphaeriella striatispora and Lignincola laevis were common at Morib mangrove
and Astrosphaeriella striatispora, Linocarpon appendiculatum, Oxydothis nypae and
Rosellinia sp. were common at Kuala Selangor mangrove. It is not clear why the
common species should vary from one mangrove site to another, but Astrosphaeriella striatispora, Lignincola laevis, Linocarpon appendiculatum and Oxydothis nypae
were common at two of the three sites investigated.
Hyde (1992a) did not look at the fungi on aerial fronds, nor did he differentiate
between palm parts. In this study only 3 fungi were identified from aerial (nonmarine) parts, and this is markedly low. Studies on terrestrial palms have shown
them to have an extremely rich and diverse mycota (Fröhlich and Hyde 1999). The
period before collecting, however, had been extremely dry and this is probably the
reason for the low numbers of fungi collected. Neolinocarpon nypicolum was notably
common on dead and senescent terrestrial petioles. We believe this is an endophyte
that changes to a saprobic lifestyle once the fronds die. In this way not only is it the
first fungus to colonise dead fronds, but it can utilise the moisture in the senescent
petioles before they dry out.
Different parts of the fronds (i.e. leaves, leaf midribs, petioles, petiole bases) were
found to support different fungi. This indicates that some fungi may preferentially
develop on certain tissue types. In fungal biodiversity studies it is therefore important
to examine different plant structures. Similar results have been found with terrestrial
palms (Yanna 1997).
The fungi occurring on aerial non-marine, as compared to intertidal parts of Nypa
palm differed, with only Linocarpon nipae occurring in both habitats. This is in
agreement with previous studies where intertidal fungi in the mangrove were found to
differ from fungi on aerial parts of the same mangrove host (e.g. Sabada et al. 1995;
Poon and Hyde 1998). The fungi developing on decaying terrestrial palm fronds have
been found to differ considerably from those (marine fungi) growing on intertidal
401
fronds (Yanna 1997). This is not surprising, as fungi growing on intertidal parts must
have the ability to grow and reproduce in saline conditions. Marine fungi generally
grow more slowly than terrestrial fungi, but have the ability to grow and reproduce
when totally or periodically submerged in seawater or estuarine water (Kohlmeyer
and Kohlmeyer 1979).
The large number of fungi known to occur on Nypa palm is remarkable considering that Hawksworth (1991) has calculated world numbers of fungal species to be
in the region of 1.5 million, based on, amongst other things, a ratio of one plant host
to six fungal species. The numbers are not incongruous when compared to the large
numbers of fungi known from other palm species (e.g. Livistona chinensis (Yanna
1997); Licuala spp. (Fröhlich and Hyde 1999). However, considering that most of the
fungi found on Nypa palm are intertidal and do not appear to occur on non-palm hosts,
the evidence indicates that there are probably more than 40 species that are unique
to Nypa fruticans. There are only two other species of mangrove palm which may
support these intertidal palm fungi (i.e. Calamus erinaceus, Oncosperma tigillarium;
Tomlinson 1986), and although there are no published reports, limited examination
of decaying parts of these intertidal hosts by KDH have not revealed many similar
fungi. The large number of probable host specific fungi occurring on Nypa fruticans
is significant in relation to global fungal diversity estimates (Hawksworth 1991). This
emphasises the pressing need for a detailed study of the fungi on other intertidal palm
hosts.
Acknowledgements
We would like to thank Helen Leung and A.Y.P. Lee for technical assistance. Siti
Aisyah Alias would like to thank the Institute of Biological Sciences, Universiti
Malaya for the award of a grant to study on Nypa fruticans.
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402
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Hyde KD (1992c) Fungi from palms. I. The genus Linocarpon, a revision. Sydowia 44: 32–54
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Hyde KD (1994a) Fungi from palms. XII. Three new intertidal ascomycetes from submerged palm fronds.
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Hyde KD and Jones EBG (1988) Marine mangrove fungi. Mar. Ecol. (Berlin) 9: 15–33
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