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Author(s)

Biodiversity of saprobic microfungi associated with bamboo in


Hong Kong and Kunming, China

Zhou, Dequn.; .

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Issued Date

URL

Rights

2000

http://hdl.handle.net/10722/65260

The author retains all proprietary rights, (such as patent rights)


and the right to use in future works.

Abstract of thesis entitled


BIODIVERSITY OF SAPROBIC MICROFUNGI ASSOCIATED WITH
BAMBOO IN HONG KONG AND KUNMING, CHINA

submitted by
Zhou Dequn
for the degree of Doctor of Philosophy
at The University of Hong Kong
in November 2000

One hundred and twenty fungi were identified from 34 bamboo species in
Hong Kong and Kunming, China. These included 64 ascomycetes, 1 basidiomycete
and 55 mitosporic taxa. Thirteen taxa, including 2 genera, Bambicola and
Striatodecospora and 1 variety, are new to science. Sixty-six fungi are new records for
Hong Kong and 24 fungi are new to Yunnan. The new species are Annulatascus
angustispora, Apiospora yunnanensis, Arthrinium yunnanensis, Bambicola filiforma,
Frondisphaeria amplispora, Linocarpon arundinarium, L. bambusae, Ophiobolus
bambusicola, Roussoella angustispora, Spadicoides bambusicola Striatodecospora
bambusae and Submersisphaeria bambusicola. The new variety is Ramichloridium
indicum var. microsporium. The new combinations, Ellisembia bambusicola (=
Sporidesmium

bambusicola) and Ellisembia pseudoseptata

(= Sporidesmium

pseudoseptatum) are also made in this study.


The occurrence of saprobes on Phyllostachys bambusoides was studied in
single collections of decaying bamboo in Hong Kong and Kunming. Fifty-nine taxa,
including 37 ascomycetes and 22 mitosporic fungi were identified on 300 bamboo
samples from Hong Kong and 41 taxa, including 29 ascomycetes, 1 basidiomycete and
11 mitosporic fungi were identified on 300 bamboo samples from Kunming. The
similarity indices among the fungal communities from different sampling sites in
Hong Kong or Kunming are high, but those between different regions are low. The
similarity indices among the fungal communities from the same site at different

sampling periods are also low. The fungal diversity index (Shannon-Wiener Index) on
bamboo baits in Hong Kong was 2.65, and is richer than that in Kunming (2.11). The
species richness in Hong Kong was 29.5, and is also higher than that in Kunming
(20.5). The climatic conditions, especially temperature and rainfall in Hong Kong
probably better favor growth and development of fungi than in Kunming.
Host exclusivity and recurrence of saprobes on 6 bamboo hosts were
investigated in Hong Kong and Kunming sites. The majority of species encountered
were rare species, while a few species were common. Species richness, individual
identifications and Shannon-Wiener indices of the fungi in Hong Kong were higher
than in Kunming. Species diversity varied in relation to the substratum. Bambusa
shiuyingiana yielded the richest diversity (25 species) in Hong Kong and the lowest
diversity (12 species) was found on Phyllostachys hetroclata in Kunming. Thirteen
species of the total 56 fungal taxa identified have a high recurrence or exclusivity to
the bamboo hosts. The percentage of host recurrent fungi on the bamboo hosts was
19.6%.
Fungal succession on Bambusa tuldoides was observed in Hong Kong during
August 1998 to September 2000. Fungal succession was effected by seasonality.
Rainfall positively impacted on fungal occurrence, but temperature and relative
humidity appear to have little influence. The fungal communities on bamboo baits
were categorized into early colonizers, middle-stage colonizers, late colonizers,
regular inhabitants and sporadic inhabitants. Fungal communities mainly comprised
rare species and middle-stage colonizers.

BIODIVERSITY OF SAPROBIC MICROFUNGI


ASSOCIATED WITH BAMBOO IN HONG KONG AND
KUNMING, CHINA

by
Zhou Dequn
B. Sc. China; M. Sc, China; MDM, Philippines

A thesis submitted in partial fulfillment of the requirements for


the Degree of Doctor of Philosophy
at The University of Hong Kong.

November 2000

Declaration
I declare that this thesis represents my own work, except where due
acknowledgement is made, and that is has not been previously included in
a thesis, dissertation, or report submitted to this University or to any other
institution for a degree, diploma, or other qualification.

Signed:

Zhou Dequn

ACKNOWLEDGEMENTS
I am extremely grateful to Dr. Kevin D. Hyde, Director of the Centre for
Research in Fungal Diversity, Department of Ecology & Biodiversity, The University
of Hong Kong for his excellent supervision and encouragement. Dr. Lilian L.P.
Vrijmoed, Department of Biology and Chemistry, City University of Hong Kong, is
also thanked for her supervision. The encouragement and helpful discussions of my
supervisors stimulated my interest in mycology and made this thesis possible. Prof.
E.B.G. Jones (UK) is also thanked for useful suggestions on the project. Without his
introduction, I would not have had this opportunity to pursue my Ph.D. studies in
Hong Kong.
I would also like to thank all of my colleagues of the Centre for Research in
Fungal Diversity of the Department of Ecology & Biodiversity, The University of
Hong Kong, for their help and collegiality. Helen Leung deserves my special thanks
for her technical help. I acknowledge Dr. T.K. Goh for his generosity in sharing his
knowledge in taxonomy of mitosporic fungi, Dr. Tess Umali for providing me with
some useful references in bambusicolous fungi and sharing her experience and
knowledge as well as acting as my field guide. I would also like to thank Dr. Bing
Sheng Lu for his help in taxonomy of the genus Anthostomella, Mr. G.J.D. Smith for
his assistance in identifying xylariaceous fungi and technical computer assistance. Dr.
Sally C. Fryar is thanked for her help in statistics and field experimental design. I also
owed to Drs. J.C. Kang, L.D. Guo Y.Z. Wang, My friends, in mycological group, for
their care in my daily life at Hong Kong. I am grateful to all of the other colleagues in
the mycological group for their encouragement and stimulating conversation at our
group meetings, which was very helpful to my study.
I would also like to thank Drs. E.H.C. McKenzie (New Zealand), P. Cannon
(UK), A. Aptroot (Netherlands) for their help in the identification of some fungi and
for reviewing my papers.
I extend my gratitude to Mr. A.Y.P. Lee in the photographic Unit, Faculty of
Science, The University of Hong Kong, for photographic assistance. Mr. CO. Wong
and Y.C. Fan, Field Office at Tai Po Kau Nature Reserve, Department of Agriculture

ii

and Fisheries, Hong Kong SAR, for their generous assistance. Mr. Wong used his
private car and sacrificed holidays to accompany me to collect bamboo fungi around
the New Territories. Prof. Li Dezhu and Peng Hua, of the Kunming Botanical Institute,
Academia, Sinica, Xia Lianhe, of the Botanical Institute of Southern China, Academia,
Sinica, Prof. Fan Guosheng, Mr. Du Fan, Mr. Xue Jiarong, of the Southwest Forestry
College, Kunming, Yunnan, China, Mr. Zhang Boliang, of the Yangzhonghai Forest
Farm, Kunming Forest Beureau, Kunming, Yunnan, China, are thanked for their help
in identifying and collecting bamboo samples. Prof. Zhang Keqin, Director of the Key
Laboratory of Microbial Fermentation of Yunnan, Yunnan University, is thanked for
his hospitality and help when I collected bamboo fungi at Kunming in 1998 to 2000.I
would like to show my appreciation to Prof. Wang Weiqing, of the Chemistry Institute,
Academia, Sinica, Beijing, for helping me develop a computer program for ecological
and statistics analysis, Mr. Cheng Luo, of the National University of Defense
Technology, Changsha, China, for fixing my computer problems and Mr. Li Wenpeng,
the Key Laboratory of Microbial Fermentation of Yunnan, Yunnan University, for
helping print my thesis.
I thank The University of Hong Kong for granting me a Studentship during my
Ph.D. study and a conference trip to attend the IX International Congress of Mycology
held in Sydney, Australia, from 14 to 25 August 1999.
Finally I would like to thank my parents, my wife, and my daughter for their
patience and wholehearted encouragement during this study.

iii

Dedication
This thesis is dedicated to my parents
and
My wife and my daughter
For their encouragement, support and love

CONTENTS
Abstract

Declaration

Acknowledgements

ii

Dedication

Contents

iv

List of Figures

xii

List of Tables

xiii

TABLE OF CONTENTS
CHARPTER 1:

LITERATURE REVIEW

1.1

RATIONALE FOR THIS STUDY

1.2

FUNGAL DIVERSITY ON BAMBOO

1.3

HOST SPECIFICITY, HOST EXCLUSIVITY AND HOST RECURRENCE

1.4

FUNGAL SUCCESSION

21

1.5

PLAN OF THE THESIS

27

REFERENCES
CHARPTER 2:
2.1

28

TAXONOMY

41

INTRODUCTION

41

2.2

MATERIALS AND METHODS

42

2.3

TAXONOMY

44

2.3.1 Ascomycetes

44

2.3.1.1 Amphisphaeriaceae

44

Arecophila

44

Arecophila bambusae

44

2.3.1.2 Annulatascaceae

45

Annulatascus

45

Annulatascus anguslispora (Figs 2.1.1-2.1.12)


Submersisphaeria

45
46

Submersisphaeria bambusicola

46

2.3.1.3 Apiosporaceae

47

Apiospora

47

IV

Apiospora montagnei

47

Apiospora sinensis

48

Apiospora yunnanensis (Figs 2.2.1-2.2.9)

49

Arthrinium state of Apiospora yunnanensis (Figs 2.3.1-2.3.4)

50

2.3.1.4 Diatrypaceae

50

Eutypella

50

Eutypella gliricidia

51

2.3.1.5 Didymosphaeriaceae

51

Roussoella

51

Roussoella anguslispora (Figs 2.4.1 -2.4.8)

52

Roussoella hyslerioides

53

Roussoella intermedia

54

Roussoella pustulans
Didymosphaeria

55
56

Didymosphaeria futilis

56

Didymosphaeria massarioides

57

2.3.1.6 Hyponectriaceae

58

Oxydothis

58

Oxydothis aequalis

58

Oxydothis oraniopsis

59

Linocarpon

60

Linocarpon arundinarium (Figs 2.5.1-2.5.12)

61

Linocarpon bambusae (Figs 2.6.1-2.6.11)

61

Linocarpon livistonae

62

Linocarpon pandani

63

2.3.1.7 Lasiosphaeriaceae

64

Ophioceras

64

Ophioceras sorghi

65

2.3.1.8 Leptosphaeriaceae

65

Ophiobolus

65

Ophiobolus bambusicola (Figs 2.7.1-2.7.6)

66

2.3.1.9 Lophiostomataceae

67

Massahna

67

Massarina arundinariae

67

Massarina desmonci

68

Massarina hepaticarum

68

Massarina cf. immersa

69

Massarina palmicola

70

Massarina talae

70

2.3.1.10 Melanommataceae

70

Astrosphaeriella

70

Astrosphaeriella cf. bakerian

70

Astrosphaeriella

fissuristoma

71

Astrosphaeriella maculans

71

Astrosphaeriella splendida

72

Astrosp/iaeriella stellata

72

Astrosphaeriella trochus

73

2.3.1.11 Myxotrichaceae

73

Myxotrichum

73

Myxotrichum berkeleyi

74

2.3.1.12 Thyridiaceae

74

Thrydium

74

Thyridium chrysomallum

74

2.3.1.13 Valsaceae

75

Endothia

75

Endothia gyros a

75

2.3.1.14 Xylariaceae

76

Anthostomella

76

Anthostomella bromeliaceae

76

Anthostomella bruneiensis

77

Anthostomella caffrariae

77

Anthostomella contaminans

77

A nthostomella

flagellariae

78

Anthostomella longa

79

Anthostomella irregularispora

80

Anthostomella raphiae

80

Anthostomella rehmi

81

Anthostomella sepelibilis

81

Anthostomella uniseriata
Astrocystis

81
82

Astrocystis cocoes

82

Astrocystis mirabilis

83

Astrocystis palmarum

84

Hypoxylon

84

VI

Hypoxylon aucklandiae

85

Hypoxylon brevirimum

86

Hypoxylon cf. hughesii

86

Hypoxylon cf. intermedium

87

Hypoxylon karii

87

Hypoxylon rubiginosum

88

Hypoxylon subgilvum

88

Kretzschmariella

89

Kretzschmariella culmorum

89

Rosellinia

89

Rosellinia buxi

90

Rosellinia saccardii

90

Spirodecospora

91

Spirodecospora bambusicola

91

Striatodecospora

91

Striatodecospora bambusae

91

2.3.1.15 Familia Incertae Sedis

91

Bambicola

91

Bambicola filiforma (Figs. 2.8.1-2.8.7)


Frondisphaeria

92
93

Frondisphaeria amplispora (Figs 2.9.1-2.9.10)


2.3.2 Basidiomycetes

93
94

2.3.2.1 Tricholomataceae

94

Marasmius

94

Marasmius rotula

94

2.3.3 Coelomycetes (mitosporic fungi)

95

Conidiocarpus

95

Conidiocarpus caucasicus

95

Hyalopycnis

96

Hyalopycnis blepharistoma

96

Pestalotiopsis

96

Pestaloliopsis uvicola

97

2.3.4 Hyphomycetes (mitosporic fungi)

97

Acrodictys

97

Acrodictys bambusicola

98

VII

Acrodicytys erecta

98

Acrodictys fuliginosa

98

Acrodictys sacchah

99

Alternaria

99

Alternaria alternat

99

Arthrinium

100

Arthrinium state of Apiospora montagnei

100

Arthrinium saccharicola

101

Arthrinium yunnanensis (Figs 2.10.1-2.10.8)

101

Brachysporiella

102

Brachysporiella pulchra

102

Cordelia

103

Cordelia coniosporioides

103

Cordelia johnstonii

103

Dictyosporium

104

Dictyosporium zeylanicum

104

Didymobotryum

104

Didymobotryum rigidum

105

Didymobotryum verrucosum

105

Doratomyces

106

Doratomyces microsporus

106

Ellisembia

106

Ellisembia bambusicola (M.B. Ellis) D.Q. Zhou and K.D. Hyde, comb. nov
Ellisembia coronata

106
107

Ellisembia pseudoseptata (M.B. Ellis) D.Q. Zhou and K.D. Hyde, comb. nov

107

Endophragmiella

108

Endophragmiella oblonga

109

Endophragmiella subolivacea

109

Exserticlava

109

Exserticlava vasiformis

110

Gilmaniella

110

Gilmaniella bambusae

110

Graphium

111

Graphium putredinis

112

Janetia

112

VIII

Janetia synnematosa

112

Penzigomyce

112

Penzigomyces

flagellata

Penzigomyces napacae

113
113

Periconia

113

Pehconia minutissim

114

Phaeoacremonium

114

Phaeoacremonium aleophilium

115

Phaeoacremonium parasiticum

115

Phaeoisaria

116

Phaeoisaria clematidis

116

Phialogeniculata

117

Phialogeniculata dimorpha

117

Plevrophragmium

118

Pleurophragmium simplex

118

Podosporium

119

Podosporium elongatum

119

Podosporium nilgirense

120

Ramichloridium

121

Ramichloridium indicum var. microsporum (Fig 2.11.1)

121

Ramichloridium musae

122

Repetophragma

123

Repetophragma subulata

123

Spadicoides

123

Spadicoides bambusicola
Spadicoides cordanoides

124
124

Spadicoides xylogena

124

Sporidesmium

125

Sporidesmium ehrengergii

125

Sporidesmium eucalypt

125

Sporidesmium eupatoriicola

126

Sporidesmium fragilissimium

126

Sporidesmium penzigii

126

Sporidesmium verrucisporum
Stachybotrys

127
127

IX

Stachybotrys dichroa

127

Stachylidium

128

Stachylidium verticillatum

128

Stilbella

128

Stilbella bambusae

129

Tetraploa

129

Tetraploa aristata

129

Veronaea

130

Veronaea coprophila

130

Veronaea parvispora

130

References

131

CHARPTER 3

BIOGEOGRAPHICAL STUDY

146

3.1

INTRODUCTION

146

3.2

MATERIALS AND METHODS

148

3.3

RESULTS

152

3.4

DISCUSSION

159

3.5

LIMITATIONS TO THIS STUDY AND FUTURE WORK

References
CHARPTER 4

163

164
HOST RECURRENCE AND BIODIVERSITY

167

4.1

INTRODUCTION

167

4.2

MATERIALS AND METHODS

169

4.3

RESULTS

170

4.4

DISCUSSION

181

4.5

LIMITATIONS TO THIS STUDY

185

4.6

FUTURE STUDIES

185

References

186

CHARPTER 5

FUNGAL SUCCESSION

190

5.1

INTRODUCTION

190

5.2

SITE DESCRIPTION

191

5.3

MATERIALS AND METHODS

191

5.4

RESULTS

192

5.5

DISCUSSION

199

5.6

LIMITATIONS TO THIS STUDY

204

References

205

CHARPTER 6
6.1

GENERAL CONCLUSIONS

DOES DECAYING BAMBOO SUPPORT A HIGH FUNGAL DIVERSITY?

208
208

6.2

ARE FUNGAL COMMUNITIES GEOGRAPHICALLY DIFFERENT?

209

6.3

DOES A ' CORE GROUP OF FUNGI' EXIST ON BAMBOO?

209

6.4

ARE SAPROBES ON BAMBOO RECURRENT OR EXCLUSIVE?

210

6.5

FUNGAL SUCCESSION

210

6.6

FUTURE STUDIES

210

References

212

XI

LIST OF FIGURES
Fig. 3.1 Species-area curve of fungi on Phyllostachys bambusoides in Hong Kong

156

Fig. 3.2 Species-area curve of fungi on Phyllostachys bambusoides in Kunming

156

Fig. 3.3 Fungal diversity on Phyllostachys bambusoides In Hong Kong and Kunming

158

Fig. 4.1 Species-area curves of the fungi on Bambusa shiuyingiana in Hong Kong

172

Fig. 4.2 Species-area curves of the fungi on Bambusa textilis in Hong Kong

172

Fig. 4.3 Species-area curves of the fungi on Bambusa tuldoides in Hong Kong

172

Fig. 4.4 Species-area curves of the fungi on Fargesia yunnanensis in Kunming

173

Fig. 4.5 Species-area curves of the fungi on Neosinocalamus affinis in Kunming

173

Fig. 4.6 Species-area curves of the fungi on Phyllostachys hetroclata in Kunming

173

Fig. 4.7 Taxonomic groups of fungi on Bambusa spp. in Hong Kong

176

Fig. 4.8 Taxonomic groups of fungi on bamboo hosts in Kunming

176

Fig. 5.1 Species-area curve of the fungi on naturally dead bamboo in December 1998

197

Fig. 5.2 Species-area curve of the fungi on naturally dead bamboo in June 2000

197

Fig. 5.3 Climatic factors and frequency of occurrence of fungi on bamboo baits and naturally dead
bamboo samples

198

XII

LIST OF TABLES
Table 2.1 Bamboo hosts and collecting regions

43

Table 2.2 Comparison of Oxydothis aequalis collections with the type

59

Table 2.3 Comparison of Linocarpon species from bamboo in this thesis

63

Table 2.4 Comparison of Ophiobolus bambusicola with other similar species

67

Table 2.5 Comparison of Frondisphaeria species

94

Table 3.1 Frequency of occurrence of fungi in Hong Kong and Kunming (1999)

154

Table 3.2 Frequency of occurrence of fungi in Hong Kong and Kunming (2000)

155

Table 3.3 Fungal species diversity in Hong Kong and Kuming

157

Table 3.4 Similarity analysis (Bray-Curtis Coefficient) of mycota identified from different
sampling sites

157

Table 3.5 Similarity analysis (Bray-Curtis Coefficient) of mycota identified from different
sampling sites

157

Table 3.6 Similarity analysis (Bray-Curtis Coefficient) of mycota identified in Hong Kong at
different sampling periods

158

Table 3.7 Similarity analysis (Bray-Curtis Coefficient) of mycota identified in Kunming at


different sampling periods

158

Table 3.8 Comparison of climatic factors in Hong Kong and Kunming in July 1999 and September
2000

160

Table 3.9 Core group of fungi on Phyllostachys bambusoides in Hong Kong and Kunming

163

Table 4.1 Bamboo hosts sampled in Hong Kong and Kunming

169

Table 4.2 Frequency of occurrence of fungi on Bambusa in Hong Kong

174

Table 4.3 Frequency of occurrence of fungi on bamboo in Kunming

175

Table 4.4 Summary of species diversity on 3 hosts from Hong Kong

177

Table 4.5 Summary of species diversity on 3 hosts from Kunming

177

Table 4.6 Similarity analysis (Bray-Curtis coefficient) of mycota on bamboo in Hong Kong

178

XIII

Table 4.7 Similarity analysis (Bray-Curtis coefficient) of mycota on bamboo in Kunming

178

Table 4.8 Similarity analysis (correlation coefficient) of mycota on bamboo in Hong Kong

179

Table 4.9 Similarity analysis (correlation coefficient) of mycota on bamboo in Kunming

179

Table 4.10 Pearson correlation of saprobic fungi on bamboo in Hong Kong

180

Table 4.11 Pearson correlation indices of saprobic fungi on bamboo in Kunming

180

Table 4.12 Recurrent fungi on bamboo hosts in Hong Kong and Kunming

184

Table 5.1 Frequency of occurrence of fungi on naturally dead bamboo

195

Table 5.2 Frequency of occurrence fungi on bamboo baits

196

Table 5.3 Fungal species diversity on naturally dead bamboo and bamboo baits

197

Table 5.4 Ecological classification of fungi on bamboo baits

197

Table 5.5 Similarity analysis of fungal communities on naturally dead bamboo samples

199

Table 5.6 Similarity analysis of fungal communities on bamboo baits

199

APPENDIX 1
APPENDIX 2
PUBLICATIONS (inside back cover)

XIV

Chapter 1: Literature Review

CHAPTER 1: LITERATURE REVIEW


ABSTRACT
The diversity of fungi on bamboo, concepts of host specificity and fungal
succession are reviewed in this chapter. The number of fungi recorded from bamboo
hosts has been estimated at about 1350 taxa. Of these, 189 species have been recorded
in mainland China, from 75 bamboo genera. One hundred and ten species of
bambusicolous fungi are now known from Hong Kong. The most common fungi are
ascomycetes. following by mitosporic fungi and basidiomycetes.
Evidence for host specificity, which is basic to our understanding of fungal
species and host to fungus ratios, is explored in this chapter. There is considerable
evidence that some endophytes, pathogens and mycorrhizae are host specific. Host
specificity however, may be an unwise term for saprobic fungi, and "host exclusivity"
and "host recurrence" are proposed in this chapter as appropriate terms. Concepts of
host specificity, host exclusivity and host recurrence are therefore defined and
discussed. Suggestions for future work needed in order to establish host exclusivity
and host recurrence in saprobic fungi are also made.
Fungal succession is the sequence in which fungi replace one another as their
dynamic communities alter temporally and spatially. Fungal successions on different
substrata, mainly in terrestrial environments, are discussed. The problems in
investigating fungal succession are also addressed.

Key words: bambusicolous fungi, host recurrence, host specificity, succession.


1. 1 RATIONALE FOR THIS STUDY
1.1.1 Bamboo in the world
Bamboos range from short, grass-like plants, to large woody plants as high as
30 meters tall, with individual culms up to 30 cm diameter (Hui, Du and Yang, 1996).
Bamboos are placed in the Gramineae, tribe Bambuseae, subfamily Bambusoideae.
They often have a tree-like habit and can be characterized as having woody, usually
hollow culms, complex rhizomes and branching systems, petiolate leaf blades and

Chapter 1: Literature Review

prominent sheathing organs (Dransfield, 1995). Over 1000 bamboo species in ca 80


genera have been recorded worldwide, but bamboos are predominantly distributed in
the tropics and subtropics of Asia (Xue et. al, 1995). Bamboos are versatile
commodities used in making domestic dwellings, handicrafts, pulp and paper. They
are also used as structural support e.g. scaffoldings for different construction works in
Hong Kong), while some bamboo shoots are edible (Xue et. al, 1995).
Bamboos grow one third faster than the fastest growing tree (Sastry, 2000),
which suggests they can be used in reforestation of degraded areas and generate more
oxygen than an equivalent stand of trees. In tropical forest areas, overlogging has
caused serious environmental problems (Sastry, 2000). Developing the use of bamboo
to replace trees, especially in natural forests in tropical regions, is very important and
may reduce worldwide wood shortages and solve environmental problems (Xue et al.,
1995).
The international society has recently paid some attention to the utilization of
bamboo resources (Xue et al., 1995). On 6 November 1997, Canada and ten other
countries including China signed a treaty establishing the first international
development and research organization for bamboo. The International Network for
Bamboo and Rattan (INBAR) has its headquarters in Beijing, China. It is thus relevant
that a more intensive study of bamboo and related investigations in China should be
undertaken.
1.1.2. Bamboo in China
China has the richest bamboo diversity in the world and has more than 400
species in 40 genera. There are approximately 20,000,000 hectares of bamboo forest in
the world and about 7,000,000 hectares are in China, which therefore accounts for
about 35% of the world's total area of bamboos (Hui et. al., 1996).
Yunnan is the richest province for bamboo diversity in China. There are 210
woody bamboo species in 28 genera naturally distributed in Yunnan, which represents
40% of the total genera in the world and 70% of those in China (Xue et. al., 1995). In
Hong Kong, there are 57 bamboo species in 13 genera, accounting for 16.3% of the
world total and 46.4% of the total in China (But, Fung and Hu, 1985).

Chapter 1: Literature Review

1.1.3 Bambusicolous fungi


Microfungi, especially saprobic microfungi, are rarely considered

for

conservation (Cannon, Mibey and Siboe, 2000). They are small, poorly known,
extremely diverse, and their fruit bodies are often ephemeral. They are frequently
perceived as not being charismatic, and even as threats to other species. Fungi
represent an enormous spectrum of genetic and metabolic resources. Many species
from different nutritional groups (saprobes, parasites and symbionts) play important
roles in the ecosystem through interactions with other organisms. It is therefore crucial
to study all forms of fungi including saprobic microfungi on bamboo.
Although there is a rich diversity of bamboo in China, there has been little
work on bambusicolous fungi. To date 189 species of fungi in 75 genera have been
recorded on bamboo in mainland China (Tai, 1979; Kuai, 1996; Teng, 1996, Zhou,
Hyde and Vrijmoed, 2000). Only 18 bambusicolous fungi have been reported in
Yunnan (Tai, 1979; Kuai, 1996; Teng, 1996; Zhou, Hyde and Vrijmoed, 2000). Dalisay
(1998) reported 51 fungi, comprising 14 ascomycetes and 37 mitosporic fungi on
Bambusa and Dendrocalamus in Hong Kong. Leung (1998) also described 37 fungal
species on Bambusa in Hong Kong. Other reports include Zhou, Goh and Vrijmoed
(1999) who reported a new species, Spadicoides bambusicola D.Q. Zhou, Goh and
K.D. Hyde and another five new records for Hong Kong. Submersisphaeria
bambusicola D.Q. Zhou and K.D. Hyde, a new species from Arundinaria hindsii in
Hong Kong, was also described from Hong Kong by Zhou and Hyde (2000). A new
genus and species, Striatodecospora bambusicola was also reported on bamboo from
Hong Kong by Zhou, Hyde, and Lu. (2000). These few reports indicate that our
knowledge of fungi on bamboo is poor. Most fungi are listed in checklists. Other
relevant information is scattered in regional publications in Asia (e.g. Tai, 1979).
Bambusicolous fungi in China have been poorly studied (Kuai, 1996; Zhou, Hyde and
Vrijmoed, 2000). With this in mind a study was therefore initiated to provide data of
the diversity of fungi on the rich sources of bamboo in Hong Kong and Kunming.

Chapter 1: Literature Review

1.2 FUNGAL DIVERSITY ON BAMBOO


1.2.1 Global estimates of bambusicolous fungi
Dalisay (1998) and Leung (1998) listed bambusicolous fungi reported globally.
There were more than 1200 species of fungi recorded on bamboo, comprising 587
ascomycetes, 145 basidiomycetes and 526 mitosporic fungi (Dalisay, 1998). A further
303 species have been recorded from bamboo since 1998 (Boidin, 1998; Goh, Hyde,
and Dalisay. 1998; Goh, et al., 1998; Leung, 1998; Dalisay, 1998; White and Reddy,
1998). This includes 224 ascomycetes, three basidiomycetes and 76 mitosporic fungi
on 20 bamboo genera (Appendix 1). Among these 303 species of bambusicolous fungi,
16 are new species to science (Dalisay, 1998) and 206 are new records.
Eriksson and Yue (1998) listed all pyrenomycetes described from bamboos, in
all 585 species including 6 varieties. They found that the most intensively studied
areas for bambusicolous pyrenomycetes were Japan (ca. 28% of the type collections),
South America (ca. 20%), the Philippines (ca. 14%), and Indonesia (ca. 9%). There
are about 20-25 species described from each of Central America, China, India and the
United States, but very few from Africa and other tropical and subtropical parts of the
world (Eriksson and Yue, 1998). In mainland China, the most intensive areas where
bambusicolous fungi have been studied are Jiangsu, Guangxi and Sichuan with 36, 35
and 27 fungal species recorded respectively (Tai, 1979; Teng, 1996).

1.2.2 Diversity of bambusicolous fungi in China


1.2.2.1 Mainland China
Oedocephalum glomerulosum (Bull.) Sacc. var. cantonense Teng was the first
fungus reported from bamboo on a culm of Phyllostachys in Guangdong, China in
1932 (Tai, 1979; Teng, 1996). Between 1934 to 1938, 82 species of bambusicolous
fungi were reported by Teng and Ou (Teng, 1996), notably in the paper of Teng and
Ou (1936), where 38 bambusicolous fungi were recorded. Kuai (1996) listed 190
pathogenic bambusicolous fungi in Mainland China, but did not give detailed
descriptions of hosts or fungi. There are 11 pathogenic bambusicolous fungi known
from Yunnan (Kuai, 1996). Of the bambusicolous pyrenomycetes listed by Eriksson
and Yue (1998), 20-25 were from China.

Chapter 1: Literature Review

A review of the literature has revealed that 189 fungal species in 75 genera
have been recorded on bamboo in mainland China (Zhou, Hyde and Vrijmoed, 2000).
The most common fungi are ascomycetes, following by mitosporic fungi and
basidiomycetes (Appendix 2). The most common families are the Pucciniaceae with
20 species, followed by the Phyllachoraceae (15 spp.), Xylariaceae (9 spp.)
Capnodiaceae (4 spp.) and Ustilaginaceae (4 spp.). The genera which have been
recorded most often are Puccinia (10 species), Phyllachora (7 species) and Meliola (6
species). Of the mitosporic fungi found on bamboo in mainland China, Coniosporium
and Fusarium have three species and are the most rich genera in terms of species
diversity.
There are ten bamboo genera from which fungi are known in mainland China
(Xue et al., 1995). These are Bambusa, Phyllostachys, Arundinaria, Pseudosasa, Sasa,
Dendrocalamus, Brachystachyum, Chimonobambusa, Pleioblastus and Yushania
species. There are still 23 genera of bamboo from which no fungi are recorded.
Yunnan has 18 bamboo genera without fungal records, which accounts for 78% of the
total bamboo genera in Yunnan lacking fungal records.

1.2.2.2 Hong Kong


One of the earliest records of bambusicolous fungi from Hong Kong is
Puccinia tenella from Bambusa sp. (Cummins, 1971). Subsequently, Rosellinia
congesta was reported from Bambusa sp. (Laessoe and Spooner, 1994) and
Sporoschisma mirabile was also found on submerged dead culm of Bambusa sp. (Goh,
Hyde and Dalisay, 1997). Dalisay (1998) identified 51 fungal taxa from Bambusa and
Dendrocalamus spp. Leung (1998) also found 37 bambusicolous fungi on unidentified
bamboo. Taking to account the overlap between fungi identified by Dalisay (1998) and
Leung (1998), 91 species of bambusicolous fungi are now known from Hong Kong.
Recently, Umali et al (1999) reported 19 endophytic taxa from leaves of Bambusa
tuldoides and thus to date, 110 fungi have been recorded on bamboo in Hong Kong.
New

species,

Spadicoides

bambusicola,

Striatodecospora

bambusae,

Submersisphaeria bambusicola and other 5 new records to Hong Kong were described
(Zhou et al, 1999, Zhou, Hyde and Lu, 2000, Zhou and Hyde, 2000).

Chapter 1: Literature Review

Bambusicolous fungi in Hong Kong are mostly xylariaceous (5 species),


amphisphaeriaceous (3 species) and lasiosphaeriaceous (3 species). Annulatascus has
the highest species diversity (3 species), followed by Anthostomella, Astrosphaeriella,
Rosellinia and Roussoella (2 species respectively). With the exception of Acrodictys,
which has 5 species recorded from bamboo, most of the mitosporic genera known to
occur on bamboo in Hong Kong were represented by one species.

1.2.2.3 Estimation of number of bambusicolous fungi in China


Hawksworth (1991) used amongst other evidence, a ratio of ca. six unique fungi to
every plant species, to estimate that there were 1.5 million fungal species globally. If
this ratio is applied to fungi on bamboo in Mainland China, where there are 400-500
bamboo species, there should be between 2,400-3,000 unique fungi (Hui et al., 1996).
We presently only know 189 species. Sawada (1961) reported 33 fungal species from
13 bamboo hosts In Taiwan. This included 8 ascomycetes, 14 basidiomycetes and 11
mitosporic fungi. While, at present, 149 species of bambusicolous fungi are known
from Taiwan (http://nt.ars-grin.gov/fungaldatabases, 2000). There are ca. 50 bamboo
species in nine genera in Taiwan (Huang, 1993, 1994) and if the ratio of 6:1 is applied
there should be ca 300 bambusicolous fungi in Taiwan. Fifty-seven bamboo taxa are
distributed in Hong Kong (But et al., 1985). Therefore ca 340 bambusicolous fungi
should occur in Hong Kong. To date 110 bambusicolous fungi (including endophytes,
parasites and saprobes) have been identified which accounts for 30% of the estimated
species.

1.3. HOST SPECIFICITY, HOST EXCLUSIVITY AND HOST RECURRENCE


1.3.1 Introduction
This part investigates whether fungi are host specific, relatively exclusive, or
occur recurrently on certain hosts. Many fungi are considered to be host specific.
There are also numerous examples of saprobic fungi that have only been recorded on a
single host and may be host specific. For example, the mangrove saprobes,
Hypophloeda rhizophora K.D. Hyde and E.B.G. Jones and Rhizophila marina K.D.
Hyde and E.B.G. Jones, have only been found on Rhizophora species, despite the large

Chapter 1: Literature Review

numbers of studies on mangrove fungi (Hyde and Lee, 1995). The fact that these taxa
have only been recorded on a single host, however, is not proof that these fungi only
occur on that host. It could also be argued that if these fungi were isolated and then
inoculated onto different hosts, then they would probably develop on those hosts. Such
an experiment, however, is artificial and probably does not really represent what
happens in nature. Host specificity may therefore not be a particularly good term
relating to saprobes and therefore new terms are needed.
The term host preference has regularly been used by mycologists (Schultz,
Kormanik and Bryan, 1981; Hughes, 1981; Hollins, Jellis and Scott, 1983; Pearson,
Lesline and Schwenk, 1987; Malvick, Grau and Percich, 1998; Augustin et ai, 1999)
to indicate a common occurrence or uniqueness of occurrence of a fungus on a
particular host. This term, however, indicates that a fungus makes a conscious decision
to occur on that host. It is unimaginable that a fungus, like animals, can make a
conscious choice for their food. This is probably why host preference appears far more
often in entomological and zoological literature than in mycological literature. For
example, since 1980, Biological Abstracts has cited 651 scientific papers in which host
preference has been discussed. Among these papers, there were 587 papers in the field
of entomology or animal science. The factors governing why a certain saprobe occurs
regularly or uniquely on a host are rarely understood, but it is certainly not due to a
fungus making a conscious decision to occur on that host. The term host preference is
therefore not suitable for fungal use. Therefore two new more appropriate terms, "host
exclusivity" and "host recurrence" are introduced in this chapter. These terms as well
as host specificity are discussed, and the definitions are given with examples from
saprobic fungi. Evidence of host specificity, host exclusivity and host recurrence in
fungal endophytes, plant pathogens, mycorrhizae and saprobes in terrestrial and
aquatic substrata are provided.

1.3.2 Importance of establishing whether saprobic fungi are host specific


1.3.2.1 Estimates of global fungal numbers
Estimates of fungal diversity are important in order to place value on
threatened natural habitats. This is crucial to prevent unsustainable exploitation of

Chapter 1: Literature Review

these habitats, by disturbances such as over logging, urbanization, or pollution


(Cannon, 1997b). Global estimates of fungal diversity are based on extrapolations of
fungal numbers from the better-known temperate ecosystems. Based on welldocumented literature in the British Isles, Hawksworth (1991) used a ratio of
approximately six fungal species to every plant species, to extrapolate to an estimated
1.5 million fungal species worldwide. These figures were based on the hypothesis that
fungi exhibit a certain degree of host specificity worldwide (Huhndorf and Lodge,
1997). Hawksworth's estimates depend heavily on consistency in the ratios of
phanerogams to fungi in different ecosystems, and the average distribution range of
phanerogams versus fungi (Hammond, 1992).
Most authors appear to have accepted that the estimate of Hawksworth (1991)
is reasonably accurate and the figure of 1.5 million has often been quoted (Rossman,
1994; Cannon, 1997b; Wildman, 1997, Frohlichand Hyde, 1999). Hawksworth (1991)
considered that 1.5 million species may be a conservative estimate, while other
mycologists were skeptical of the large estimate. Huhndorf and Lodge (1997) argued
that Hawksworth's estimates lead to higher than probable global fungal numbers. May
(1991) suggested that fungal species in the tropics might be more closely allied to the
physical variety of tree habitats and the pattern of plant species distribution than to the
number of plant species present. May (1991) argued that fungi might be more
generalized with respect to hosts in the tropics, and that it was therefore unacceptable
to apply a host specific fungal to host species ratio of 6 to 1 to tropical taxa. Huhndorf
and Lodge (1997) further pointed out if May (1991) is correct in his assumptions, then
the fungus/plant ratio in the tropics would be lower and worldwide estimates of
numbers of fungal species would be substantially reduced. Their data indicated that for
large, decayed woody substrates, host specificity or preference might not play a large
part in the distribution of the pyrenomycetes (Huhndorf and Lodge, 1997).
On the other hand, several mycologists working in the tropics have argued that
Hawksworth's (1991) estimate is too low (Lodge, 1997). Cannon (1997a) argued that
there were even more fungi (9.9 million) based on extrapolation of data from
Phyllachora species in the tropics. Frohlich and Hyde (1999) suggested that 1.5
million was a very conservative estimate of the number of fungal species according to

Chapter 1: Literature Review

their investigation of palm fungi in Australia and Brunei. They stated that in the case
of palms, the ratio of 6:1 may be low and that numbers should be assessed upwards.
Following a study of the fungi associated with palms in north Queensland, Hyde (1996)
stated that as many as 26 fungal taxa were likely to be associated with each host
species. He calculated this ratio based on the ca. three plant pathogens, 100
endophytes and 10 saprobes that he expected to develop on each palm species, and a
belief that 25% of these fungi might be host specific (compared with 67% host
specificity assumed by Hawksworth, 1991).
There are many examples of fungal plant pathogens being recorded from only
one host (Shivas and Hyde, 1997). This is particularly true of most Phyllachora
species (Cannon, 1991), and species of Cercospora, Mycosphaerella, and Septoria
(Sutton, 1980; Corlett, 1995; Braun and Sivapalan, 1999). Various researchers have
used presence on a host species or host subspecies as criteria for describing new
species (Zheng and Yu, 1987). Powdery mildews and rusts may also fall in this
category. Zheng and Yu (1987) have used occurrence on a host in delimiting species of
Erysiphales occurring in China. Erysiphe graminis has been divided into subspecific
units, e.g., forms based on host species occurrence (Jenkyn and Bainbridge, 1978).
Species in the Phyllachoraceae have also largely been defined in terms of their plant
partners (Cannon, 1997a). This approach may, however, be incorrect, as although
many plant pathogens are identified based on their host, evidence is not always
available to support this.
Evidence for host specificity in endophytes and saprobes is less compelling.
Endophytes are rarely identified to species level, often remaining identified as mycelia
sterilia. There is little quantifiable evidence in support of endophytes being host
specifici (Brown and Lomolino, 1998), although publications often indicate the
opposite. Saprobes although usually identified to species level, are also generally not
well enough studied to be able to provide meaningful statements in support of host
specificity.
Host specificity is probably the most important single factor used in estimating
global fungal numbers. These estimates rely heavily on the deduction, that host
specificity exists universally in fungi. When Hawksworth (1991) examined the

Chapter 1: Literature Review

number of fungi recorded associated with vascular plants in the British Isles he
suggested that 2/3rds of these fungi may be host specific. Some mycologists have also
used the term "host preference" to indicate that some fungi occur more often on a
particular host, as compared to other surrounding hosts (Hollins et al, 1983; Pearson et
al., 1987; Augustin et al., 1999). We therefore consider that it is important to discuss
the use of these terms and introduce two new terms that we considered more suitable
in discussing the unique or repeated occurrence of fungi on particular substrates.

1.3.2.2 Host specificity, exclusivity and recurrence


Host specificity
In 1923, specificity first appeared in a paper of the British Association for the
Advancement of Science and was defined as the degree to which a parasite or
symbiont was restricted in its range of hosts (Simpson and Weiner, 1989). The concept
of host specificity was originally proposed almost a century ago by plant pathologists
(Rapilly, 1998). In " A Dictionary of Plant Pathology" compiled by Holliday (1998),
"host specific" was defined as a term "describing pathogens that attack certain species
of hosts". In their letter to the editor of Phytopathology, Browder and Eversmeyer
(1986) discussed the relationship between host specificity and resistance/susceptibility.
Specificity is a characteristic of parasite-host associations, rather than of single
organisms within the associations. Plant pathologists often view specificity either from
the host side or from the parasite side, but host specificity usually works in both ways.
Lucas (1998) suggested that host specificity inferred a relationship between hosts and
fungi, and has therefore mainly applied to plant pathogens. Vanderplank (1975)
analyzed the genetic basis of host specificity, which are due to both the parasite
genotype and host genotype influencing the outcome of the relationship. Lucas (1998)
stated that each pathogen is restricted to particular plant types and has a characteristic
host range. Host specificity is therefore a term generally used in the field of plant
pathology. The reader should consult the reviews of Wood (1976), Daly (1979),
Browder and Eversmeyer (1986), Singh and Singh (1988) and Kohmoto, Singh and
Singh (1995) for detailed information concerning host specificity amongst pathogenic
fungi occurring on plants.

10

Chapter 1: Literature Review

Host preference
Preference is a term indicating "which one prefers; the object of prior choice;
the favorite" (Simpson and Weiner, 1989). Some mycologists have used host
specificity and host preference in the same paper and alternate between each term
without giving strict definitions. For example, in her paper "host specificity among
wood-inhabiting pyrenomycetes in a wet tropical forest in Puerto Rico" Lodge (1997)
alternately used host specificity and host preference. So far, it appears that nobody has
defined host preference for use in mycology. Host specificity may not be a good term
for use with saprobes, as "host specific" describes pathogens which cause disease of
certain host species (Lucas, 1998) and host specificity is the reaction between two
living organisms (fungus vs. host). Unfortunately host preference cannot be a more
suitable term to describe the relationship between saprobic fungi and their hosts, as it
cannot simply refer to the occurrence of fungi. Preference is a conscious choice and
fungi are assumed not to have do not have such an ability.
Host specificity can be extended not only to include pathogenic fungi but also
to include other non-saprobic fungi, which are confined to certain hosts. However, in
most cases this relationship cannot be proven and therefore "host exclusivity" may be
a better term. Some fungi occur more frequently on a particular host, as compared to
surrounding hosts, but are certainly not confined to the first host. In this case the term
"host recurrence" may be more suitable. Based on the above discussion, host
specificity, host exclusivity and host recurrence are defined.

Host specificity as the relationship between a plant and a fungus, which is


restricted to the host plant, but does not occur on other plants in the same
habitat (Holliday, 1998).

Host exclusivity as the exclusive occurrence of a saprobic fungus on a


particular host. If other potential hosts in the same habitat are also studied and
the fungus is absent, this gives strength to the notion of host exclusivity.

Host recurrence as the frequent occurrence of a saprobic fungus on a particular


host. If other potential hosts in the same habitat are also studied and the fungus
is infrequent, this gives strength to the notion of host recurrence.

11

Chapter 1: Literature Review

1.3.2.3 Are fungi host specific?


Host specific plant pathogens
There is much evidence to support the view that fungal pathogens are host
specific and this has been reviewed by Wood (1976), Daly (1979) and Singh and Singh
(1988). The basis for host specificity lies on genetic basis, e.g., gene-for-gene
association in the diseases and the pathogenic fungi (Singh and Singh, 1988; Singh,
Singh, and Kohmoto, 1995) and also host-specific toxins, which are produced by the
pathogens, which are toxic to some hosts but not others (Browder and Eversmeyer,
1986). There is also cladistic, genetic, molecular and chemical evidence that fungi are
host specific (Hart, 1988; Otani et al, 1995, 1998; Wada et al, 1996; Ellis et al., 1997;
Oyarzun^o/.. 1998).
Evidence of host specificity amongst pathogens has resulted in numerous fungi
being described based on host occurrence (Zheng and Yu, 1987; Cannon, 1991; Shivas
and Hyde. 1997). There are about 1614 species of Mycosphaerella, 710 species of
Colletotrichum. 913 species of Phomopsis and 206 species of Pestalotiopsis (Funindex:
http: 194.131,255.3/cabipages/Names/NAMES.ASP, 2000). The large numbers of
epithets in these genera are due to the fact that mycologists have historically described
new species based on host. Recent evidence has shown that this practice may have
been unwise (Jeewon, personal communication), although Crous (2000) have found
that by using molecular sequencing, many pathogenic Mycosphaerella species deserve
species recognition. Dennis, Faeth and Wingfield (1999) have shown that isolates of
endophytes were not distinguishable on the basis of molecular studies (ITS and
presence of ds RNA), but could be separated based on other characteristics (e.g.
pathology) when they examined differences between isolates from completely healthy
tree tissue and morphologically indistinguishable isolates from diseased tissue in
South Africa.

Host specificity in mycorrhizae


There

are contradictory

observations

concerning

host

specificity

in

mycorrhizae. Some researchers have found that mycorrhizal fungi are symbiotic with a
comparatively wide host range (Malajczuk et al, 1982; Lee and Kim, 1987). Horton

12

Chapter 1: Literature Review

and Bruns (1998) found that most ectomycorrhizal fungi associated with Douglas Fir
(Pseudotsuga menziesii) and Bishop Pine {Pinus muricata) showed no host specificity.
Kropp and Trappe (1982) reported that few mycorrhizal taxa appeared to form host
specific symbiosis with Truga heterophylla (Hemlock). The evidence for lack of host
specificity in endomycorrhizal fungi may have been due to the fact that their
taxonomy was not well known (Kropp and Trappe, 1982).
On the other hand, Newton and Haigh (1998) concluded that ectomycorrhizal
fungi were host specific. They examined the host range of ectomycorrhizal fungi in
Britain by compiling a data matrix from the literature, based primarily on accounts of
sporocarp associations with particular host genera. Galan and Moreno (1998) have
also reported that Ruhlandiella berolenensis, an exotic and subhypogenous fungus in
Europe was specific to Eucalyptus spp. Borowicz and Juliano (1991) compared host
records for ectomycorrhizal fungi (mutualists) to those for biotrophic shoot fungi and
necrotrophic root fungi (both antagonists) and concluded that many species of
ectomycorrhizal fungi were host specific.
Nelson (1979) suggested that evolutionary and ecological processes that
determine specificity act differently on hosts and their fungal symbionts. Each partner
is influenced not only by its symbiotic association, but also by other environmental
factors. The level of host specificity amongst mycorrhizal fungi is dynamic, and
depends not only on symbiotic partners, but also on ecological opportunities.
Some researchers considered that host specificity could occur at different host
taxonomic levels. When Malajczuk et al. (1982) studied ectomycorrhizae formation in
Eucalyptus, they found no evidence for host specific fungal associations with
Eucalyptus species. Pinus radiata and most Eucalyptus species formed associations
with several broad-host-ranging ectomycorrhizae. However, taxa which were known
to associate exclusively with Pinaceae, e.g. Suillus and Rhizopogon species, did not
form ectomycorrhizae with any Eucalyptus species. The reverse situation also did not
occur. They further pointed out that successful, long-term development of exotic
stands of these hosts is largely related to appearance of host specific ectomycorrhizal
fungi, rather than broad host species. After investigation of the ectomycorrhizal genus
Rhizopogon, Massicotte et al. (1994) found that none of the fungal species had broad

13

Chapter 1: Literature Review

host range affinities. A variety of specific responses were exhibited by different fungal
taxa, ranging from genus-restricted to species-restricted species.
The association between mycorrhizae and plants has resulted during
coevolution (Barrett, 1983). A high degree of specificity between the interacting
partners will promote coevolution with each other (Clay, 1988). Barrett (1988)
indicated that mycorrhizal fungi might infect the roots of many plant species and,
conversely, that plant species may be infected by many fungal species. A large
proportion of the variance in fitness will necessarily result from genetic differences
among hosts for species with restricted host ranges (Clay, 1988). Host specificity of
mycorrhizal fungi is therefore probably a consequence of coevolution between the
fungi and plant hosts.
Recently Guillot (1997) considered that the recognition mechanisms between
trees and fungus likewise the causes of the specificity which characterizes many
associations. A recent hypothesis that, not only in the case of ectomycorrhizae but also
in other types of symbiosis, the involvement of a lectin present at the surface of a
partner and of a saccharidic structure belonging to the other, is a possible explanation.
Host specificity amongst endophytes
Endophytes are organisms that live within plants. Often the terms "endophyte"
and "endophytic" are used with particular meaning by different workers and for
particular groups of hosts and microbes (Bacon and White, 2000). Endophyte are
"fungi colonizing living plant tissue without causing any immediate, overt negative
effects" (Bacon and White, 2000). This definition includes virtually the entire
spectrum of symbiotic interactions in which fungi and plants participate: parasitism,
communalism, and mutualism (Bills, 1996). The term "endophyte" has been used in a
variety of ways, giving rise to semantic confusion and ambiguities (Hawksworth et al.,
1995). Mycologically, endophyte refers to fungi that invade the stems and leaves of
plants, but cause no symptoms of disease (Hawksworth et al., 1995). Although some
researchers found no evidence of host specificity in endophytes (Dalisay, 1998), Butin
(1986) and Dominik and Adrian (1999) observed that some endophytes were host
specific.

14

Chapter 1: Literature Review

Petrini (1986) observed that host specificity in endophytes is evident only at


the family level. Reddell and Bowen (1986) found that host-endophyte specificity was
expressed mainly at the generic level. Fisher et al. (1995) found that some endophytic
coelomycetes in the genera Cryptosporiopsis, Gloeosporidiella and Phomopsis were
host specific. Some researchers believed that host specificity were mainly at the
generic level (Reddell and Bowen, 1986). Chapela et al. (1991) demonstrated specific
recognition of two endophytes in the genus Hypoxylon for wound exudates of their
respective hosts, beech and hazel, in contrast to other hosts. Peters et al. (1998) found
that growth of the endophytes was specifically enhanced by calli of the host plant.
Geniculosporium is known as an ubiquitous endophytic genus (Carrol, 1988). Thus, it
was conceivable that isolates of Geniculosporium exhibit little host specificity and
respond positively to both host and non-host calli. Nevertheless, the growth
stimulation was greater in the host than in the non-host combination (Peters et al.,
1998).
Clay (1988) pointed out that the endophytes might play a defensive role that
benefits both partners. Host plants may be subjected to reduce herbivores, because
some endophytes are toxic or "bad taste" to herbivores. This association between host
plants and endophytes enhance the host specificity. This means that, because
herbivores passively "choose" the plants, which have no or few endophytes, this gives
more survival opportunities to the plants in which endophytes are found. Host
specificity probably plays a role in the coevolution between both partners.
Recently studies by Guo, Hyde and Liew (1998, 2000) may provide evidence
for host specificity of endophytes on palms. Palm saprobes are a rather unique group
of fungi with examples of many genera that appear to be unique to, or mostly occur on
palms (e.g. Astrosphaeriella, Monokwaria, Myelosperma, Oxydothis, Palmicola,
Pemphidium). When palm fronds senesce they often remain attached to the tree,
become yellow and then brown and often become dry. This is particularly true of
fronds of Cocos nucifera, which usually grow in xeric regions. In this situation,
Oxydothis and Cocoicola species develop on the fronds before they completely dry out
(Hyde et al, 1997). It appears that Oxydothis and Cocoicola species utilize moisture
within the fronds. These fungi appear to have the ability to do this, while most other

15

Chapter 1: Literature Review

fungi that develop on the palm fronds only do so when the fronds are wet. This
phenomenon indicates that these fungi may have an advantage over other fungi, and
are probably endophytes within the palm, changing to a saprobic lifestyle with the
onset of palm senescence. A similar situation occurs with Pemphidum nypicolum on
Nypa palm. The blackened stroma of this fungus forms throughout the senescing palm
material well before it has the appearance of being rotten (Hyde and Alias, 1999). If
this is the case and many of the saprobes develop from endophytes, then there are
compelling arguments in support of host specificity in saprobic fungi.
Oxydothis species are not named according to their host and there are examples
of species known only from a single host and others known from more than one host
(Hyde. 1994). Oxydothis alexandrae was collected on fronds of Archontophoenix
alexandrae on decaying petioles, but was not identified from Calamus or Licuala in
the same habitat or region. This taxon is clearly a good example of a fungus showing
host specificity. There are few morphological characters on which to distinguish
between Oxydothis species. Species delineation is based on spore size and shape, ascal
ring size and orientation of the ascomata. Some species are very distinct and cannot be
confused (e.g. O. alexandrae). It is probable that Oxydothis species are host specific or
at least specific to a few closely related hosts.

Saprobes
A saprobe is an organism using dead organic material as food, and commonly
causing its decay and is the preferred term for fungi (Hawksworth et al., 1995). Our
knowledge of host specificity of saprobic fungi is mostly based on field observations
from biodiversity assessment (Gonczol, 1975; 1989; Kohlmeyer and Kohlmeyer, 1979;
Sridhar and Kaveriappa, 1988).

Trrestrial fungi
There are many examples of terrestrial fungi being recorded as common on a
single plant host, Family or Order. Francis (1975) emphasized that species of
Anthostomella occurred more frequently on conifers than on Angiosperms.
Hawksworth and Boise (1985) observed that Astrosphaeriella species mostly occurred

16

Chapter 1: Literature Review

on palms, bamboo and other larger monocots, while Kretzschmariella species only
occurred on bamboo culms (Ju and Rogers, 1996). Huhndorf and Lodge (1997) also
observed that Rosellinia bunodes at El Verde, Puerto Rico, occurred more frequently
on plants in the Rubiaceae and Malvaceae. Lodge (1997) found that the larger
ascomycetous fungi in the Xylariaceae were usually restricted to fruiting on leaves and
fruits of particular host plant genera or families in the tropics. Xylaria axifera has been
shown to be specific to dead petioles of Araliaceae (Lasss0e and Lodge, 1994).
Gonzales and Rogers (1989) observed that a high proportion of Xylaria species
growing on fruits in Mexico were host specific. Tokumasu et al. (1994) reported that
Verticicladium trifidum and Sympodiella acicola were specific to pine needles.
Caudatispora palmicola was restricted to palms species, e.g. Phytelaphas sp. and
Jessenia sp. (Frohlich and Hyde, 1995).
Various degrees of host specificity are exhibited by wood-inhabiting
pyrenomycetes in temperate regions (Huhndorf and Lodge, 1997). For example,
Rosellinia buxi occurs only on Buxus wood, and R. herpotrichoides occurs only on
Truga canadensis. Other species of Rosellinia are specific to deciduous or coniferous
hosts or may be confined to certain host genera (Petrini, 1992). Mathiassen (1993)
reported several pyrenomycetes, e.g. Saccardoella kanderana and Amphisphaerella
erikssonii that occurred on species of Salix in Scandinavia. In the temperate regions of
North America, there are many examples of pyrenomycetes growing on certain host
genera, such as Cucurbitaria berberidis on Berberis spp. and C. elongata on
leguminous trees (Barr, 1990). Camillea verruculospora is found only on Miconia sp.
in Puerto Rico and Ecuador (Lodge and Laessoe, 1995). Hyde (1992, 1995) reported
that host specificity of fungi on palms was exhibited by several taxa including species
of Ascotaiwania and Linocarpon. Taylor (1998) observed that Oxydothis was
exclusive to and frequently occurred on Archontophoenix alexandrae.
Wong (2000) investigated saprobic fungal diversity among six grass and one
sedge hosts in Hong Kong. She observed that a group of fungi, including Niptera
excelsior on Phragmi/es australis; Ceratosporella sp., Linocarpon augustatum,
Massarina purpurascens, Phaeoisaria clematidis, and Stachybotrys kampalensis on
Miscanthus floridulus; Paraphaeosphaeria schoenoplecti and Septoria-like sp. on

17

Chapter 1: Literature Review

Schoenoplectus litoralis, occurred extensively on one host only during each sampling
period, with frequencies of occurrence around 50%, showing obvious host specificity.
She thought that the main reasons for host specificity was related to the presence of
host specific endophytes that become saprobes, and/or differences in physical
structures and nutritional levels of the hosts.
When Lodge (1997) studied diversity of decomposer fungi at El Verde in the
Luquillo Mountains of Puerto Rico, she observed that host recurrence (as preference)
were common among certain groups of tropical wood decomposers and strong host
specificity appeared to be rare among wood decomposer fungi. Many wood-inhabiting
ascomycetes had distinct preferences for the state of substratum decay and relative
position to the ground, in addition to substratum and diameter class. Such microhabitat
preferences have also been noted for tropical wood decomposer fungi in Africa
(Ryvarden and Nunez, 1992; Laessoe et al., 1996). Similarly, Hedger (1985) showed
that, in Ecuador, leaf decomposer basidiomycetes from the lower litter required
partially decomposed leaves, in contrast to agarics of the upper layer that required
freshly fallen leaves. The availability of different microhabitats and substrata in
different states of decay are likely to influence the diversity of decomposer fungi in
tropical forests. This was further discussed by Hedger (1985), Lodge and Cantrell
(1995) and Laessoe et al. (1996).

Marine fungi
Kohlmeyer and Kohlmeyer (1979) reported 8 fungi that were specific to
individual host species or genera, and Hyde (1990) observed some fungi (e.g. Aigialus
mangrovis and Caryospora mangrovei) to be specific to a single mangrove tree species.
Jones (2000) reviewed the factors affecting habitat preference in marine fungi, where
he addressed host specificity. He concluded that some fungi occurred more readily on
test blocks of one wood type than another, but overall there was little evidence for host
specificity in marine fungi. This was also true of intertidal mangrove fungi, although
in Brunei, Hyde (1990) found that some fungi were only associated with certain
mangrove species, indicating a host specificity. The fungi occurring on intertidal Nypa
fruticans are however, remarkable in that more than 40 species are unique to this host

18

Chapter 1: Literature Review

(Hyde and Alias, 2000). It is unclear if the same fungi also occur on the other two
intertidal palms, but even if they did the ratio of these palm hosts to specific intertidal
fungi is extremely high. These species are not found on terrestrial palms and new
species are still being discovered on Nypa fruticans (Hyde et al., 1999).
In the case of the saprobic fungi, host preference would probably be better
termed host recurrence. The examples of host specific saprobes are derived from lack
of presence on other hosts, rather than having been proven by Koch's postulates
(Gonczol, 1975; 1989; Kohlmeyer and Kohlmeyer, 1979; Sridhar and Kaveriappa,
1988; Gonzales and Rogers, 1989). Host exclusivity amongst saprobic fungi is
difficult to prove, as all other hosts in the same habitat need investigating. Taylor
(1998) found that Oxydothis was exclusive to Archontophoenix alexandrae as it did
not occur on any other palms in the same habitat. Perhaps this genus has host
exclusivity to Archontophoenix alexandrae. Astrosphaeriella species occur on palms,
bamboo and other larger monocots. This genus may occur exclusively on monocots.

1.3.3 Future work


The notion of host specificity in plant pathology is complex, but well-defined
and future research will modify concepts further. Although many authors have
observed that host recurrence exists in saprobic fungi, there is a lack of chemical and
genetic evidence. Much work has been carried out on species concepts in
Colletotrichum and Phomopsis and the relationship with hosts, and this needs to be
expanded to other large genera such as Pestalotiopsis, Phoma and Septoria.
In the case of endophytes and pathogenic fungi, it may be possible to analyze
phylogenic development based on host plant phylogeny. This is because "parasite
phylogeny mirrors host phylogeny and "the more primitive the host, the more
primitive the parasites that it harbours" and "parasites evolve more slowly than their
hosts" (Mitter and Brooks, 1983). The longer the association with a host group, the
more pronounced the specificity exhibited by the parasite group (Mitter and Brooks,
1983). A good way to detect the phylogenetic routes is to carry out comparative
studies of host phylogeny with that of endophytes or parasitic fungi.
Astrosphaeriella bakeriana is one of the first and most common taxa to appear

19

Chapter 1: Literature Review

on dead fronds and leaves of Livistona chinensis in Hong Kong and is presently not
known from any other host, although it appears to be common throughout the Asian
region (Hyde and Frohlich, 1997). Guo et al. (1998) isolated endophytes from
Livistona chinensis, where Astrosphaeriella bakeriana was a common saprobe, but did
not find this species in their initial studies of endophytes on the host. However, by
careful manipulation of the mycelia sterilia, they were able to identify this species,
which grew on strips of palm petiole in conical flasks. This discovery has shown that
this common saprobe is in fact an endophyte. This has important implications. One of
the most likely scenarios is that endophytes within the plant tissues change their mode
of life once the host tissues die. As Guo et al. (1998) pointed out, some endophytes
and saprobes are interrelated, i.e. some saprobes have a latent period inside plant
tissues, or some endophytes become saprobes after a plant senesces. Wong (2000) also
indicated that the fungal mode of life could alternate from endophytes to saprobes. On
the other hand, some endophytes could also become pathogens under certain condition
(Brown and Lomolino, 1998). Gindrat and Pezet (1994) observed that the endophytes,
e.g. Fusarium spp. could cause latent infections in wheat. Northover and Cerkauskas
(1998) also made similar observations on European plums (Prunus domestica 'Stanley')
with a high incidence of symptomless latent infections of Monilinia fructicolathis. In
this way the endophytes gain an advantage over other non-internal fungi. It is quite
compelling to make clear the mechanism of conversion of life mode in fungi. It is
doubtless that this conversion impacts fungal diversity.
Common palm genera such as Anthostomella, Arecomyces, Arecophila,
Capsulospora and Linocarpon may fall into the category, but this needs to be proven.
If this is the case, as we suspect, these fungi at least, are probably host specific when
they become endophytes. These fungi perhaps, at some stage of their life cycle and
under certain environment conditions, alternate their life mode as endophytes, which
may strengthen the relationship with the host plants and favor the establishment or
development of host specificity. Guo et al. (2000) identified most of the mycelia
sterilia they identified from Livistona chinensis to genus, family or ordinal level using
molecular data and many of these were xylariaceous. These may turn out to be species
of the above genera, but further work is necessary in order to establish this, as none of

20

Chapter 1: Literature Review

these genera have yet been sequenced.


Cannon (1997a) pointed out that lack of knowledge of host specificity for most
species is a major obstacle in estimating fungal diversity even for small areas. There is,
however, even less information on host exclusivity or recurrence (previously termed
preference). Further surveys of various hosts in the same and different habitats are
needed to reveal examples of host exclusivity or recurrence of saprobic fungi. This can
be carried out in two stages: 1) to statistically make observations on various plants and
2) to establish the basis for host recurrence. In the latter case we need to establish if
host exclusivity or recurrence has a chemical base, is based on enzyme production,
results from endophytic infection or is a result of some other factor(s).

1.4 FUNGAL SUCCESSION


1.4.1 Concept of succession
Succession of fungi on decaying substrates can be defined as a directional
change in the composition, relative abundance and spatial patterns of species
communities (Frankland, 1992). Fungal succession is the sequential occupation of the
same site by thalli (normally mycelia) either of different fungi, or of different
associations of fungi. Fungi replace one another as their dynamic communities of
mycelia alter in space and time, each species is adapted for the occupation of
particular niches. Early workers concentrated on replacement of species in time rather
than in space (Frankland, 1998). Frankland (1998) suggested that each succession is
unique, dependent on the host material and its environment.

1.4.2 Fungal succession on angiosperms


1.4.2.1 On dicotyledons
Sugar maple
Fungal populations on green and senescent foliage were dominated by a few
ubiquitous taxa, namely Aureobasidium pullulans, Cladosporium cladosporioides, and
Phoma exigua, that remained abundant for up to eight months after leaf fall (Kuter,
1986). The results showed that the composition of populations that colonized the litter
during at least the first months of exposure, were influenced in part by the prevailing

21

Chapter 1: Literature Review

seasonal conditions. Thus, popular concepts of fungal succession based on changes in


the nutritional composition of the substrate cannot entirely explain the changes in the
composition of micro-fungal populations associated with the decay of the leaves.
Aspen
Wildman and Parkinson (1979) observed micro-fungal succession on surfaces
of attached aspen poplar Populus tremuloides leaves. The outer sheathing bud scales
were colonized by a limited range of fungi, but the enclosed leaves were free from
fungal colonization. The adaxial surfaces of young, newly expanded leaves were
sparsely colonized by fungi. As the leaves matured, they were extensively colonized
by the common phylloplane fungi (such as Alternaria altemata, Aureobasidium
pullulans. Cladosporium spp. and Epicoccum purpurascens) especially on the adaxial
surface.

1.4.2.2 On monotyledons
Sandhu and Sidhu (1980) reported that Aspergillus fumigatus, A. niger, A.
terreus, A. flavus, Mucor pusillus, Penicillium sp. Rhizopus microsporus and
Trichoderma longibrachiatum were isolated from the fungal succession on bagasse.
The most common species were A. fumigatus and A. terreus which were present
throughout the 20 week study period. During the succession, the pH of the bagasse
changed from acidic to neutral and finally became slightly alkaline, while the moisture
content dropped gradually and the maximum temperature reached was 50C.

1.4.3 Fungal succession on gymnosperms


Fungal succession on decaying pine needles has been well documented. Pine
leaf litter has several characteristics that are advantageous in studying fungal
succession (Kendrick, 1959) and this is reviewed by Tokumasu et al. (1994). Fungal
succession of pine needles was first demonstrated by Kendrick and Burges (1962) in
England. Since this work, there have been many studies on the fungal succession on
pine needles in the British Isles. Tokumasu et al. (1994) studied vertical distribution
and fungal succession on decaying needles of Pinus sylvestris in southwest Germany.

22

Chapter 1: Literature Review

Verticicladium trifidim was apparently the most prevalent fungus and Verticicladium
trifidim was the first colonizer of freshly fallen needles. Kendrick and Burges (1962)
noted that the chemical nature of the pine needles determines the fungi involved in the
succession.
Kasai. Morinaga and Horikoshi (1995) investigated fungal succession in the
early decomposition of pine cones on the trees and the floor of Pinus densiflora forest.
Pestalotiopsis spp. were dominant on withered cones attached to the tree. Phomopsis
sp. and Xylaria sp. which colonized the interior tissues, occurred with higher
frequencies on the cones on the tree, but their frequency decreased after cones had
fallen onto the forest floor. Conversely, the occurrence frequencies of Mortierella spp.
and Trichoderma spp. increased on fallen cones, as compared to those on the tree.
Trichoderma koningii numbers increased rapidly and showed high occurrence
frequencies. Thysanophora penicillioides, which prefers coniferous substrates, showed
higher frequencies of occurrence in the early stages of decay of pine cones on the
forest floor. The mycota on cones on the forest floor did not, however, change during
the investigation period.
1.4.4 Fungal succession on baits
Fungal succession on baits has been well documented. On woollen baits,
Ghawana. Shrivastava and Kushwaha (1997) studied fungal succession under
laboratory conditions for more than one year. The initial colonizers on woollen baits
were non-keratinophilic fungi, while the late colonizers were keratinophilic. The
fungal communities found during the decomposition of woollen in soil samples
collected from plain and hilly areas were almost the same, except for the dominant
colonizers, which comprised Chrysosporium tropicum for plains, but Microsporum
gypseum and M. fulvum in the hilly areas.
Leung (1998) used senescent culms of Bambusa mutabilis as baits to observe
fungal succession in terrestrial, freshwater and marine environments in Hong Kong.
He categorized the fungi on baits in terrestrial environment into two groups: early
colonizers, including Arthrinium state of Apiospora montagnei, Giliocladium roseum,
Nectria sp., Periconia minutissima, Phomopsis sp., Stilbella sp., Wiesneriomyces

23

Chapter 1: Literature Review

javanicus are regular inhabitants, i.e., Arthrinium state of Apiospora montagnei and
Periconia minutissima which existed throughout study period. Leung (1998) did not
find any seasonal pattern of occurrence but more fungi occurred in the summer. The
mycota from baits in freshwater mainly comprised sporadic and infrequent fungi.
Leung (1998) also revealed that the mycota from baits in marine habitats were
composed of several dominant and frequent species. Occurrence of the fungi showed a
peak in the summer (Leung, 1998). Ho (1998) used sterilized wood (Machilus velutina
and Pinus massoniana) as baits to examine fungal succession on wood in a freshwater
stream, at Tai Po Kau, Hong Kong. He categorized fungi from the baits into early
colonizers, late colonizers, regular inhabitants and sporadic inhabitants. Ho (1998)
also observed that higher species richness and Shannon-Wiener Index were recorded
in the rainy season.

1.4.5 Biological factors impacting on the succession process


Frankland (1998) observed that decomposing bracken provided new resources
and microhabitats for small animals. After two or more years, Collembola and Acarina
were abundant inside the petioles, grazing voraciously, but selectively on fungal
mycelium and fructifications, pushing the softer tissues aside to form longitudinal
channels and adding feces. These animal activities undoubtedly influenced fungal
succession. Elton (1966) remarked that fungal activity on plant substrata had almost
ceased by the end of 5-6 years when bacteria predominated. Solhein (1992) observed
that infestation of healthy Norway spruce trees by the bark beetle, Ips typographus,
was the crucial factor in fungal succession. The earliest invaders, such as Ophiostoma
pencicillatum and O. ainoae, were the species most frequently carried by the beetles.
Ponge (1991) observed a co-succession of nematodes, amoebae, enchytraeids,
sciarid larvae, oribatid mites and earthworms with fungi on fallen Scots pine litter.
Bacterial development was observed in the L-2 layer, following penetration by microfauna (nematodes, amoebae). After this stage pine needles were actively tunneled by
enchytraeids, sciarid larvae and oribatid mites and at the same time were nibbled on by
epigeic earthworms (L-2 and F-1 layers). When the fine root system of pine developed
through accumulated old needles (F-1 layer), mycorrhizal fungi penetrated the needles

24

Chapter 1: Literature Review

and seemed to impede any further bacterial development. Pine foliar tissues were
progressively incorporated into the fecal material of earthworms and other members of
the soil fauna.
Klironomos et al. (1992) investigated the feeding preferences of two strains of
the collembolan, Folsomia Candida, for some common primary and secondary
saprobes of spruce and fir litter. They found that generally the insects preferred
primary saprobes to secondary saprobes. The microcosm study showed that the
absence of the grazers slowed down that rate at which primary saprobes on litter were
replaced by secondary saprobes. These data therefore support the hypothesis that
fungal successions observed in the field on decaying litter may result from preferential
grazing by micro-arthropods.

1.4.6 Some crucial physical factors responsible for fungal succession


1.4.6.1 PH
When Sandhu and Sidhu (1980) observed fungal succession on decomposing
sugarcane bagasse, they found that during succession, the pH of the bagasse changed
from acidic to neutral and finally became slightly alkaline. The moisture content
gradually dropped and maximum temperature reached was 50 C. Kanaujia (1981)
however, found a poor correlation between moisture content and fungal succession
when he investigated decay of aboveground parts of Pennisetum typhoides.
1.4.6.2 Moisture
Reddy and Reddy (1983) reported on fungal succession on seeds of Sesamum
indicum with different moisture levels. The percentage incidence of fungi varied with
the moisture content of seeds. Alternia alternata was abundant only in the initial
stages of decay. Aspergillus flavus was a dominant fungus. Macrophomina phaseolina
and Rhizoctonia solani were associated only with seeds of high moisture. The seed
mycota first increased on the seeds with storage time and subsequently decreased
significantly. Seed germination increased with storage time.

25

Chapter 1: Literature Review

1.4.6.3 Seasonal effect


Tokumasu (1998a) observed that the temperature at the surface of decaying
needle litter of Pinus densiflora was a major factor contributing to seasonal changes in
interior fungal communities. For example, Verticicladium trifidum first appeared and
Selenosporella curvispora appeared later and there was no overlap. The surface
colonizers at four different periods were roughly similar to each other and Tokumasu
(1998b) concluded that seasonal variation in climate had a stronger effect on internal
colonizers than external colonizers of needles.

1.4.6.4 Sugars
The composition of fungal communities is thought to mostly depend on
nutritional changes in the substrata. Nusbaumer and Aragno (1996) found that fresh
compost was colonized by saprobic micro-flora that used simple sugars. With time,
populations of ascomycetes and basidiomycetes appeared, first using the remaining
free sugars, then decaying the cellulose. Their action was accompanied by the
development of a population of sugar saprobes and continued until the cellulose of the
remaining internal cell walls became difficult to access because of lignin.
1.4.6.5 Tannin
Fungi will grow over a range of C:N ratios, but have optimum C:N ratios for
maximum activity (Dix and Webster, 1995). The activities of fungi colonizing aged
litter are inhibited by natural water-soluble condensed tannins (Frankland, 1998).
Tannin concentrations are crucial for succession differences. For example, Quercus is
one of the slowest trees to decay, because Quercus is rich in tannin and Quercus leaf
litter has one of the highest C:N ratios. Because Fraxinus litter contains low quantities
of tannin, the litter was rotted by fungi in a relatively short time (Frankland, 1998).
Savoie and Gourbiere (1989) also suggested that tannin could inhibit the activities of
fungi colonizing aged needle litter of Abies alba.

26

Chapter 1: Literature Review

1.5 PLAN OF THE THESIS

This thesis has four objectives:

To investigate the diversity of bambusicolous fungi in Hong Kong and


Kunming, China;

To investigate the biogeography of bambusicolous fungi in Hong Kong and


Kunming, China;

To investigate host recurrence of saprobic fungi on bamboo hosts in Hong


Kong and Kunming, China and

To investigate fungal succession on bamboo in Hong Kong.


In this chapter, the current state of knowledge of the biodiversity, taxonomy

and ecology of bambusicolous fungi after 1997 is reviewed. For information from
1997 and before, the reader should refer to Dalisay (1998) and Leung (1998). Host
specificity, host preference, and fungal succession are also reviewed in this chapter.
The new terms, host recurrence and host exclusivity, are proposed for saprobes.
Taxonomic descriptions of the bambusicolous fungi found in this study are presented
in Chapter Two. The results of biogeographical studies are presented in Chapter Three,
and in Chapter Four, host recurrence and biodiversity are addressed. Fungal succession
on bamboo is discussed in Chapter Five. In Chapter Six, a general conclusion is
provided.

27

Chapter 1: Literature Review

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Chapter 1: Literature Review

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40

Chapter 2: Taxonomy

CHAPTER 2: TAXONOMY
ABSTRACT
One hundred and twenty taxa occurring on 34 bamboo species in Hong Kong
and Kunming, Yunnan, China are described and discussed in this chapter. These
include 64 ascomycetes, one basidiomycete and 55 mitosporic taxa. Thirteen taxa,
including 2 genera, Bambicola and Striatodecospora and 1 variety are new to science.
Sixty-six fungi are new records to Hong Kong and 24 fungi are new records to Yunnan.
The new species are Annulatascus angustispora, Apiospora yunnanensis, Arthrinium
yunnanensis,

Bambicola filiforma,

Frondisphaeria

amplispora,

Linocarpon

arundinarium, L. bambusae, Ophiobolus bambusicola, Roussoella angustispora,


Spadicoides

bambusicola, Striatodecospora bambusae and Submersisphaeria

bambusicola. The new variety is Ramichloridium indicum var. microsporium. The new
combination, Ellisembia bambusicola (= Sporidesmium bambusicola) and Ellisembia
pseudoseptata (= Sporidesmium pseudoseptatum) are also made in this chapter.

Key

words:

Ascomycetes,

basidiomycetes,

coelomycetes,

mitosporic

taxa,

hyphomycetes, new species.


2.1 INTRODUCTION
The fungi identified in this study are discussed under ascomycetes,
basidiomycetes and mitosporic taxa (hyphomycetes and coelomycetes). Ascomycetes
and basidiomycetes are discussed in alphabetical order of the families, and
subsequently the genera and species. Hyphomycete and coelomycete genera are also
discussed in alphabetical order. For each genus a summary account is given and for
each species, the following information is presented: anamorph or teleomorph (if
found), known hosts and known distribution with information from the literature and
the relevant references cited, material examined with the area and country, date of
collection, collector, species code (the mycological herbarium number of The
University of Hong Kong, i.e., HKU(M)). Finally, remarks justifying and discussing
the identification are provided. For new taxa, a detailed description is given. The

41

Chapter 2: Taxonomy

etymology of the specific epithet is provided and the holotype is listed under the
" material examined". Formally the new taxa, described in this thesis are invalid until
they are described (with a Latin diagnosis) in a journal or book which is distributed to
the public, or at least botanical institutions, through sale, exchange or gift
(Hawksworth et a/., 1995).

2.2 MATERIALS AND METHODS


Dead bamboo samples ca 25-30 cm in length were randomly collected from
Hong Kong, and Kunming, Yunnan, China, and returned to the laboratory where they
were incubated in zip-lock polythene bags or boxes lined with moistened tissue.
Material was periodically examined for fungal fruiting bodies within one month.
Squash mounts and sections of fungal fruiting bodies were mounted in water for
measurement and photography. Single-spore isolations were used to obtain each taxon
in culture. During this study, more than 5,500 samples of dead bamboo were examined
from Hong Kong and Kunming following 24 field forays.
All new taxa were photographed and are illustrated in this chapter with
differential interference micrographs.
The bamboo hosts and collection regions are listed in Table 2.1. With the
exception of Phyllostachys bambusoides, all other bamboo species are confined to
either Hong Kong or Kunming. Among the 34 bamboo species collected in this study,
24 species (70.6%) occurred in Hong Kong and 10 (29.4%) species occurred in
Kunming.

42

Chapter 2: Taxonomy

Table 2.1 Bamboo hosts and collecting regions


No.
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.

Bamboo name
Arundinaria hindsii
Bambusa basihirsuta
Bambusa beecheyana
Bambusa chungii
Bambusa comigera
Bambusa glaurescens cx.fernleaf
Bambusa mutabilis
Bambusa shiuyingiana
Bambusa sp.
Bambusa spinrosa
Bambusa textilis
Bambusa tuldoides
Bambusa vulgaris var. vittata
Bambusa vulgris cv. Wamin
Dendrocalamus asper
Dendrocalamus bambusoides
Dendrocalamus brandisii
Dendrocalamus pulverulentus
Fargeisa yunnanensis
Indocalamus longiauritus
Indocalamus sinicus
Jndosasa sinica
Melocanna baccifera
Neosinocalamus affinis
Phyllostachys nidularia
Phyllostachys aurea
Phyllostachys bambusoides
Phyllostachys glauca
Phyllostachys heteroclata
Phyllostachys nidularia
Phyllostachys nigra
Phyllostachys pubescens
Schizostachyum dumetorum
Sinobambusa tootsik

Location
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Hong Kong
Kunming
Kunming
Hong Kong
Kunming
Hong Kong
Hong Kong
Kunming
Hong Kong
Kunming
Hong Kong
Kunming
Hong Kong and Kunming
Hong Kong
Kunming
Hong Kong
Kunming
Kunming
Hong Kong
Hong Kong

43

Chapter 2: Taxonomy

2.3 TAXONOMY
2.3.1 Ascomycetes
2.3.1.1 Amphisphaeriaceae
Arecophila K.D. Hyde, Nova Hedwigia 63: 81. 1996.
Hyde (1996a) established Arecophila, which is typified by A. gulubiicola K.D.
Hyde. The genus comprised 6 species and 2 combinations. Recently Dalisay et al.
(1999) described A. bambusae Dalisay and K.D. Hyde and A. coronata (Rehm)
Dalisay and K.D. Hyde as a new combination. Species in this genus are mainly found
on palms and other monocotyledonous hosts (Hyde, 1996a; Wong, 2000).
Arecophila bambusae Dalisay and K.D. Hyde, Mycoscience 40: 185. 1999.
Anamorph: Unknown. Colony slow growing on PDA, ca 2-3 cm diam. after
two weeks, white, fluffy, with sparse aerial hyphae. No sporulating structures formed
(HKUCC3391).
Known hosts: Bambusa sp. (Dalisay, Hyde and Quimio, 1999), B. chungii, B.
tuldoides (this study) and Dendrocalamus pulverulentus (Dalisay et al., 1999).
Known distribution: China (Hong Kong) and the Philippines (Dalisay et al.,
1999).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the dead culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HKU(M) 9140); on
dead culm of B. tuldoides, 23 Oct. 2000, Dequn Zhou (HKU(M) 9377); Kadoorie
Farm, on the senescent culm of Dendrocalamus publverulentus, 5 Aug. 1998, Dequn
Zhou (HKU(M) 9090); ibid. (HKU(M) 9092); Hong Kong Island, Pokfulam Country
Park, on dead culm of B. tuldoides, 5 June 1999, Dequn Zhou (HKU(M) 9244).
Remarks: The collections here are identified as A. bambusae, except that the
ascospores are slightly larger (22-24 x 6-8 jam vs. 19-22.5 x 5.5-7 um) (Dalisay et al,
1999).

44

Chapter 2: Taxonomy

2.3.1.2 Annulatascaceae
Annulatascus K.D. Hyde, Australian Systematic Botany 5: 117. 1992.
Annulatascus was introduced by Hyde (1992) for an ascomycete occurring on
wood submerged in a stream in North Queensland, Australia. It was first placed in the
Lasiosphaeriaceae, Sordariales on account of its black, immersed to superficial
ascomata, which lack a clypeus, paraphyses which are wide, septate and tapering, and
asci which are cylindrical and with an relatively massive refractive apical ring (Hyde,
1992a). The Lasiosphaeriaceae sensu lato include fungi with a large variety of
ascospore forms, peridial shapes, ascoma morphologies and anamorphs (Carroll and
Munk 1964; Barr, 1990a). Wong et al. (1998) established the Annulatascaceae to
accommodate Annulatascus and Annulatascus-like species. So far 8 species of
Annulatascus have been reported (Hyde and Wong, 2000).
Annulatascus angustispora D.Q. Zhou, K.D. Hyde and T.E. Umali, sp. nov.
(Figs 2.1.1-2.1.12)
Etymology: From the Latin prefix angusti meaning "narrow" and spora
meaning " spores" in reference to the narrow ascospores.
Pseudostromata forming beneath the host surface, visible as raised domes,
sometimes erumpent, 220-250 um diam. Ascomata globose to subglobose, two to
three in a stroma, in section 245-350 um diam. x 220-340 um high (3c = 225 um diam.
x 230 um high, n = 10) (Figs. 2.1.2-2.1.3). Peridium 15-30 um wide along side (3c =
20 um, n = 10); 12-15 (am at the base (3c = 13.5 um, n = 10), composed of 5-6 layers
of angular cells (Fig. 2.1.4). Paraphyses 4 um wide at base, tapering to 2 um wide at
the apex, hyaline, septate, filamentous (Fig. 2.1.5). Asci 158-170 x 8-10 um) (x =164
x 10 um, n = 25), 8-spored, cylindrical, relatively long-pedicellate (Figs. 2.1.6-2.1.7),
with a relatively massive apical ring, 4 um wide x 2 um high (Figs. 2.1.8-2.1.9).
Ascospores 22-24 x 5-6 um (3c =23x 6 um, n = 30), overlapping uniseriate, fusiform,
hyaline, unicellular, smooth, surrounded by thin mucilaginous sheath (Figs. 2.1.102.1.12).
Known hosts: Bambusa sp. (Dalisay, 1998), B. chungii (this study) and

45

Figs. 2.1.1-2.1.12. Interference micrographs of Annulatascus angustispora (from (HKU(M) 9225,


holotype). 1. Appearance of ascomata on bamboo culm (arrowed). 2-3. Sections of the ascomata. 4.
Section of peridium. 5. Paraphyses. 6-7. Asci. 8-9. Asci with massive refractive apical rings (arrowed). 1012. Ascospores. Scale bars: 1 = 300 um, 2-3 = 50 urn, 4-12 = 10 urn.

Chapter 2: Taxonomy

Dendrocalamus sp. (Dalisay, 1998).


Material examined: Hong Kong, New Territories, Wu Tong Village, on
senescent culm of Bambusa chungii, 18 May 1999, Dequn Zhou (HKU(M) 9225,
holotype).
Other material examined: Hong Kong, New Territories, Tai Po Kau Nature
Reserve, on senescent culm of Bambusa sp., T.E. Umali and F. Layung, TD96SH
(HKU(M) 8740); ibid, on senescent culm of Dendrocalamus sp. (HKU(M) 5652).
Remarks: Annulatascus

angustispora differs

from

other

species

in

Annulatascus by its quite narrow and unicellular ascospores (Hyde, 1992, 1995; Wong
et al., 1999; Ho et ah, 1999a, b; Hyde and Wong, 2000). Annulatascus angustispora is
comparable with A. velatisporus in having a massive apical ring and fusiform, smoothwalled ascospores. The ascospores of A. angustispora are however, much narrower
than those of A. velatispora (5-6 (im vs. 9-12 um). Annulatascus velatispora,
moreover, is an aquatic species (Hyde, 1992) while A. angustispora is terrestrial in
origin.
Submersisphaeria K.D. Hyde, Nova Hedwigia 62: 172. 1996.
Hyde (1996b) introduced Submersisphaeria, a monotypic genus, with the type
species S. aquatica K.D. Hyde, based on an ascomycete on submerged wood in a
stream on Mt Lewis, north Queensland, Australia. The morphology of the ascomata,
asci and paraphyses were similar to those of Annulatascus K.D. Hyde.
Submersisphaeria differs from Annulatascus in having brown ascospores, with polar
hyaline germ pores (Hyde, 1996b).
Submersisphaeria bambusicola D.Q. Zhou and K.D. Hyde, Fungal Diversity 4: 182.
2000.
Material examined: Hong Kong, New Territories, Tai Lam Country Park, on
dead culms of Sinobambusa tootsik, 21 June 2000, Dequn Zhou (HKU(M) 9388); ibid.,
Tai Po Kau Nature Reserve, on dead culm of Phyllostachys bambusoides, 5 Sept. 2000,
Dequn Zhou (HKU(M) 9399).
This species has been formally published (see Appendix).

46

Chapter 2: Taxonomy

2.3.1.3 Apiosporaceae
Apiospora Sacc, Atti della Societa Veneto-Trentina di Science Naturali Resendente in
Padova 4:85. 1875.
Apiospora was introduced by Saccardo with A. montagnei Sacc. as the type
species in 1875. The Apiosporaceae was established to accommodate the genera with
Arthrinium-like anamorphs (Hyde, Frohlich and Taylor, 1998). Apiospora species
occur on grasses and palms throughout the world, except in the subarctic and arctic
regions (Miiller, 1992). The genus is very common on bamboo where the anamorph is
readily distinguishable as dusty blackened regions on recently dead culms (Hyde,
Frohlich and Taylor, 1998). Teleomorphs and anamorphs often develop simultaneously
adjacent to each other. Apiospora is a small genus with 12 species, most of which are
graminicolous or cypericolous (Hyde, Frohlich and Taylor, 1998).
Apiospora montagnei Sacc, Atti della Societa Veneto-Trentina di Science Naturali
Resendente in Padova 4: 85. 1875.
Anamorph: Arthrinium arundinis (Hyde, Frohlich and Taylor, 1998). Colonies
ca 6 cm diam. on PDA after one month, grey, cottony, low convex, with edge entire
and dark grey (in reverse side) but failing to sporulate.
Known hosts: Arundo mauritanica (Hyde, Frohlich and Taylor, 1998,
Helianthus annuus (Roberts et ah, 1986) and Phyllostachys bambusoides (this study),
Phyllostachys sp. (McKenzie, 1992).
Known distribution: Algeria (Hyde, Frohlich and Taylor, 1998), China (Hong
Kong and Yunnan) (this study), New Zealand (McKenzie, 1992) and USA (Roberts et
ah, 1986).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa tuldoides, 17 June 1999, Dequn Zhou (HKU(M) 9255);
China, Yunnan, Kunming, Lunan, on the dead culm oi Phyllostachys bambusoides, 30
Sept. 1998, Dequn Zhou (HKU(M) 9190).
Remarks: The specimens are very similar to Apiospora montagnei, but the
ascospores are slightly wider (26-32 x 8-10 urn vs. 23-28 x 6-8 um) (Hyde et al.,
1998).

47

Chapter 2: Taxonomy

Apiospora sinensis K.D. Hyde, J. Frohl. and J.E. Taylor, Sydowia 50: 27. 1998.
Anamorph: Arthrinium phaeospermum (Corda) M.B. Ellis (Hyde, Frohlich and
Taylor, 1998), which often occur simultaneously alongside the teleomorph on the same
bamboo culm. Cultures on PDA white, cottony, with aerial sparse mycelia and milky
white (reverse side), but no sporulating structures produced (HKUCC 3460, 3461).
Known hosts: Arundinaria hindsii, Bambusa textilis, Neosinocalamus affinis
(Hyde, Frohlich and Taylor, 1998), Phyllostachys bambusoides (this study) and
Trachycarpus fortunei (Hyde, Frohlich and Taylor, 1998).
Known distribution: China (Hyde, Frohlich and Taylor, 1998) and Hong Kong
(this study).
Material examined: Hong Kong, Hong Kong Island, Long Fu Shan Country
Park, on the dead culm of Arundinaria hindsii, 30 July 1998, Dequn Zhou (HKU(M)
9047); ibid. (HKU(M) 9081); ibid. (HKU(M) 9085); ibid. (HKU(M) 9222); on the
dead culm of Bambusa textilis, 7 June, 1998, Dequn Zhou (HKU(M) 8340); New
Territories. Tai Po Kau Nature Reserve, on senescent culm of B. tuldoides, 19 Apr.
1999. Dequn Zhou (HKU(M) 9212); ibid., 17 June 1999, Dequn Zhou (HKU(M)
9247); ibid. (HKU(M) 9297); ibid. (HKU(M) 9301); China, Yunnan, Kunming, Anlin,
Qiu Mu Yuan, on the dead culm of Neosinocalamus affinis, 5 Oct. 1998, Dequn Zhou
(HKU(M) 9164); West Hill, on senescent culm of Fargesia yunnanensis, 4 July 1999,
Dequn Zhou (HKU(M) 9262); ibid., 29 July 1999, Dequn Zhou (HKU(M) 9310); ibid.,
(HKU(M) 9312); ibid. (HKU(M) 9319); Anlin, on senescent culm of Neosiocalamus
affinis, 20 June 1999, Dequn Zhou (HKU(M) 9258); Lunan, on the dead culm of
Phyllostachys bambusoides, 30 Sept. 1998, Dequn Zhou (HKU(M) 9180); 25 June
1999, Dequn Zhou (HKU(M) 9271); ibid. (HKU(M) 9282); Yiliang, on dead culm of
Phyllostachys bambusoides, 27 July 1999, Dequn Zhou (HKU(M) 9328).
Remarks: Hyde, Frohlich and Taylor (1998) found that Apiospora sinensis was
distinctive within the genus in possessing ascospores surrounded by a wide,
mucilaginous sheath. Ascospore size is considered an unreliable character for species
identification due to the degree of overlap in ascospore length between species. The
anamorphs are thought to have undergone a greater degree of diversification and are
important in the identification of the teleomorph (Hyde, Frohlich and Taylor, 1998).

48

Chapter 2: Taxonomy

The ascospore dimension of the collections are within the size range of those of
Apiospora sinensis, but are slightly shorter (22.5-34 x 6-8 urn vs. 26-34 x 6-8 (am)
(Hyde, Frohlich and Taylor, 1998).
Apiospora yunnanensis D.Q. Zhou and K.D. Hyde, sp. nov.

(Figs 2.2.1-2.2.9)

Etymology: In reference to the location.


Ascomata immersed or semi-immersed, visible as raised blackened areas on the
host surface, with neck and upper surface visible through splits in the host tissue,
gregarious, in linear rows; individual ascoma in section 165-365 um diam. x 75-185
um high (mean = 273.5 um diam. x 129.5 um high, n = 15), globose to subglobose,
longitudinal axis vertical to the host surface (Figs 2.2.1-2.2.2). Peridium up to 24 urn
wide, comprising elongate, angular cells with brown cells (Fig. 2.2.4). Paraphyses up
to 6 um wide, hypha-like (Fig. 2.2.3). Asci 86-128 x 24-30 urn (3c = 106 x 26 um, n =
20), 8-spored, clavate or cylindric-clavate, thin-walled, slightly flattened apically,
lacking any apical apparatus (Figs 2.2.5-2.2.6). Ascospores 30-54 x 10-12 um (3c =
41.3 x 11 urn, n = 50), biseriate, hyaline, smooth-walled, apiosporous, straight or basal
cell slightly bent, basal cell 1/4-1/5 the size of the larger apical cell (Figs 2.2.7-2.2.9).
Material examined: CHINA, Yunnan, Kunming, An Lin, Yu Long Wan, on
dead culm of Neosinocalamus affinis, 19 Aug. 1998, Dequn Zhou (HKU(M) 9100,
holotype)
Other material examined: CHINA, Yunnan, Kunming, An Lin, Yu Long Wan,
on dead culm of Neosinocalamus qffinis, 9 Aug. 1998, Dequn Zhou (HKU(M) 9093)
Remarks: Apiospora yunnanensis is distinct from other species of Apiospora
(Samuels et al, 1981; Miiller, 1992; Hyde et al, 1998) as it has very larger ascospores.
This species has affinities with A. sinensis, but differs in its larger ascospores (30-54 x
10-12 urn vs. 26-34 x 6-8.4 um) and in lacking a mucilaginous sheath (Hyde, Frohlich
and Taylor, 1998). Its anamorph is also different from Arthrinium phaeospermum
(Corda) M.B. Ellis in the latter having much smaller conidia (9-12 x 6-8 urn), mainly
rounded in face view, and mainly lenticular (Hyde, Frohlich and Taylor, 1998).

49

Figs. 2.2.1-2.2.9. Interference micrographs of Apiopsora yunnanemis (from (HKU(M)


9100, holotype). 1. Appearance of ascomata on bamboo culm. 2. Section of ascomata. 3.
Paraphyses. 4. Section of peridium. 5-6. Asci. 7-9. Ascospores. Scale bars: 1 = 200 um, 2
= 50 urn, 3-9 =10 urn.

Chapter 2: Taxonomy

Arthrinium state of Apiospora

+yunnanensis

(Figs 2.3.1-2.3.4)

Colonies developing beneath the epidermis which splits longitudinally to


expose the shiny black spore masses, 5 x 1 mm (Fig 2.3.1). Conidiophore mother cells
5-10 x 3-5 um, lageniform. Conidiophores 65 x 1-1.5 urn, erect, simple, flexuous,
cylindrical, colourless, smooth, with colourless highly refractive or pale brown
transverse septa (Figs 2.3.2-2.3.3). Conidia 4-6 um diam. in face view, 2-3 (-4) um
thick, lenticular, rather dark golden brown with a hyaline band at the junction of the
two sides (Figs 2.3.2-2.3.4).
Cultural characteristics: Colony reaching 8 cm diam. on PDA after two months.
The aerial hyphae sparse, white, flat, with brown pigmentation in media (HKUCC
3534).
Material examined: CHINA, Yunnan, Kunming, An Lin, Qiu Mu Yuan, on
dead culm of Neosinocalamus affinis, 19 Aug. 1998, Dequn Zhou (HKU (M) 9095);
on dead culm of Neosinocalamus affinis, 5 Oct. 1998, Dequn Zhou (HKU(M) 9166);
Kunming International Gardening Exposition, on senescent culm of Fargesia
yunnanensis, 28 Sept. 1998, Dequn Zhou (HKU(M) 9167); Chengong, on senescent
culm of Dendrocalamus bambusoides, 4 Oct. 1998, Dequn Zhou (HKU(M) 9184).
2.3.1.4 Diatrypaceae
Eutypella (Nitschke) Sacc, Atti della Societa Veneto-Trentina di Science Naturali
Resendente in Padova 4: 80. 1875.
Eutypella is a widespread genus comprising 76 species (Hawksworth et al.,
1995). Rappaz (1987) discussed the history and taxonomy of this genus and gave a
key to 73 species. Eutypella occurs on the dead branches of hardwood and host
specificity varies between species (Glawe and Rogers, 1984).
Eutypella is very similar in overall morphology to Eutypa. Both genera have
perithecia which are immersed in a stroma with elongate, black necks, asci which are
unitunicate and long pedicellate, and ascospores which are unicellular and allantoid
(Hanlin, 1990). The two taxa differ in stromata morphology. Eutypa has perithecia
immersed in a single layer within a white or green entostroma surrounded by a black
exostroma, while in Eutypella, perithecia are immersed in a stromata of both host and

50

Figs. 2.3.1-2.3.4. Interference micrographs of Arthrinium state of Apiospora


yunnanensis (from (HKU(M) 9095, holotype). 1. Appearance of shiny black conidia
masses on bamboo culm. 2-4. Conidiophores and conidia. Scale bars: 1 = 1000 (am, 2-4
= 10 (am.

Chapter 2: Taxonomy

fungal tissue which is delimited by a blackened zone (Rappaz, 1987). They also differ
in the staining reaction of the apical rings of their asci; weakly amyloid in Eutypa,
non-amyloid in Eutypella (Hanlin, 1990). This genus could be confused with Diatrype,
which is distinguished from Eutypella and Eutypa, in having a less well developed
stroma, and usually embedded in the substrate and formed of host tissue or a mixture
of host and fungal tissue (Glawe and Rogers, 1984).
Eutypella gliricidiae Rehm, Philipines Journal of Science 8: 189. 1913.
Anamorph: Unknown. Cultures on PDA is white, woolly and flat, with white
aerial mycelium, the underside is creamy, and the agar in centre stained yellow, slow
growth 1.5 cm diam. after one week.
Known hosts: On Bambusa chungii (this study), Erythhna indica, Gliricidia
maculatum (Rappaz, 1987), Phyllostachys nidulaha (this study) and Streblus asper
(Rappaz, 1987).
Known distribution: China (Hong Kong) (this study) and the Philippines
(Rappaz, 1987).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the dead culm of Phyllostachys nidularia, 8 Sept. 1998, Dequn Zhou (HKU(M)
9108); on the dead culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HKU(M)
9146); on dead culm of P. bambusoides, 5 Aug. 1999, Dequn Zhou (HKU(M) 9340);
Wu Tong village, on the senescent culm of B. chungii, 18 May 1999, Dequn Zhou
(HKU(M) 9224).
Remarks: The specimens are identified as Eutypella gliricidiae based on a key
in Rappaz (1987), but the ascospores are slightly larger (6-8 x 1-2 um vs. 5-7.5 x 1-2
(am) (Rappaz, 1987).
2.3.1.5 Didymosphaeriaceae
Roussoella Sacc, Atti Accademia Scientifica Veneto Trentino Istriana 6: 410. 1888.
Roussoella was established by Saccardo (Saccardo and Paoletti, 1888) for the
single species R. nitidula Sacc. and Paol. from bamboo in Malacca. Hohnel (1919)
found an earlier name in Dothidea hysterioides Ces., and combined this to represent

51

Chapter 2: Taxonomy

the type of Roussoella. Muller and von Arx (1962) accepted R. hysterioides (Ces.)
Hohn. as the correct name of the single species. They placed Roussoella in the
Amphisphaeriaceae, a position partially followed by Aptroot (1995a). Aptroot (1995b)
described asci of Roussoella as unitunicate and 'IKI negative or positive (blue)', but
later he considered the asci to be bitunicate after he examined further material. Hyde,
Eriksson and Yue (1996) and Hyde (1997a) reported Cytoplea hysterioides, to be the
anamorph of R. hysterioides and added two new species, R. aequatoriensis K.D. Hyde
and R. saltuensis K.D. Hyde. In the same paper, he provided a modified key to
Roussoella, based on the key proposed by Ju et al. (1996). Hyde (1997a) suggested the
nature of the ascus in Roussoella needed further discussion, because he examined
numerous slides from both fresh and dried material and found no evidence that the asci
discharged their spores fissitunicately.
Roussoella angustispora D.Q. Zhou and K.D. Hyde, sp. nov.

(Figs 2.4.1-2.4.8)

Etymology: From Latin angustus meaning " narrow" and spora meaning
'spore', referring to the narrow ascospores.
Ascomata forming under black, shiny, dome-like areas up to 2 mm diam. on the
host surface, with an inconspicuous central ostiole, solitary; in section 750-900 urn
diam., up to 300 um high, subglobose, surrounded by stromatic tissues (Fig 2.4.1).
Peridium up to 3 um wide, comprising several layers of compressed elongate brownwalled cells. Trabeculae embedded in a gelatinous matrix (Fig 2.4.2). Asci 200-240 x
14-17 um (x = 210 x 15.5 um, n = 20), 8-spored, cylindrical, with a knob-like pedicel,
relatively thick-walled, bitunicate, with an ocular chamber and faint ring (Figs 2.4.32.4.4). Ascospores 24-28 x 6-8 um (x = 25.5 x 7 um, n = 50), uniseriate, ellipsoidfusiform,

1-septate, constricted at the septum, brown, with reticulate wall

ornamentations (Figs 2.4.5-2.4.8).


Material examined: HONG KONG, New Territories, Tai Po Kau Nature
Reserve, on senescent culm of Bambusa changii, 8 Sept. 1998, Dequn Zhou (HKU (M)
9144, holotype);
Other material examined: BRUNEI, Tembong, Kuala Belalong, Field Studies

52

Figs. 2.4.1-2.4.8. Interference micrographs of Roussoella angustispora (from (HKU(M)


9144, holotype). 1. Appearance of ascomata on bamboo culm (arrowed). 2. Trabeculae.
3-4. Asci. 5-8. Ascospores. Scale bars: 1 = 500 urn, 2, 5-8 = 10 um, 3-4 = 20 urn.

Chapter 2: Taxonomy

Centre, on bamboo, at edge of river, 12 Dec. 1997, K.D. Hyde (HKU(M) 8313).
Remarks: Roussoella scabrispora is the only one species of the genus having
reticulate ascospores. The ascospores ornamentation in R. alveolata comprises
longitudinal ribs and numerous horizontal links between the ribs which give the pits an
angular to square appearance. Roussoella alveolata also has larger ascospores and
larger pits (Ju et al., 1996). Roussoella angustispora resembles R. scabrispora, but the
ascospores are conspicuously narrower (24-28 x 6-8 um vs. 26.5-35 x 10-12.5 um)
(Hyde, 1997a).
Roussoella hysterioides (Ces.) Hohn., Sitzungsberichten der Akademie der
Wissenschaften in Wien, Mathematischnaturwissenschaftliche Klasse, Abteilung I 128:
563. 1919.
Anamorph and cultural characters: Cytoplea hysterioides K. D. Hyde (Hyde,
Eriksson and Yue, 1996). Colonies growing up to 5 cm diam. on PDA after one month,
mycelium white to grey, cottony, underside creamy grey (HKUCC 3379).
Known distribution: China (Hong Kong and Kunming) (this study), tropical,
subtropical and warm temperate regions (Hyde, 1997a).
Known hosts: Arundinaria hindsii, Bambusa spp., B. basihirsuta, B. multabilis,
Dendrocalamus pulverulentus, Indocalamus sinicus, Phyllostachys glauca, P.
pubscens (this study), Pennisetum sp. and Zingiberaceae sp. (Hyde, 1997a).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa textilis, 14 Apr. 1998, Dequn Zhou (HKU(M) 8329); on
senescent culm of B. multabilis, 15 July 1998, Dequn Zhou (HKU(M) 9026); ibid.
(HKU (M) 9031); ibid. (HKU(M) 8330); on senescent culm of B. tuldoides, 17 June
1999, Dequn Zhou (HKU(M) 9252); ibid. (HKU(M) 9290); on senescent culm of
Phylostachys glauca, 6 June 1998, Dequn Zhou (HKU(M) 8360); ibid. (HKU(M)
8364); on dead culm of Bambusa vulgaris var. vittata, 25 June 1998, Dequn Zhou
(HKU(M) 8381); Sai Kung, on dead culm of Indocalamus sinicus, 27 June 1998,
Dequn Zhou (HKU(M) 9001); on dead culm of Dendrocalamus pulverulentus, 27 June
1998, Dequn Zhou (HKU(M) 8392); 29 July 1998, Dequn Zhou (HKU(M) 9065); on
dead culm of Arundinaria hindsii, 29 July 1998, Dequn Zhou (HKU(M) 9067); Hong

53

Chapter 2: Taxonomy

Kong Island, Victoria Peak, on dead culm of Arundinaria hindsii, 8 June 1998, Dequn
Zhou (HKU(M) 8344); on senescent culm of Bambusa basihirsuta, 30 June 1998,
Dequn Zhou (HKU(M) 9012); China, Yunnan, Kunming, West Hill, on dead culm of
Fargesia yunnanensis, 4 July 1999, Dequn Zhou (HKU(M) 9265); 29 July 1999,
Dequn Zhou (HKU(M) 9315); ibid. (HKU(M) 9317); Yiliang, on dead culm of P.
bambusoides, 25 June 1999, Dequn Zhou (HKU(M) 9268); ibid. (HKU(M) 9273); ibid.
(HKU(M) 9283); ibid. (HKU(M) 9285); ibid. (HKU(M) 9287); 27 July 1999, Dequn
Zhou (HKU(M) 9305); ibid. (HKU(M) 9324); ibid. (HKU(M) 9325); ibid. (HKU(M)
9327); ibid. (HKU(M) 9329); ibid. (HKU(M); ibid. (HKU(M) 9331); ibid. (HKU(M)
9333); ibid. (HKU(M) 9308); ibid. (HKU(M) 9309); 13 Sept. 1999, Dequn Zhou
(HKU(M) 9360); An Lin, Qiu Mu Yuan, on senescent culm of Phyllostachys pubscens,
5 Oct. 1998, Dequn Zhou (HKU(M) 9197); on dead culm of Neosinocalamus affinis, 4
July 1999, Dequn Zhou (HKU(M) 9288).
Remarks: The collections are very similar to R. hysterioides, except that the
ascospores are slightly shorter (16-27.5 x 6-8 urn vs. 18-34 x 6-8 um) (Hyde, 1997a).
The length of ascospores is quite variable (18-34 um) (Hyde, 1997a) and the shape of
the ascospores from different habitats and hosts are also variable. A molecular study is
needed to confirm if this species is heterogeneous. This species is very common on
bamboo culms in Hong Kong and Kunming, China. It seems that the fungus has a host
recurrence to monocotyledons, although sometimes this fungus also occurs on
dicotyledons (Hyde, Eriksson and Yue, 1996).
Roussoella intermedia YM. Ju, J.D. Rogers, and Huhndorf, Mycotaxon 58: 447. 1996.
Cultural characters: Colonies slow growing, ca. 5 cm diam. after one month,
white, downy, concentric rings, flat, underside smokey-grey with creamy margin
(HKUCC 3009).
Known hosts: Bambusa beecheyana, B. vulgaris var. vittata (this study),
Bambusa sp. and Dendrocalamus latiflorus (Ju, Rogers and Huhndorf, 1996).
Known distribution: Philippines, Taiwan (Ju and Rogers, 1996) and Hong
Kong (this study).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,

54

Chapter 2: Taxonomy

on senescent culm of Bambusa vulgaris var. vittata, 24 June 1998, Dequn Zhou
(HKU(M) 8382); on dead culm of Bambusa beecheyana, 15 July 1998, Dequn Zhou
(HKU(M) 9039); on senescent culm of Phyllostachys bambusoides, 5 Aug. 1999,
Dequn Zhou (HKU(M) 9341); ibid. (HKU(M) 9343); Hong Kong Island, Pokfulam
Country Park, on senescent culm of B. tuldoides, 5 June 1999, Dequn Zhou (HKU(M)
9245).
Remarks: Ju and Rogers (1996) pointed out that Roussoella intermedia has an
ascospore size range between that of R. hysterioides and R. pustulans. The collections
are similar to R. intermedia based on description and illustrations by Ju et al. (1996),
but the ascospores are longer (17.5-20 x 4-5 um vs. 13.5-18 x 4.5-5.5um) (Ju and
Rogers. 1996).
Roussoella pustulans (Ellis and Everh.) Y.M. Ju, J.D. Rogers and Huhndorf,
Mycotaxon 58: 448. 1996.
Anamorph: Unknown. Colony 6.5 cm diam. after two weeks on PDA, blackish
brown, raised and flutty, with rich aerial hyphae. No fruiting structures produced after
one month.
Known hosts: Bambusa sp., B. mutalilis, B. textilis, B. sinospinosa,
Dendrocalamus pulverulentus, Neosinocalamus affinis, Phyllostachys glauca, P.
nidularia (this study), palms and unidentified bamboo (Hyde, 1997a).
Known distribution: China (Kunming) (this study), Brunei, Indonesia, Taiwan
and USA (Hyde, 1997a).
Material examined: China, Yunnan, Kunming, An Lin, Qiu Mu Yuan, on
senescent culm of Neosinocalamus affinis, 5 Oct. 1998, Dequn Zhou (HKU(M) 9163);
West Hill, on dead culm of Fargesia yunnanensis, 4 July 1999, Dequn Zhou (HKU(M)
9266); 29 July 1999, Dequn Zhou (HKU(M) 9313); Yiliang, on senescent culm of
Phyllostachys bambusoides, 25 June 1999, Dequn Zhou (HKU(M) 9270); ibid. (HKU
(M) 9189); Hong Kong, New Territories, Tai Po Kau Nature Reserve, on dead culm of
P. glauca, 6 June, 1998, Dequn Zhou (HKU(M) 8359); on senescent culm of P.
bambusoides, 5 Aug. 1999, Dequn Zhou (HKU(M) 9344); on dead culm of Bambusa
textilis, 24 June 1998, Dequn Zhou (HKU(M) 8386); on dead culm of Bambusa sp., 15

55

Chapter 2: Taxonomy

July 1998, Dequn Zhou (HKU(M) 9022); on senescent culm of B. mutabilis, 15 July,
1998, Dequn Zhou (HKU(M) 9025); Sai Kung, on senescent culm of B. sinospinosa, 2
July, 1998, Dequn Zhou (HKU(M) 8399); on senescent culm of Dendwcalamus
pulverulentus, 27 June 1998, Dequn Zhou (HKU(M) 8393); ibid. (HKU(M) 9008); on
senescent culm of B. sinospinosa, 2 July 1998, Dequn Zhou (HKU(M) 8399); Dao
Moushan, on dead culm of P. nidularia, 19 May 1999, Dequn Zhou (HKU(M) 9238).
Remarks: This species is easily confused with Didymosphaeria futilis (Berk,
and Broome) Rehm. The main differences between these two species are that in
Didymosphaeria, pseudoparaphyses are trabeculae, richly anastomosing above the asci,
and ascospores are thinly pseudosepta (Aptroot, 1995b). Didymosphaeria futilis also
has wider ascospores (7-13 x 4-6 um vs. 9.5-13 x 3.5-4.5 um) (Aptroot, 1995b; Hyde,
1997a). The specimens are identified as R. pustulans except that the ascospores are
shorter and wider (8-10 x 4-5 um vs. 11-14 x 2.5-3 um) (Hyde, 1997a). This species
repeatedly is identified on bamboo as very common on many bamboo hosts in Hong
Kong and Kunming.

Didymosphaeria Fuckel, Jabrbuch des nassauischen Vereins fur Naturkunde 23-24:


140. 1870.
The genus Didymosphaeria was erected by Fuckel (1870) for three species of
ascomycetes with two-celled spores. It soon became a popular genus for
didymosporous pyrenocarpus ascomycetes (Aptroot, 1995b). Several attempts have
been made to redefine the genus, notably by Saccardo (1882) who restricted the genus
to brown-spored members and Scheinpflug (1958) who excluded several species, but
investigated only 20% of the species then known. About 100 taxa have already been
excluded from the genus (Hawksworth and Boise, 1985; Hawksworth, 1985a, 1985b;
Hawksworth and Diederich, 1988; Barr, 1989a, 1989b, 1990a, 1990b, 1992a, 1992b,
1993). Aptroot (1995b) accepted 7 species in his world monograph.
Didymosphaeria futilis (Berkeley and Broome) Rehm, Hedwigia 18: 167. 1879.
Anamorph: A PhomaAike coelomycete and Fusicladiella (Aptroot, 1995b).
Known hosts: Cosmopolitan in and on stems of many plants, also on dead

56

Chapter 2: Taxonomy

leaves, wood and even linoleum (Aptroot, 1995b), on senescent culm of Phyllostachys
bambusoides (this study) and Trachycarpus fortunei (Frohlich, 1997).
Known distribution: China (Hong Kong and Hunan) (Frohlich, 1997, this study)
and cosmopolitian (Aptroot, 1995b).
Material examined: Hong, Kong, The New Territories, Tai Po Kau Nature
Reserve, on senescent culm of Phyllostachys bambusoides, 5 Aug. 1999, Dequn Zhou
(HKU(M) 9336); ibid. (HKU(M) 9338); ibid. (HKU(M) 9342); on Bambusa tuldoides,
23 Oct. 1999, Dequn Zhou (HKU(M) 9380).
Remarks: Aptroot (1995b) pointed out that the species is very uniform and very
little variation exists, except in the degree of blackening of the stromatic tissues. The
collections are very similar to Didymosphaeria futilis, except that the ascospores are
slightly longer (10-14 x 4-5 urn vs. 7-13 x 4-6 um) (Aptroot, 1995b).

Didymosphaeria massarioides Sacc. and Brunaud, Michelia 2: 592. 1882.


Anamorph: Phoma-like coelomycete (Aptroot, 1995).
Known hosts: Dryas octopetala (Rosaceae), Fraxinus excelsior (Oleaceae), Ilex
aquifolium (Aquifoliaceae), Lycium barbarum (Solanaceae) (Aptroot, 1995b),
Phyllostachys bambusoides (this study), Rubus idaeus, R. sanctus (Rosaceae) and
Sambucus nigra (Caprifoliaceae) (Aptroot, 1995b).
Known distribution: China (Hong Kong) (this study), Cosmopolitan, in and on
stems and roots of various plants (Aptroot, 1995b).
Material examined: Hong Kong, The New Territories, Tai Po Kau Nature
Reserve, on dead culm of Phyllostachys bambusoides, 5 Aug. 1999, Dequn Zhou
(HKU(M) 9345).
Remarks: The collection mostly agrees with Didymosphaeria massarioides
except that the ascospores are narrower (16-18 x 4-6 urn vs. 15-22 x 6-9 urn) (Aptroot,
1995b).

57

Chapter 2: Taxonomy

2.3.1.6 Hyponectriaceae
Oxydothis Penzig and Sacc, Malpighia 11: 505. 1897.
Oxydothis was described from Cibodas, Java, Indonesia by Penzig and
Saccardo (1897) and reviewed by Hyde (1994a, b, c), who suggested that the taxon
should be moved from the Amphisphaeriaceae to the Hyponectriaceae on the basis of
ascus, ascospore and peridium structure. The genus is most similar to Ceriospora
Niessl, Frondispora (Ces.) K.D. Hyde, Lasiobertia Sivan. and Leiosphaerella Hohn.
These four genera were discussed and compared with Oxydothis by Hyde (1994a).
Oxydothis was characterized by having long cylindrical asci with a J+ subapical
apparatus and long fusiform to filiform hyaline bicelled ascospores, which taper from
the center to spine-like, pointed or rounded processes (Hyde, 1994b; Hyde, Taylor and
Frohlich, 2000).
Frohlich (1997) suggested that Oxydothis was the most common saprobic genus
encountered on palms. Prior to Frohlich, Hyde (1994a) pointed out that most
Oxydothis species are from palm hosts, Pandanus and bamboo. Frohlich (1997)
concluded that it was not surprising that these hosts shared some species of saprobic
fungi, as the three plant hosts often share tropical habitats.
Oxydothis aequalis Syd. and P. Syd., Annales Mycologici 15: 208. 1917.
Known hosts: On bamboo (Hyde, 1994a), Calamus moti, C. radicalis, C.
sordicus (Frohlich, 1997), on senescent culm of Bambusa chungii (this study).
Known distribution: Australia, Brunei (Frohlich, 1997), China (Hong Kong)
(this study), Malaysia and Philippines (Hyde, 1994a).
Material examined: Hong Kong, New Territories, Wu Tong Village, on
senescent culm of Bambusa chungii, 18 May 1999, Dequn Zhou (HKU(M) 9228).
Remarks: This specimen resembles Oxydothis aequalis. The ascospores and
ascal rings have a similar size to Oxydothis aequalis (Table 2.2).

58

Chapter 2: Taxonomy

Table 2.2 Comparison of Oxydothis aequalis collections with the type


Taxa
HKU (M) JF94
(Frohlich. 1997)
HKU(M) JF356
(FrOhlich. 1997)
Dequn Zhou
(HKU (M)) 9228
O. aequalis
(Hyde, 1994a)

Ascus length
(urn)
180-270

Ascus
Width (nm)
10-12(-14)

Ascospores
length (nm)
70-90

Ascospores
width (|jm)
(4.6-)5.2-6.4

Ascal ring
height (|jm)
4.6-6.8

Ascal ring
diam. (^m)
2.4-3.2

180-200

10.4-12

74-90

5.6-6

2.8-4

2.2-2.6

200-245

10-12.5

70-78(-82)

6(-8)

5-6

3-4

100

12

74-90

4.6.5

3.5-5

2.4-3.2

Asci of the specimens in this study are also much longer than those of O.
aequalis (Table 2.2). Ascus dimensions given by Hyde (1994a) were based on a
"single measurement as few intact asci were seen". The measurement of asci in my
specimen are quite close to those of Frohlich's specimens (HKU(M) JF 94, HKU(M)
IF 356). Ascomata dimensions of O. aequalis are not given by Hyde (1994a), but the
other description and illustration are consistent with my collection. The ascospores and
apical ring shape of O. aequalis also agree with my specimen. Hyde (1994a) suggested
that O. aequalis is similar to O. grisea, from which it differs in having clustered
ascomata and narrow ascospores and in its host, a bamboo.
Oxydothis oraniopsis J. Frohl. and K.D. Hyde, Mycological Research 98: 215. 1994.
Known hosts: Living leaf of Oraniopsis appendiculata, Laccospadix
australasicus (Hyde, 1994a) and on dead culm of Bambusa tuldoides (this study).
Known distribution: Australia (Hyde, 1994a) and China (Hong Kong and
Kunming) (this study).
Material examined: Hong Kong, The New Territories, Tai Po Kau Nature
Reserve, on senescent culm of Bambusa tuldoides, 17 June 1999, Dequn Zhou
(HKU(M) 9253); ibid. (HKU(M) 9298); China, Yunnan, Kunming, Yiliang, on dead
culm of B. bambusoides, 25 June 1999, Dequn Zhou (HKU(M) 9280).
Remarks: The specimens are identified as Oxydothis oraniopsis, because they
are consistent with the description and illustrations by Hyde (1994a), but the habitats
are different: Oxydothis oraniopsis was found on living leaves (Hyde, 1994a) whereas
my collections are on dead bamboo culms.

59

Chapter 2: Taxonomy

Linocarpon Syd. and P. Syd., Annales Mycologici 15:210. 1917.


Linocarpon was introduced by Sydow and Sydow (1917) with the type species
L pandani (Syd. and P. Syd.) Syd. and P. Syd., a fungus colonising dead Pandanus
leaves in the Philippines. Several species were later described and Petrak (1952)
transferred to Linocarpon a number of species previously placed in Ophiobolus Riess
and reduced Gaeumannomyces von Arx and Olivier to synonymy. Walker (1980)
could not accept Petrak's concept of Linocarpon since it incorporated several
characteristics not shown by the type species of Linocarpon. Hyde (1988) described a
new species of Linocarpon, L. appendiculatum K.D. Hyde, from Nypa fruticans and
transferred three Ophiobolus species to Linocarpon. Walker (1980) limited
Linocarpon to mostly palmicolous species. He described and illustrated the type
species, L. pandani, and provided notes on other Linocarpon species. Hyde (1997b)
also pointed out that Linocarpon is predominantly saprotrophic on palms, although
hosts include Cajanus (Fabaceae), Miscanthus (Poaceae), Pandanus (Pandanaceae),
Phenakospermum (Zingiberaceae) and Quercus (Fagaceae). Frbhlich (1997) deduced
that palms, Pandanus and bamboo should share some species of saprobic fungi,
because the basic morphology of bamboo stems (culms) is very similar to that of rattan
stems and Pandanus fronds. The three plants often share tropical habitats. Our finding
supports Frohlich's inference and has extended the host range. The salient features of
Linocarpon are listed by Hyde (1997b):
Lenticular ascomata immersed beneath a clypeus, usually shiny and black;
Cylindrical unitunicate asci with rounded apices and a J-, ring-like, apical
apparatus;
Filiform ascospores with refringent bands. Their apices may or may not contain
mucilage;
Tapering paraphyses and
A n unusual Phialophora, anamorph.
In 1998 four more Linocarpon species were added by Dulymamode, Cannon
and Anderson (1998).

60

Chapter 2: Taxonomy

Linocarpon arundinarium D.Q. Zhou and K.D. Hyde, sp. nov.

(Figs 2.5.1-2.5.12)

Etymology: referring to host.


Ascomata developing beneath the clypeus, which is composed of epidermal
cells packed with fungal hyphae, lenticular in section, with variable amounts of
stromatic tissue laterally; in section425-440 umdiam., and 120-155 urn high, brown
to black and with a central periphysate ostiolum (Figs 2.5.1-2.5.3). Paraphyses hyphalike, wide septate and tapering (Fig 2.5.7). Asci 120-137.5 x 10-12.5 urn, 8-spored,
cylindrical, shortly pedicellate, with rounded apex and ring-like apical apparatus (Figs
2.5.8-2.5.10). Ascospores 85-115 x 2.5 um (3c = 95.5 x 2.5 urn, n = 50), filiform,
multiseriate, apex rounded, with abasal appendage (Figs 2.5.4-2.5.6, 2.5.11-2.5.12).
Material examined: HONG KONG, Hong Kong Island, Lung Fu Shan Country
Park, on dead culm of Arundinaria hindsii, 5 June, 1998, Dequn Zhou (HKU(M) 8338,
holotype).
Other material examined: Hong Kong, Hong Kong Island, Lung Fu Shan
Country Park, on dead culm of Arundinaria hindsii, 5 June, 1998, Dequn Zhou
(HKU(M) 9407).
Remarks: This species is most similar to L. livistonae (Henn.) K.D. Hyde, but L.
livistonae has larger ascomata (700 urn diam. and 195 um high vs. 425-440 um diam.
and 120-155 um high) and ascospores are narrower (70-140 x 1.6-2.3 um vs. 85-115 x
2.5 um) (Hyde, 1988). In most cases Linocarpon livistonae occurs on palms and
Pandanus, whereas L. arundinariae is from Arundinaria hindsii, a bamboo host.

Linocarpon bambusae D.Q. Zhou and K.D. Hyde, sp. nov.

(Figs 2.6.1 -2.6.11)

Etymology: Referring to bamboo host.


Ascomata appearing on host culm, shiny, raised, visible as dome-shaped, single
or sometimes gregarious in groups of two, developing below the clypeus (Fig 2.6.1),
in vertical section 415-460 um diam. (x = 437.5 um, n = 10), 145-155 um high (x =
150 um, n = 10). Clypeus 40-65 um, of epidermal cells packed with fungal tissues
(Fig 2.6.2). Peridium up to ca. 10 um wide, comprising a few layers of brown-walled
elongate cells (Fig. 2.6.3). Paraphyses 4-5 um wide, septate, tapering, embedded in

61

Figs. 2.5.1-2.5.12. Interference micrographs of Linocarpon arundinarium (from (HKU(M) 8338,


hoiotype). 1. Appearance of ascomata on bamboo culm. 2. Section of ascoma. 3. Section of stroma. 4-6.
Ascospores with 2-3 septa (arrowed). 7. Paraphyses (arrowed). 8-9. Asci. 10. Ascus with a subapical
ring (arrowed). 11-12. Ascospores with a fringe-like appendage (arrowed). Scale bars: 1 = 500 urn, 2 =
200 urn, 3-6, 7,11-12 = 10nm. 8-10 = 5 \ua.

Figs. 2.6.1-2.6.11. Interference micrographs of Linocarpon bambusae (from (HKU(M)


9223, holotype). 1. Appearance of ascoma on bamboo culm. 2. Section of ascoma
showing stromatal structure 3. Section of peridium. 4. Paraphyses. 5-6. Ascospores
showing a fringe-like appendage (arrowed). 7-8. Asci. 9-11. Ascospores. Scale bars: 1 =
500 urn, 2 = 100 urn, 4 = 20 urn, 3, 5-11 = 10 urn

Chapter 2: Taxonomy

gelatinous matrix (Fig 2.6.4). Asci (118-) 124-134 x (10-) 12-14 um (3c = 127 x 12.5
(am, n = 20), cylindric-clavate, pedicellate, rounded at the apex, lacking an apical ringlike apparatus (Figs 2.6.7-2.6.8). Ascospores 98-108 (-110) x 3.5-4 um (3c = 103 x 3.5
um, n = 30), filiform, rounded at the apex, with fringe-like mucilage at the base (Figs
2.6.5-2.6.6,2.6.9-2.6.11).
Material examined: HONG KONG, Hong Kong Island, Long Fushan Country
Park, on senescent culm of Arundinaria hindsii, 18 May 1999, Dequn Zhou (HKU(M)
9223, holotype); 5 June 1999, Dequn Zhou (HKU(M) 9242); The New Territories, Tai
Po Kau Nature Reserve, on senescent culm of Arundinaria hindsii, 6 June 1998,
Dequn Zhou (HKU(M) 8348).
Remarks: Linocarpon bambusae differs from L. sulcatum Dulymamode, P.F.
Cannon and Peerally in its larger ascomata (415-460 um wide vs. 260-340 um wide;
145-155 um high vs. 70-130 um high) (Dulymamode et ah, 1998) and as L. sulcatum
lacks a clypeus. Linocarpon sulcatum occurs on dead attached leaves of Pandanus
barklyi (Dulymamode et ai, 1998), but L. bambusae occurs on bamboo culms.
Linocarpon bambusae cannot be confused with Linocarpon carinisporum K.D. Hyde,
or L. elaeidis Petr. Ascospores of I. bambusae are larger than those of L. carinisporum
K.D. Hyde (98-108 x 3.5-4 um vs. 84-98 x 2.4-3.2 um). Linocarpon bambusae is
distinct from L. elaeidis in having longer ascospores (98-108 x 3.5-4 um vs. 72-97 x
3-4 um um). Both L. carinisporium and L. elaeidis occurred on woody plants (Hyde,
1992b; 1997b).
Linocarpon livistonae (Henn.) K.D. Hyde, Transactions of Mycological Society of
Japan 29: 346. 1988.
Anamorph: Phialophora sp. (Hyde, 1997b).
Known distribution: Australia, Brazil, French Guiana, Indonesia, Philippines,
Taiwan (Hyde, 1992b, 1997b) and Hong Kong (this study).
Known hosts: Numerous palm hosts and Pandanus spp. (Hyde, 1992b),
Archontophoenix (Taylor, 1998) and Arundinaria hindsii (this study).
Material examined: Hong Kong, Hong Kong Island, Lung Fu Shan Country

62

Chapter 2: Taxonomy

Park, on dead culm of Arundinaria hindsii, 19 July 1998, Dequn Zhou (HKU(M)
9046); No. 1 Pokfulam Reservoir, on senescent culm of Melocanna baccifera, 30 June,
1998, Dequn Zhou (HKU(M) 9051).
Remarks: Linocarpon pandani Rehm is almost identical to L. livistonae and is
considered synonymous, but Hyde (1992b) maintained L. pandani as an independent
species. Minor differences include the slightly smaller, but overlapping ascal ring and
smaller fruiting bodies. Linocarpon livistonae is most similar to L. elaeidis, but differs
in having narrow ascospores (1.6-2.3 urn vs. 3-4 urn) (Hyde, 1992b). The collection
closely resembles L. livistonae, but the ascomata are smaller (370-430 um in diam. vs.
585-780 um in diam.), asci are smaller (112.5-125 x 10-12.5 um vs. 170-216 x 8-12
(am) and ascospores are smaller (82.5-107.5 x 1.5-2.5 um vs. 124-140 x 1.6-2.3 (am)
(Hyde, 1992b).

Linocarpon pandani (Syd. and P. Syd.) Syd. and P. Syd., Annales Mycologici 15:210.
1917.
Known hosts: On Pandanus laevis, Miscanthus sp. (Hyde, 1992b),
Phyllostachys nidularia (this study).
Known distribution: Philippines, Hong Kong (this study) and Taiwan (Hyde,
1992b).
Material examined: Hong Kong, New Territories, Dao Mou Shan, on senescent
culm oi Phyllostachys nidularia, 19 May 1999, Dequn Zhou (HKU(M) 9236); Tai Po
Kau Nature Reserve, on dead culm oi Phyllostachys bambusoides, 8 July 1999, Dequn
Zhou (HKU(M) 9274).
Remarks: This collection is identified as L. pandani based on the description
and illustrations given by Hyde (1992b), however, the ascospores in this collection are
slightly longer (78-86 x 3-4 (am vs. 62-80 x 2-4 (im) (Hyde, 1992). Linocarpon
pandani is different from L. pandanicola in the ascospores lacking appendages.
All species of Linocarpon found on bamboo in this thesis are compared in
Table 2.3.

63

Chapter 2: Taxonomy

Table 2.3 Comparison of Linocarpon species from bamboo in this thesis


Taxa

Ascomata (um)

Asci (um)

Ascospores (urn)

L. arundinarium
(this thesis)
L.bambusae
(this thesis)
L.livistonae
(Hyde, 1992b)
L. pandani
(Hyde, 1992b)

425-440 diam. x 120155 high


415-460 diam. x 145155 high
700 diam x 195 high

120-137.5x1012.5
124-134x12-14
100-140x6-12

85-115x2.5
(x= 95.5x2.5)
98-108x3.5-4
(x= 103x3.5)
70-104x1.6-2.3

600-650 diam. x 200300 high

100-140x8-10

62-80x2-4

Appendages
of ascospores
With a base
appendage
With fringelike mucilage
With a base
mucilage
No
appendages

Key to Linocarpon species occurring on bamboo in this study


1 .Ascospores lacking appendages, ascospores 62-80 x 2-4 um

L. pandani

Ascospores having appendages

2 Ascospores 3.5-4 um wide, with fringe-like mucilage

L. bambusae

Ascospores less than 3 urn wide

3. Ascomata 700 um diam x 195 um high, ascospores 85-115 x 1.6-2.3 um


L. livistonae
Ascomata 425-440 um diam x 120-155 um high, ascospores 85-115x2.5 um
L. arundinarium
2.3.1.7 Lasiosphaeriaceae
Ophioceras Sacc, Sylloge Fungorum 2: 358-360. 1883.
Twenty-five species have been included in Ophioceras Sacc. (Miiller and
Dennis, 1965; Tilak and Kale, 1969; Conway and Barr, 1977; Lai, 1987; Teng, 1996,),
but five are currently accepted (Shearer, Crane and Chen, 1999). The genus is in need
of revision, but unfortunately, most of the descriptions are not detailed or clear
(Walker, 1980; Teng, 1996). Ophioceras species have black, immersed or superficial,
globose perithecia with elongate necks, tapering paraphyses, unitunicate, cylindrical
asci with refractive, non-amyloid subapical rings and filiform, multiseptate ascospores
which may be hyaline or slightly tinted. The ascomata of Ophioceras have distinctive
beaks, which are easily identified with comparison to other lasiosphaeriaceous genera
with filiform ascospores. Ophioceras is similar to Pseudohalonectria, but differs in the
pigmentation and firmness of the ascomata, and the method of ascospore discharge.
Pseudohalonectria species have bright yellow, membraneous ascomata, while those of

64

Chapter 2: Taxonomy

Ophioceras species are dark brown to black (Hyde, Taylor and Frohlich, 2000).
Ophioceras is accommodated in the Lasiosphaeriaceae (Conway and Barr, 1977;
Hawksworth et al., 1995). This placement was supported with preliminary molecular
studies (Chen, Shearer and Klopp, 1995), but recently Shearer, Crane and Chen (1999)
discussed placement of Ophioceras and provisionally placed Ophioceras in the
Magnaporthaceae. Hyde et al. (2000) also agreed with this placement. The anamorphs
are presently unknown (Shearer et al, 1999).
Ophioceras sorghi Saccas, L'Agronomie Tropicale 9: 285. 1954.
Known hosts: Calanus radicalis, Phyllostachys bambusoides (this study),
Licuala ramsayi and Sorghum vulgare (Taylor, 1998).
Known distribution: Australia, Central African Republic (Taylor, 1998) and
China (Kunming) (this study).
Material examined: China, Yunnan, Kunming, Yiliang, on senescent culm of
Phyllostachys bambusoides, 27 June 1999, Dequn Zhou (HKU(M) 9367); Hong Kong,
The New Territories, Tai Shui Kang, on dead culm of Phyllostachys bambusoides, 29
July, 1998, Dequn Zhou (HKU(M) 9070).
Remarks: The specimens are identified as Ophioceras sorghi (Walker, 1980),
but ascospores are slightly shorter (74-84 um vs.75-95 um).

2.3.1.8 Leptosphaeriaceae
Ophiobolus Riess, Nova Hedwigia 1: 27. 1854.
Ophiobolus (Leptosphaeriaceae) was introduced by Riess (1854). The generic
concept is relatively broad (Holm, 1948; Muller, 1952). Holm (1957) however,
accepted a much narrower concept of the genus (two species, with provisional
inclusion of a third) and transferred others to Nodulosphaeria and Leptospora. The
narrow concept was followed by Luttrell (1973), von Arx and Muller (1975) and
Walker (1980). Walker (1980) described and discussed Ophiobolus on the basis of the
type specimen, which includes 63 taxa, but most were transferred to other genera.
Shoemaker (1976) chose to use Ophiobolus in a broad sense. He reported on 31
species with detailed descriptions for each species, and provided a key. The main

65

Chapter 2: Taxonomy

differences between Ophiobolus and other similar genera was also discussed. About
100 species have now been reported and these are widespread (Hawksworth et al.,
1995). Some species of Ophiobolus are plant pathogens (Shoemaker, 1976).

Ophiobolus bambusicola D.Q. Zhou and K.D. Hyde, sp. nov.

(Figs 2.7.1-2.7.6)

Etymology: referring to the host.


Ascomata 270-330 urn diam. x 340-390 urn high, scattered, immersed
becoming erumpent, globose, papillate, glabrous (Fig 2.7.1). Pseudoparaphyses ca. 24 um wide, sparse, cellular, tapering distally (Fig 2.7.2). Asci 114-138 x (8-) 10-12 urn
(x = 128 x 10 um , n = 15), 8-spored, cylindrical, pedicel, bitunicate, apex rounded
(Figs 2.7.3, 2.7.5). Ascospores 88-102 x 2-2.5 um (x = 93.4 x 4 um, n = 30),
scolecosporous, 5-8 transversely septate, straight to slightly curved, hyaline, smooth,
not constricted at some septa, no mucilaginous sheath, terminal mucilaginous
appendages or characteristic nodose swellings (Figs 2.7.4,2.7.6).
Material examined: China, Yunnan, Kunming, Chengong, on senescent culm
of Phyllostachys bambusoides, 4 Oct. 1998, Dequn Zhou (HKU(M) 9189, holotype).
Other material examined: China, Yunnan, Kunming, Chengong, on senescent
culm of Phyllostachys bambusoides, 4 Oct. 1998, Dequn Zhou (HKU(M) 9381).
Remarks: Ophiobolus bambusicola resembles O. acuminatus (Sow.) Duby, but
differs as O. acuminatus has disarticulating ascospores with more septa (Khashnobish
and Shearer, 1995). Ophiobolus bambusicola is distinguished from O. fulgidus
(Clinton and Peck) Sacc, which has shorter, wider ascospores with obviously enlarged
cells or constrictions (Shoemaker, 1976). Ophiobolus bambusicola is also distinct
from O. galii Richon which has a greater number of septa, and narrower ascospores
(Shoemaker, 1976). Ophiobolus bambusicola cannot be confused with O. miscanthi
Sivan. and Hsieh, as ascospores are wider and pale brown to brown (Sivanesan and
Hsieh, 1989). Ophiobolus bambusicola is also different from O. troakei Hawksw. and
Sivan., which has narrower asci and longer ascospores with a greater number of septa
(Hawksworth and Sivanesan, 1975). These species are compared in Table 2.4.

66

Figs. 2.7.1-2.7.6. Interference micrographs of Ophiobolus bambusicola (from (HKU(M)


9189, holotype). 1. Appearance of ascomata on bamboo culms (arrowed). 2.
Pseudoparaphyses. 3, 5. Asci. 4, 6. Ascospores. Scale bars: 1 = 500 um, 2-6 = 10 um.

Chapter 2: Taxonomy

Table 2.4 Comparison of Ophiobolus bambusicola with other similar species


Ascus (urn)

Ascospore (u.m)

Number septa
on ascospore

O. acuminatas
(Sow.) Duby
O. bambusicola

118-133 x 10-13

64-100x2.2-3

8-15

Ascospore
disarticulating or
not
disarticulating

114-138 x 10-12

88-102x2-2.5

5-8

not disarticulating

O. fulgidus (Clinton

90-115 x 12-15

60-95 x 3.0-3.5

not disarticulating

O. galii Richon

100-130x8-10

85-130 x 1.5-2.0

11-16

not disarticulating

O. miscanthi Sivan.

112-165 x 12-13

70-90 x 3.0-4.5

6-11

not disarticulating

120-140x8-9

115-138x2.0-2.5

7-14

not disarticulating

Scientific name

and peck) Sacc.

and Hsieh
O. troakei Hawksw.

2.3.1.9 Lophiostomataceae
Massarina Sacc, Sylloge Fungorum 12: 153. 1883.
Massarina was introduced by Saccardo (1883) with M. eburnea (Tul. and C.
Tul.) Sacc. as the type species. Aptroot (1998) monographed Massarina and accepted
43 species. Three species, M. arundinariae (Ellis & Everh.) M.E. Barr on Arundinaria
sp., M. carolinensis Kohlmeyer on unidentified bamboo and M. bambusina Teng on
bamboo were accepted by Aptroot (1998), but he did not see the type material of M.
bambusina. Dalisay (1998) found M. arundinariae on Bambusa sp. in the Philippines
and M. desmonci (Syd. and P. Syd.) K.D. Hyde and Aptroot on Bambusa sp. in Hong
Kong.
Massarina arundinariae (Ellis and Everhart) M. Barr, Mycotaxon 45: 211. 1992.
Known hosts: On Arundinaria sp. (Aptroot, 1998), Bambusa sp. (Dalisay, 1998)
and B. glaucescens cv. Fernleaf (this study).
Known distribution: China (Hong Kong) (Dalisay, 1998), Philippines (Dalisay,
1998) and USA (Aptroot, 1998).
Material examined: Hong Kong, New Territories, Wu Tong Village, on
senescent culm of Bambusa glaucescens cv. Fernleaf, 18 May 1999, Dequn Zhou
(HKU(M) 9230).
Remarks: Aptroot (1998) and Barr (1992b) regarded Didymosphaeria

67

Chapter 2: Taxonomy

arundinariae Ellis and Everh. a synonym of Massarina arundinariae. My collection is


similar to Massarina arundinariae (Aptroot, 1998), but the ascospores are slightly
narrower (5-6 urn vs. 6-7 urn) and close to the measurement given by Dalisay (1998)
(24-26 x 5-6 um vs. 23-25 x 5.5-6 jun).

Massarina desmonci (Syd. and P. Syd.) K.D. Hyde and Aptroot, Nova Hedwigia 64:
493.1997.
Cultural characters: Colonies slow growing, up to ca. 4 cm after one month,
green, granular, with granular structures in centre, underside milky grey (HKUCC
3128).
Known

hosts: Desmoncus sp.

(Aptroot,

1998)

and Dendrocalamus

pulverulentus (this study).


Known distribution: Brazil (Hyde and Aptroot, 1997) and China (Hong Kong)
(this study).
Material examined: Hong Kong, New Territories, Sai Kung, on dead culm of
Dendrocalamus pulverulentus, 26 June 1999, Dequn Zhou (HKU(M) 9054).
Remarks: This collection is similar to Massarina desmonci (Hyde and Aptroot,
1997), however, ascospores are slightly narrower (11.5-14 um vs. 12-15 um).
Massarina

hepaticarum

(Crouan)

Dobbeler,

Mitteilungen

Botanischen

Staatssammlung Miinchen 14: 197. 1978.


Known hosts: Phyllostacys glauca (this study) and Reboulia hemisphaerica
(Aptroot, 1998).
Known distribution: China (Hong Kong) (this study) and France (Aptroot,
1998).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Phyllostacys glauca, 6 June 1998, Dequn Zhou (HKU(M) 8362).
Remarks: This collection is similar to Massarina hepaticarum (Aptroot, 1998),
but the ascospores are slightly longer (20-25 urn vs. 16-23 urn).

68

Chapter 2: Taxonomy

Massarina cf. immersa Dobbeler, Mitteilungen Botanischen

Staatssammlung

MQnchen 14: 198. 1978.


Known hosts: Phyllostachys bambusoides (this study) and Polytrichum
juniperinum (Aptroot, 1998).
Known distribution: China (Hong Kong) (this study) and Switzerland (Aptroot,
1998).
Material examined: Hong Kong, The New Territories, Tai Po Kau Nature
Reserve, on senescent culm oi Phyllostachys bambusoides, 8 July 1999, Dequn Zhou
(HKU(M) 9277); on dead culm of Bambusa tuldoides, 17 June 1999, Dequn Zhou
(HKU(M) 9295).
Remarks: Aptroot (1998) found that the ascomata opened on both sides of moss
leaves, with more ascomata opening on the dorsal than on the ventral side. Although
similar to M. immersa, the identification is doubtful, as M. immersa was found on
moss leaves (Aptroot, 1998).
Massarina palmicola K.D. Hyde and Aptroot, Nova Hedwigia 64: 499. 1997.
Cultural characters: Colonies on PDA slow growing, ca. 5 cm diam. after one
month, grey, cottony, underside dark brown, agar stained brown (HKUCC 8363).
Known hosts: Livistona (Hyde and Aptroot, 1997) and Phyllostachys glauca
(this study).
Known distribution: Hong Kong (this study) and Malaysia (Hyde and Aptroot,
1997).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Schizostachyum dumetorum, 6 June 1998, Dequn Zhou (HKU(M)
8363).
Remarks: The ascospores of this collection are larger (47-57.5 x 10-15 um vs.
37-53 x 10-12.5 um) and the shape of ascospores is inequilaterally fusiform with one
side flattened (Aptroot, 1998).

69

Chapter 2: Taxonomy

Massarina talae Speg., Anales Mus, Nac. Hist. Nat. Buenos Aires, Ser. 2, 6: 278.
1899.
Known hosts: Celtis talae (Aptroot, 1998), on dead culms of Arundinaha
hindsii and Schizostachyum dumetorum (this study).
Known distribution: Argentina (Aptroot, 1998) and Hong Kong (this study).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Schizostachyum dumetorum, 8 Sept. 1998, Dequn Zhou (HKU(M)
9105); ibid., on the dead culm of Arundinaria hindsii, 6 June 1998, Dequn Zhou
(HKU(M) 8349).
Remarks: The collections are similar to Massarina talae Aptroot (1998), but the
ascospores are slightly longer (38-54 urn vs. 43-50 urn).

2.3.1.10 Melanommataceae
Astrosphaeriella Syd. and P. Syd. Annales Mycologici 11: 260. 1913.
Hyde and Frohlich (1997) monographed the genus and accepted 31 species
(including 10 new species and 5 new combinations Javaria. Boise (1984) is similar
and it was treated as congeneric.
Astrosphaeriella cf. bakeriana (Sacc.) K.D. Hyde and J. Frohl., Sydowia 50: 93. 1997.
Known hosts: Calamus, Livistona (Hyde and Frohlich, 1997), Bambusa chungii,
Dendrocalamus puherulentus and Phyllostachys pubscens (this study).
Known distribution: China (Hong Kong) (Hyde and Frohlich, 1997), China
(Kunming) (this study), Japan, Papua New Guinea and Singapore (Hyde and Frohlich,
1997).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa sp., 15 July 1998, Dequn Zhou (HKU(M) 9021); on
senescent culm of Bambusa chungii, 18 May 1999, Dequn Zhou (HKU(M) 9223); on
dead culm of B. glaurescens CV Fernleaf, 18 May 1999, Dequn Zhou (HKU(M) 9233);
Tai Shi King, on dead culm of Dendrocalamus puherulentus, on 29 July 1998, Dequn
Zhou (HKU(M) 9063); China, Yunnan, Kunming, An Lin, Qiu Mu Yuan, on senescent
culm of Phyllostachys pubscens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9158).

70

Chapter 2: Taxonomy

Remarks: This species is most similar to A. floridana, which may be a synonym


(Hyde and Frohlich, 1997). In introducing A. floridana, Barr (1990a) compares A.
aosinemis, and concludes that this species has shorter and wider asci. In the
illustration (and description) of A. floridana provided by Barr (1990a), however,
ascospores lack a mucilaginous sheath, while those of A. bakeriana have a distinct
mucilaginous sheath. Hyde and Frohlich (1997) maintained A. floridana and believed
that further collections may prove it to be a synonym. The specimens match
Astrosphaeriella bakeriana, which was redescribed by Hyde and Frohlich (1997).
Astrosphaeriella bakeriana is an endophytes of Livistona chinensis (Guo et ah, 1998),
therefore we tentatively place these collections as A. bakeriana.

Astrosphaeriella fissuristoma J. Frohl., K.D. Hyde and Aptroot, Nova Hedwigia 70:
147.2000.
Known hosts: Bambusa shiuyingiana (this study), Calamus australis, C.
caryotoides, C. conirostris, C. flabellatus, C. moti, C. pogonacanthus, C. radicadis, C.
sordidus, Lucuala sp. and Mauritia flexuosa (Hyde, Taylor and Frohlch, 2000).
Known distribution: Australia, Brunei (Hyde, Aptroot, Frohlch and Taylor.,
2000) and China (Hong Kong) (this study).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa shiuyingiana, 27 Aug. 1999, Dequn Zhou (HKU(M)
9352); ibid., on dead culm of B. tuldoides, 23 Oct. 1999, Dequn Zhou (HKU(M) 9352).
Remarks: This collection is identical to A. fissuristoma based on the description
by (Hyde, Aptroot, Taylor and Frohlch, 2000).
Astrosphaeriella maculans (Rehm) Aptroot and K.D. Hyde, Nova Hedwigia 70: 152.
2000.
Known hosts: Archontophoenix alexandrae, Arenga mindorensis

(Hyde,

Aptroot, Taylor and Frohlch, 2000) and Bambusa chungii (this study).
Known distribution: Australia, China (Hong Kong) (this study) and the
Philippines (Hyde, Aptroot, Taylor and Frohlch, 2000).
Material examined: Hong Kong, New Territories, Kadoorie Farm, on dead

71

Chapter 2: Taxonomy

culm of Bambusa chungii, 10 Aug. 1998, Dequn Zhou (HKU(M) 9091), Tai Po Kau
Nature Reserve, on senescent culm of B. chungii, 26 June 1998, Dequn Zhou (HKU(M)
8385).
Remarks: This specimen is identical to Astrosphaeriella maculans according to
description and illustrations given by Hyde, Aptroot, Taylor and Frohlch (2000).
Astrosphaeriella splendida K.D. Hyde and J. Frohl., Sydowia 50: 81-132. 1997.
Known hosts: Arundinaria hindsii (this study), Astrocaryum, Iriartia, Jessenia
and Mauritia (Hyde and Frohlich, 1997).
Known distribution: China (Hong Kong) (this study) and Ecuador (Hyde and
Frohlich, 1997).
Material examined: Hong Kong, Hong Kong Island, Lung Fu Shan Country
Park, on senescent culm of Arundinaria hindsii, 5 June 1999, Dequn Zhou (HKU(M)
9240).
Remarks: Hyde and Frohlich (1997) pointed out that this species differs from
other Astrosphaeriella species as its ascospores have unusual appendages at each end.
The collection has affinities with Astrosphaeriella splendida in morphology of
ascomata, asci and ascospores, but ascospores are slightly narrower (48-54 x 6-8 um
vs. 42-63 x 7.5-10 um) (Hyde and Frohlich, 1997).

Astrosphaeriella stellata (Pat.) Sacc, Sylloge Fungorum 24: 938. 1928.


Cultural characters: Colonies growing up to 4 cm diam. after one month, white,
dusty, underside creamy white at margin with cottony mycelia in centre (HKUCC
3150).
Known hosts: Bambusa textilis (this study), Bamboo, Calamus (Hyde and
Frohlich, 1997) and Dendrocalamus pulverulentus (this study).
Known distribution: Australia and China (Hong Kong and mainland) (Hyde and
Frohlich, 1997 and this study), French Guiana, India, Indonesia (Java), Japan, Papua
New Guinea, Philippines and Vietnam (Hyde and Frohlich, 1997).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa textilis, 6 June 1998, Dequn Zhou (HKU(M) 8351); on

72

Chapter 2: Taxonomy

senescent culm of B. shiuyingiana, 27 Aug. 1999, Dequn Zhou (HKU(M) 9362); 3


Sept. 1999, Dequn Zhou (HKU(M) 9206); on dead culm of B. tuldoides., 23 Oct. 1999,
Dequn Zhou (HKU(M) 9379); Tai Shui Kang, on senescent culm of Dendrocalamus
pulverulentus, 29 July 1998, Dequn Zhou (HKU(M) 9064); Kadoorie Farm, on
senescent culm of D. pulverulentus, 5 Aug. 1998, Dequn Zhou (HKU(M) 9089).
Remarks: The collections are identical to Astrosphaerieall stellata (Hyde and
Frohlich, 1997). This species is a very common species on bamboo in Hong Kong.
Perhaps this species has host recurrence on bamboo.
Astrosphaeriella trochus (Penz. and Sacc.) D. Hawksw., Botanical Journal of the
Linnean Society 82: 46. 1981.
Known hosts: Old bamboo stems and stout grasses (Hyde and Frohlich, 1997)
and Phyllostachyspubscens (this study).
Known distribution: Chile (Hyde and Frohlich, 1997), China (Kunming) (this
study), Colombia, Ecuador, French Guiana, Indonesia (Java), Japan, South Africa,
Taiwan and Uganda (Hyde and Frohlich, 1997).
Material examined: China, Yunnan, Kunming, An Lin, Qiu Mu Yuan, on
senescent culm of Phyllostachys pubscens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9154);
Yiliang, on dead culm of P. Bambusoides, 27 July 1999, Dequn Zhou (HKU(M) 9350);
13 Sept. 1999, Dequn Zhou (HKU(M) 9359).
Remarks: The collections match Astrosphaeriella trochus however, ascomata
have no teeth-like flanger (Hawksworth, 1981) and the colour of the ascospores is pale
olivaceous-brown (vs. reddish-brown) (Hyde and Frohlich, 1997).

2.3.1.11 Myxotrichaceae
Myxotrichum Kunze, Mykologische Hefte 2: 109 (1823)
According to Schroeter (1893) the first described species of this genus is
Myxotrichum chartarum Kunze. Saccardo (1886) placed the genus in the
hyphomycetes and it gradually became a deposit for very heterogenous elements.
Apinis (1964) discussed Myxotrichum and provided a key to 8 species. Hawksworth et
al. (1995) presumed there were 7 species in Myxotrichum. Eriksson and Yue (1998)

73

Chapter 2: Taxonomy

reported Myxotrichum simile Berk, and M. A. Curtis from decaying Arundinaria sp. in
USA.

Myxotrichum berkeleyi Apinis, Mycological Papers 96: 17. 1964.


Anamorph: Geotrichum-\\ke species have been found on the type specimen, but
it was not certain whether these are related to Myxotrichum berkekeyi (Apinis, 1964).
Known hosts: Bambusa textilis (this study) and Pinus silvestris (Apinis, 1964).
Known distribution: China (Hong Kong) (this study) and UK (Apinis, 1964).
Material examined: Hong Kong, The New Territories, Tai Po Kau Nature
Reserve, on senescent culm of Bambusa textilis, 3 June 1998, Dequn Zhou (HKU(M)
9050).
Remarks: The collection is almost consistent with Myxotrichum berkekeyi but
ascospores are slightly larger (3-5 x 2-3um vs. 3-4.5 x 1.5-2.5 urn) (Apinis, 1964).
2.3.1.12 Thyridiaceae
Thrydium Nitschke, Pyrenomycetes Germanici: 110. 1867.
The type species, T. vestitum (Fr.) Fuckel, occurs on twigs of various deciduous
trees in temperate regions (Eriksson and Yue, 1989). Eriksson and Yue (1989)
established the Thyridiaceae based on two genera, Thyridium and Sinosphaeria, but
the latter was later considered as a synonym of Thyridium. There are 36 species in this
genus (Hawksworth et ah, 1995).
Thyridium chrysomallum (Berk, and Broome) O.E. Eriksson and J.Z. Yue, Systema
Ascomycedum 8: 12. 1989.
Known hosts: Archontophoenix alexandrae (Taylor and Hyde, 1997), on an old
culm of bamboo and palms (Yue and Eriksson, 1987) and on senescent culm of
Bambusa tuldoides, Dendrocalamus pulverulentus, Phyllostachys bambusoides and P.
pubscens, (this study).
Known distribution: China (Hainan) (Yue and Eriksson, 1987), China (Hong
Kong and Kunming) (this study), Sri Lanka (Yue and Eriksson, 1987; Taylor and Hyde,
1997).

74

Chapter 2: Taxonomy

Material examined: Yunnan, Kunming, An Lin, Qiu Mu Yuan, on senescent


culm of Phyllostachys pubscens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9160, holotype);
Chengong, on dead culm of Phyllostachys bambusoides, 4 Oct. 1998, Dequn Zhou
(HKU(M) 9188); Hong Kong, New Territories, Sai Kung, Tu Kwa Pin, on senescent
culm of Dendrocalamus pulverulentus, 27 June 1998, Dequn Zhou (HKU(M) 9060);
on senescent culm of Bambusa tuldoides, 23 Oct. 1999, Dequn Zhou (HKU(M) 9374);
ibid. (HKU(M) 9375); Sai Kung, on senescent culm of Dendrocalamus pulverulentus,
26 June 1998, Dequn Zhou (HKU(M) 9055 ).
Remarks: The golden yellow-coloured stromata of these collections are very
conspicuous, however, the ascospores are slightly shorter than those of Thyridium
chrysomallum (12-14 vs. 14-16.5 urn) (Yue and Eriksson, 1987). The specimens
collected by Taylor lacked a stroma, and ascomata were mainly individual, but the
species was thought to be variable (Taylor and Hyde, 1997).

2.3.1.13 Valsaceae
Endothia Fries. Summa Vegetabilium Scandinaviae: 385. 1849.
Endothia Fr. species are distributed in many temperate, subtropical and tropical
areas of the world (Roane, Griffin and Elkins, 1986). Barr (1978) separated the species
into Endothia and Cryphonectria and placed these genera in the Gnomoniaceae and
Valsaceae, respectively. In the Gnomoniaceae, perithecia are upright with central,
rarely eccentric, erumpent necks; and ascospores are unicellular to one-septate or
occasionally several-septate. In the Valsaceae, the perithecia are oblique or horizontal
with oblique or lateral beaks separately erumpent or converging through a stromatic
disk. Key characters for the tribe Endotheae in the family Gnomoniaceae, to which
Endothia sensu Barr (1978) has been assigned, include few to many perithecia in welldeveloped stromata with pseudoparenchymatous stromal tissue around the perithecia.
Endothia gyrosa (Schw.) Fr., Summa Begetabilium Scandinaviae: 385. 1849.
Anamorph: Endothiella gyrosa Sacc. (Martha et. ai, 1986).
Known Hosts: Acer saccharinum, Castanea dentata, Castanea spp. (Martha et
ai, 1986), Dendrocalamus pulverulentus (this study), Fagus grandifolia, F. sylvatica,

75

Chapter 2: Taxonomy

Ilex opaca, Liquidambar formosana, L. styraciflua (Martha et ai,

1986),

Phyllostachys glauca f. yunzhu (this study), Prunus laurocerasus, Quercus agrifolia Q.


bicolor, 0. coccinea, Q. falcata, Q. georgiana, Q. ilicifolia, Q. imbrcaria, Q. lyrata, Q.
macrocarpa, Q. marilandica, Q. Montana and Q. velutina (Martha et ai, 1986).
Known Distribution: China (Hong Kong) (this study), Italy, Poland, Portugal,
Spain and USA (Martha et ai, 1986).
Material examined: Tai Po Kau Nature Reserve, New Territory, Hong Kong, on
culms of Phyllostachys glauca f. yunzhu, 6 June 1998, Dequn Zhou (HKU(M) 8353);
ibid. (HKU(M) 8366); Sai Kung, To Kwa Pin, on the dead culm of Dendrocalamus
pulverulentus. 27 June 1998, Dequn Zhou (HKU(M) 9005).
Remarks: The collections have affinities with Endothia gyrosa (Martha et. al.,
1986) except that the asci are slightly longer (25-39 urn vs. 25-30 um) and the
ascospores are narrower (1.3-2.5 um vs. 2-3 um).

2.3.1.14 Xylariaceae
Anthostomella Sacc, Atti della Societa Veneto-Trentina di Science Naturali
Resendente in Padova4: 77. 1875.
Lu and Hyde (2000) has monographed the genus Anthostomella and accepted
85 species. Anthostomella species are characterized by immersed ascomata, with a
central ostiole and a periphysate ostiolar canal. Asci are 8-spored, cylindrical,
unitunicate, apically rounded with a subapical J+ or J- ring. Ascospores are unicellular,
brown, often surrounded by a mucilaginous sheath and often with a longitudinal germ
slit (Lu and Hyde, 2000).
Anthostomella bromeliaceae Rehm., Annales Mycologici 5: 525. 1907.
Known hosts: Andropogon, Bromeliaceae (Lu and Hyde, 2000) and
Sinobambusa tootsik (this study).
Known distribution: Brazil, China (Hong Kong) and USA (Lu and Hyde, 2000).
Material examined: Hong Kong, New Territories, Tai Lam Country Park, on
the dead culm of Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9119).
Remarks: The ascospores of this collection are broader (6-7 um vs. 4-5 um)

76

Chapter 2: Taxonomy

(Lu and Hyde, 2000).

Anthostomella bruneiensis K.D. Hyde, Nova Hedwigia 62: 286. 1996.


Known hosts: Licuala sp. (Hyde, 1996c) and Bambusa tuldoides (this study).
Known distribution: Brunei (Hyde, 1996c) and Hong Kong (this study).
Material examined: Hong Kong, Hong Kong Island, Pokfulam Country Park,
on senescent culm of Bambusa tuldoides, 5 June 1999, Dequn Zhou (HKU(M) 9248);
New Territories, Tai Lam Country Park, on dead culm of Sinobambusa tootsik, 8 Sept.
1998, Dequn Zhou (HKU(M) 9119); China, Yunnan, Kunming, Little White Dragon,
on dead culm of Phyllostachys bambusoides, 1 Sept. 2000, Dequn Zhou (HKU(M)
9393).
Remarks: In these collections, ascospores are not as obviously beaked as those
in Anthostomella bruneiensis, the dimensions of asci and ascospores falls within the
size range of those in Anthostomella bruneiensis (Lu and Hyde, 2000).
Anthostomella caffrariae B.S. Lu and K.D. Hyde, Mycological Research 104: 751.
2000.
Known hosts: Bambusa chungii (this study) and Phoenix reclinata (Lu and
Hyde 2000).
Known distribution: China (Hong Kong) (this study) and South Africa (Lu and
Hyde, 2000).
Material examined: Hong Kong, New Territories, Wu Tong Village, on the
senescent culm of Bambusa chungii, 18 May 1999, Dequn Zhou (HKU(M) 9226).
Remarks: The ascospores of this collection are slightly narrower than those of
Anthostomella caffrariae in face view (8-10 um vs. 9-11.5 (am) (Lu and Hyde, 2000).
Anthostomella contaminans (Dur. and Mont.) Sacc, Michelia 2: 313. 1881.
Cultural characters: Colonies ca 7 cm diam. on PDA after two months, slow
growing, brown, greyish in the margin area, flat, with sparse aerial hyphae.
Known hosts: Bumbusa chungii (this study), Chamaerops, Phoenix, Raphia,
Smilax (Lu and Hyde, 2000) and Sinobambusa tootsik (this study).
Known distribution: Algeria, Nigeria (Lu and Hyde, 2000) and China (Hong

77

Chapter 2: Taxonomy

Kong and Kunming) (Dalisay, 1998 and this study).


Material examined: Hong Kong, Hong Kong Island, Long Fushan Country
Park, on senescent culm of Arundinaria hindsii, 11 May 1999, Dequn Zhou (HKU(M)
9220); New Territories, Tai Po Kau Nature Reserve, on the dead culm of Bambusa
chungii, 8 Sept. 1998, Dequn Zhou (HKU(M) 9135); on the dead culm of B.
shiuyingiana, 27 Aug. 1999, Dequn Zhou (HKU(M) 9363); on the dead culm of B.
tuldoides, 9 Mar. 1999, Dequn Zhou (HKU(M) 9205); 17 June 1999, Dequn Zhou
(HKU(M) 9250); ibid. (HKU(M) 9291); on senescent culm of Phyllostachys
bambusoides., 8 July 1999, Dequn Zhou (HKU(M) 9275; Yuan Leung, on the dead
culm of Sinobambusa tootsik, 18 June 1998, Dequn Zhou (HKU(M) 8374); China,
Yunnan, Kunming, West Hill, on senescent culm of Fargesia yunnanensis, 4 July 1999,
Dequn Zhou (HKU(M) 9261); ibid. (HKU(M) 9286); 29 July 1999, Dequn Zhou
(HKU(M) 9321; ibid. (HKU(M) 9323); 27 July 1999, Dequn Zhou (HKU(M) 9326);
on dead culm of Phyllostachys bambusoides., 25 June 1999, Dequn Zhou (HKU(M)
9269); Little White Dragon, on dead culm of P. bambusoides, 1 Sept. 2000, Dequn
Zhou (HKU(M) 9293); on dead culm of P. bambusoides, 1 Sept. 2000, Dequn Zhou
(HKU(M) 9394).
Remarks: These collections are identified as Anthostomella contaminas (Hyde,
1996c), but the subapical ring is higher (3-4 urn diam. and 1-1.5 um high vs 2.5-4 urn
diam. and 0.5-1 um high). Anthostomella comtaminans is very common on senescent
or dead bamboo culms from Hong Kong and Yunnan.
Anthostomella flagellariae (Rehm) B.S. Lu and K.D. Hyde, Nova Hedwigia 70: 159.
2000.
Cultural characters: Colonies very slow growing, ca. 3 cm diam. after two
months, white, felty, underside brown in centre and becoming creamy towards margin
and mycelia being sparse (HKUCC 3123).
Known hosts: Arundinaria hindsii (this study), Bambusa basihirsuta, B. chungii,
B. vulgaris var. vittata (this study), Bambusa sp., Cocos sp. (Lu and Hyde, 2000),
Dendrocalamus asper, D. pulverulentus and Phyllostachys bambusoides (this study).
Known distribution: China (Hong Kong and Guangdong), Kenya, the

78

Chapter 2: Taxonomy

Philippines and USA (Lu and Hyde, 2000).


Material examined: Hong Kong, Hong Kong Island, Victoria Cape, on the dead
culm of Arundinaria hindsii, 7 Sept. 1998, Dequn Zhou (HKU(M) 9202); No.l
Pokfulam Reservoir, on the dead culm of Bambusa chungii, 30 June 1998, Dequn
Zhou (HKU(M) 9015); Victoria Peak, on dead culm of B. basihirsuta, 30 June 1998,
Dequn Zhou (HKU(M) 9016); New Territories, Tai Po Kau Nature Reserve, on the
dead culm of B. tuldoides, 17 June 1999, Dequn Zhou (HKU(M) 9251); on the dead
culm of B. vulgaris var. vittata, 23 June 1998, Dequn Zhou (HKU(M) 8379); Tai Shui
Kang, on the dead culm of Dendrocalamus pulverulentus, 29 July 1998, Dequn Zhou
(HKU(M) 9061); Wu Tong Village, on the senescent culm of B. chungii, 18 May 1999,
Dequn Zhou (HKU(M) 9227); Yuan Leung, on the dead culm of D. asper, 18 June
1998, Dequn Zhou (HKU(M) 8368); on the dead culm of Phyllostachys bambusoides,
29 July 1998, Dequn Zhou (HKU(M) 9072); Tai Po Kau Nature Reserve, on senescent
culm of P. bambusoides, 17 June 1999, Dequn Zhou (HKU(M) 9299); 8 July 1999,
Dequn Zhou (HKU(M) 9276); 5 Aug. 1999, Dequn Zhou (HKU(M) 9339); China,
Yunnan, Kunming, West Hill, on dead culm of Fargesia yunnanensis, 29 July 1999,
Dequn Zhou (HKU(M) 9316).
Remarks: These collections have affinities with Anthostomella flagellariae
based on description and illustrations by Lu and Hyde (2000). The ascospores are
slightly longer (18-22 vs. 14.5-19.5um) (Lu and Hyde, 2000). Anthostomella
flagellariae is very common on senescent or dead bamboo culms from Hong Kong
and Yunnan and seems recurrent on bamboo.
Anthostomella longa B.S. Lu, T.U. Dalisay and K.D. Hyde, Cryptogamie Mycologie
(2000, in press)
Known hosts: on senescent culm of Bambusa sp. (Dalisay, 1998) and
Arundinaria hindsii (this study).
Known distribution: China (Hong Kong) (this study) and Philippines (Dalisay,
1998).
Material examined: Hong Kong, Hong Kong Island, Long Fushan Country
Park, on the dead culm of Arundinaria hindsii, 11 May 1999, Dequn Zhou (HKU(M)

79

Chapter 2: Taxonomy

9221).
Remarks: The collection is similar to Anthostomella longa, but differs in having
slightly broader ascospores (6-7 x 4-5.5 um vs. 5.5-6.5 x 4-5 um) (Lu, Hyde and Liew,
2000).
Anthostomella irregularispora K.D. Hyde, Nova Hedwigia 62: 303. 1996.
Known hosts: Licuala spinosa (Hyde, 1996c) and Phyllostachys nidularia (this
study).
Known distribution: China (Hong Kong and Yunnan) (this study) and Papua
New Guinea (Lu and Hyde, 2000).
Material examined: Hong Kong, New Territories, Dao Mao Shan, on senescent
culm of Phyllostachys nidularia, 19 May 1999, Dequn Zhou (HKU(M) 9237); China,
Yunnan, Kunming, on dead culm of Phyllostachys bambusoides, 27 July 1999, Dequn
Zhou (HKU(M) 9349).
Remarks: The specimens are identified as Anthostomella irregularispora, but
the ascospores are slightly shorter (16-20 vs. 16-21.5p.rn) (Lu and Hyde, 2000). The
species differs from other Anthostomella species in its irregularly shaped ascospores
with a mucilaginous sheath (Lu and Hyde, 2000).
Anthostomella raphiae B.S. Lu and K.D. Hyde, Mycological Research 104: 751. 2000.
Known hosts: Phyllostachys pubscens (this study), Labiatae sp. and Raphia
australis (Lu et ai, 2000).
Known distribution: China (Kunming) (this study) and South Africa (Lu et ai,
2000).
Material examined: China, Yunnan, Kunming, An Lin, Qiu Mu Yuan, on the
dead culm of Phyllostachys pubscens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9161).
Remarks: This collection is similar to A. raphiae (ascospores 14-18 x 6-8 um
vs. 11.5-19 x 6.5-7.5 um) and this is the first record on monocotyledonous plants.

80

Chapter 2: Taxonomy

Anthostomella rehmii (Thiim.) Sacc., Bericht des Naturhistorischen Vereins in


Augsburg 26: 95. 1881.
Known hosts: Abies, bamboo (Lu and Hyde, 2000), Bambusae tuldoides (this
study), Cladium, Freycinetia, Pandanus, Phyllostachys and Sasa (Lu and Hyde, 2000).
Known distribution: Australia, Brunei, China (Hong Kong), Germany, Japan,
Romania (Lu and Hyde, 2000).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the dead culm of Bambusa tuldoides, Sept. 1999, Dequn Zhou (HKU(M) 9354);
ibid. (HKU(M) 9355).
Remarks: These collections are most similar to A. rehmii, but ascospores are
slightly narrower (6-7 um vs. 7.5-9 um) and subapical ring are higher (4 um vs. 2-2.5
Um) (Lu and Hyde, 2000).
Anthostomella sepelibilis (Berk, and M.A. Curtis) Sacc, Sylloge Fungorum 1: 281.
1882.
Known hosts: Bambusa vulgaris var. vittata (this study), Oncidium, Smilax and
unidentified bamboo (Lu and Hyde, 2000).
Known distribution: Argentina, Brazil, China (Hong Kong) and USA (Lu and
Hyde, 2000).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the dead culm of Bambusa vulgaris var. vittata, 24 June 1998, Dequn Zhou
(HKU(M) 8377).
Remarks: This collection is similar to A. sepelibilis, but ascospores are slightly
longer (14-18 um vs. 12-16 um) (Lu and Hyde, 2000).
Anthostomella uniseriata J. Frohl. and K.D. Hyde, Palm Microfungi: 295. 2000.
Known hosts: Bambusa vulgaris var. vittata (this study) and Calamus (Lu and
Hyde, 2000).
Known distribution: Brunei (Frohlich and Hyde, 2000) and China (Lu and
Hyde, 2000).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,

81

Chapter 2: Taxonomy

on the dead culm of Bambusa vulgaris var. vittata, 24 June, 1998, Dequn Zhou
(HKU(M)9113).
Remarks: The subapical ring of the ascus in this collection is larger than A.
imiseriata (4-5 urn diam., 1 urn high vs. 1.5-2 um diam., 0.5-1 urn high), but the
collections are conspecific (Frohlich and Hyde, 2000).
Astrocystis Berk, and Broome, Botanical Journal of the Linnean Society 14: 123. 1873.
Astrocystis is based on A. mirabilis Berk, and Broome, a bamboo-inhabiting
pyrenomycete (Berkley and Broome, 1873) and confined to monocotyledonous hosts
(Frohlich and Hyde, 2000). Lasssoe and Spooner (1993) considered that the most
striking differences between Astrocystis and Rosellinia De Not. are the differences in
ascus apparatus morphology and in stromatal shape. In Rosellinia asci have a
comparatively long, tapered stipe and a large barrel-shaped apical apparatus. In
addition. Rosellinia has different anamorphs, a mainly superficial stromata developing
beneath a false subiculum (a mat of hyphae which is equivalent to an ectostroma) and
in the substrate which is not limited to monocotyledonous plants as in Astrocystis.
Typical species of Astrocystis also have a carbonized basal ring or margin around the
ascomata (Laessoe and Spooner, 1994). Ju and Rogers (1990, 1996) employed a broad
concept and treated the genus as a synonym of Rosellinia. Smith, Whalley and Hyde
(2001, in press) maintained Astrocystis as a separate entity from Rosellinia. Fifteen
species have been recorded in the genus (Hawksworth et al., 1995).
Astrocystis cocoes Laessoe and Spooner, Kew Bulletin 49: 27. 1993.
Cultural characters: Colonies slow growing up to ca. 3 cm diam. after one
month, white, fluffy, effuse, with white radical hyphal veins, underside creamy
(HKUCC 3444, 3446).
Known hosts: Arundinaria hindsii (this study), Arenga, Calamus Cocos
(Laessoe and Spooner, 1993) and Sinobambusa tootsik (this study).
Known distribution: Australia (Laessoe and Spooner, 1994), China (Hong Kong)
(this study), Brazil (Lasssoe and Spooner, 1994), India, Philippines and Virgin Islands
(Laessoe and Spooner, 1994).

82

Chapter 2: Taxonomy

Material examined: Hong Kong, Hong Kong Island, Victoria Peak, on


senescent culm of Arundinaria hindsii, 17 Sept. 1998, Dequn Zhou (HKU(M) 9149);
New Territories, Tai Lam Country Park, on dead culm of Sinobambusa tootsik, 8 Sept.
1998, Dequn Zhou (HKU(M) 9125); Tai Po Kau Nature Reserve, on the senescent
culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HKU(M) 9132); on the dead
culm of Phyllostachys bambusoides, 5 Aug. 1999, Dequn Zhou (HKU(M) 9332); ibid.
(HKU(M) 9337); Yuan Leung, on the senescent culm of Sinobambusa tootsik, 18 June
1998, Dequn Zhou (HKU(M) 8271).
Remarks: Astrocystis cocoes is well characterized by its spore shape
(inequilaterally fusiform with one or both ends beaked) and host (monocotyledonous
plants) (Laessoe and Spooner, 1994). These collections are similar to A. cocoes except
that the ascospores are slightly wider (13-14

6-7 um vs. 13-14.5 * 5-6 urn) (Laessoe

and Spooner, 1994).

Astrocystis mirabilis Berk, and Broome, Botanical Journal of the Linnean Society 14:
123. 1873.
Anamorph and cultural characters: Acanthodochium Samuels, J.D. Rogers, and
Nagasawa (Ju and Rogers 1990). The colonies on PDA growing up to 3 cm diam. after
one month, white, creamy, velvety, flat, underside creamy yellow, with rhizoid-like
mycelia, underside creamy in two weeks, no fruiting structures produced (HKUCC
3447, 3576).
Known hosts: Bambusa chungii (this study), Bambusa vulgaris (Lasssoe and
Spooner 1993; Eriksson and and Yue, 1998), Dendrocalamus latiflorus (Ju and Rogers,
1990), Leleba mutiplex, Phyllostachys bambusoides (Hino, 1961), Phyllostachys
pubscens (this study), Pleioblastus linearis, P. simoni, P. vaginatus, Sasa borealis var.
purpurascens, Semiarundianaria fastuosa and Sinobambusa tootsik (Hino, 1961).
Known distribution: China (Kunming) (this study), Indonesia, Jamaica, Japan,
Mauritius, New Caledonia, Philippines, Sri Lanka and Taiwan, (Laessoe and Spooner
1994).
Material examined: China, Yunnan, Kunming, Hot Spring, Qiu Muyuan, on
dead culm of Phyllostachys pubscens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9198); ibid.

83

Chapter 2: Taxonomy

(HKU(M) 9156); ibid. (HKU(M) 9153); Hong Kong, New Territories, Tai Po Kou
Nature Reserve, on dead culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou
(HKU(M)9134).
Remarks: These collections are very similar to Astrocystis mirabilis (Laessoe
and Spooner, 1994) however, the ascospores are slightly longer (14-16 um vs. 11.5-14
urn).
Astrocystispalmarum Laessoe and Spooner, Kew Bulletin 49: 28. 1993.
Cultural characters: Colonies slow growing, ca. 5 cm after 6 weeks, white,
velvety, radiating, underside creamy (HKUCC 3124).
Known hosts: Bambusa sp. (this study) and palms (Lasssoe and Spooner, 1994).
Known distribution: China (Hong Kong) (this study) and Bermuda (Laessoe and
Spooner, 1994).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the senescent culm of Bambusa sp., 15 July 1998, Dequn Zhou (HKU(M) 9024);
Yunnan, Kunming, on senescent culm of Phyllostachys bambusoides, 27 July 1999,
Dequn Zhou (HKU(M) 9348).
Remarks: Astrocystis palmarum has a similar habit to A. sublimbata, but differs
in spore size and host (Laessoe and Spooner, 1994). These collections are identified as
Astrocystis palmarum (Laessoe and Spooner, 1994), but ascospores are slightly
narrower (5-7.5 um vs. 6.5-7.5um).
Hypoxylon Bull., Histoire des Champignons de la France 1: 168. 1791.
Hypoxylon was monographed by Ju and Rogers (1996) who considered that
Hypoxylon reaches the highest diversity in the tropics and subtropics and can live as an
endophyte, pathogen, and/or saprobe. The genus has previously been defined many
times. Pouzar (1985a, 1985b) believed that at least some of the taxa with black
stromata have little relationship with Hypoxylon and moved them to Nemania and this
view is widely accepted (Ju and Rogers 1996). Rogers (1994) separated Camillea
Hypoxylon and Biscogniauxia following electron microscopic and cultural studies.
The taxon has been divided into two sections, Hypoxylon and Annulata,

84

Chapter 2: Taxonomy

according to characters of the stroma, the position of the ostiole, and the
ornamentation of the perispore (Ju and Rogers, 1996). Hypoxylon differs from similar
genera in the Xylariaceae in its anamorph and the structure of its stromata. Nemania
and Ustulina have Geniculosporium-like anamorphs, Biscogniauxia and Camilea have
bipartite stromata whereas Hypoxylon has unipartite stromata. In Daldinia, stromata
has concentric rings and Rhopalostroma has stipitate stroma, while the stromata of
Hypoxylon species are usually broader than the tall (Ju and Rogers 1996). The
stromata in species of Hypoxylon are brightly pigmented (Ju and Rogers 1996).
Hypoxylon aucklandiae Y.M. Ju and J.D. Rogers, A Revision of the Genus Hypoxylon:
90. 1996.
Cultural characters: Colonies growing up to ca. 5 cm after two months, brown,
cottony, low convex, underside black to brown, black in centre, with black striations,
becoming light brown towards the edge, agar stained brown (HKUCC 3442).
Known hosts: On wood of Brachyglottis repanda (Ju and Rogers, 1996) and
senescent culm of Sinobambusa tootsik (this study).
Known distribution: China (Hong Kong) (this study) and New Zealand (Ju and
Rogers, 1996).
Material examined: Hong Kong, New Territories, Tai Lam Country Park, on
senescent culm of Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9128);
Hong Kong Island, Victoria Peak, on dead culm of Arundinaria hindsii, 17 Aug. 1998,
Dequn Zhou (HKU(M) 9152).
Remarks: These collections resemble Hypoxylon aucklandiae, but the
ascospores are shorter (9-10 x 4-6 urn vs. 10-13 x 5-6 urn) (Ju and Rogers, 1996). The
surface colors and morphology of this species are reminiscent of H. hypomiltum, but
differs as in H. hypomiltum the ascospores are smaller (7-9 x 3-4 urn vs. 10-13 x 5-6
urn). This species was only reported from New Zealand (Yu and Rogers, 1996) and the
ascospores are at 7-9 urn in length, which are on the lower limit of the spore size. The
specimens are worthy of further study for comparsion with the specimens collected by
Ju and Rogers (1996), as this species has only previously been found in New Zealand.
Hypoxylon brevirimum Y.M. Ju and J.D. Rogers, A Revision of the Genus Hypoxylon:

85

Chapter 2: Taxonomy

365. 1996.
Anamorph: Nodulisporium (Ju and Rogers 1996). Colonies on PDA are greyish,
felty, low convex, underside dark grey, but no sporulating structure produced
(HKUCC 3444).
Known hosts: Bambusa chungii (this study) and corticated wood (Ju and
Rogers 1996).
Known distribution: China (Hong Kong) (this study) and Brazil (Ju and Rogers
1996).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HKU(M) 9131); on
senescent culm of B. shiuyingiana, 27 Aug. 1999, Dequn Zhou (HKU(M) 9364).
Remarks: The asci in these collections are slightly longer and broader in
comparison to those of Hypoxylon brevirimum (asci 126-144 x 12-14 um vs. 110-140
x 9-12 um) (Ju and Rogers 1996).
Hypoxylon cf. hughesii Y.M. Ju and J.D. Rogers, A Revision of the Genus Hypoxylon:
130. 1996.
Known hosts: Bambusa chungii (this study), decorticated wood, Ixerba
brecioides, Melicytus remiflorus and Nothofagus menziesii (Ju and Rogers, 1996).
Known distribution: China (Hong Kong) (this study) and New Zealand (Ju and
Rogers, 1996).
Material examined: Hong Kong, The New Territories, Tai Po Kau Nature
Reserve, on senescent culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HK(M)
9133).
Remarks: The stromatal surface of H. hughesii resembles that of H
dieckmannii but differs in that ascospores of//, dieckmannii are smaller (6.5-10 x 3.54 um vs. 9-12 x 4-5.5 um) (Ju and Rogers, 1996). Our collections match H. hughesii
but the stromata in our collections are smaller (25 mm long x 13 mm broad x 5-6 mm
thick vs. 7-60 mm x 3-25 mm broad x 5-6 mm thick) (Ju and Rogers, 1996). It needs
to be further confirmed through more collections.
Hypoxylon cf. intermedium (Schwein.: Fr.) Y.M. Ju and J.D. Rogers, A Revision of

86

Chapter 2: Taxonomy

the Genus Hypoxylon: 133. 1996.


Cultural characters: Colonies on OA covering Petri dish after 3 weeks, white,
felty to floccose, azonate, with diffuse margins; reverse uncolored. No sporulating
structure produced (HKUCC 3574).
Known hosts: On twigs of Fraxinus angustifolia (Ju and Rogers, 1996), on
senescent culm of Melocanna baccifera and Schizostachyum dumetorum (this study).
Known distribution: China (Hong Kong) (this study), Czechoslovakia, Moravia,
Switzerland and USA (Ju and Rogers, 1996).
Material examined: Hong Kong, Hong Kong Island, Pokfulam Country Park,
on senescent culm of Melocanna baccifera, 30 June 1998, Dequn Zhou (HKU(M)
9019); Lung Fu Shan Country Park, on dead culm of Schizostachyum dumetorum, 19
July 1998, Dequn Zhou (HKU(M) 9044).
Remarks: The collections have affinities with Hypoxylon intermedium, but
differ in having broader ascospores (14-20.5 x 7-9.5 um vs. 16-18 x 6-8 urn) and the
perispore is indehiscent in 10% KOH.
Hypoxylon karii Y.M. Ju and J.D. Rogers, A Revision of the Genus Hypoxylon: 138.
1996.
Anamorph: Colonies 2 cm diam. on PDA after one month, slow growing,
blackish grey, convex, velvety (HKUCC 3575).
Known hosts: Arundinaria hindsii (this study) and decaying herbaceous stems
(Ju and Rogers 1996).
Known distribution: China (Hong Kong) (this study) and India (Ju and Rogers
1996).
Material examined: Hong Kong, Hong Kong Island, Lung Fu Shan Country
Park, on dead culm of Arundinaria hindsii, July 30 1998, Dequn Zhou (HKU(M)
9079).
Remarks: This collection resembles Hypoxylon karii except in the width of the
ascospores (ascospores 10-12 urn vs. 7-8.5 urn) (Ju and Rogers 1996).
Hypoxylon rubiginosum (Pers.: Fr.) Fr. Summa Vegetativa Scandinavica II: 384. 1849.

87

Chapter 2: Taxonomy

Anamorph and cultural characters: A Nodulisporium-like structure was


produced on culture (Ju and Rogers, 1996 and this study). Colonies slow growing, ca.
5 cm diam. after one month, grey, granular, flat, with brown concentric rings,
underside brown and light brown towards margin (HKUCC 3010).
Known hosts: On corticated wood of Aesculus sp. (Dennis, 1986), dead culm of
Bambusa cornigera (this study), Fraxinus sp., Fagus sp., Wuercus tocemeyi, Populus
alba, Acer macrophyllum (Ju and Rogers, 1996).
Known distribution: China (Hong Kong) (this study), France, Germany, USA
(Ju and Rogers, 1996) and UK (Dennis, 1986).
Material examined: Hong Kong, The New Territories, Sai Kung, on dead culm
of Bambusa cornigera, 27 June 1998, Dequn Zhou (HKU(M) 8390).
Remarks: This collection resembles Hypoxylon rubiginosum, but differs in that
the perispores are indehiscent in 10% KOH.
Hypoxylon subgilvum Berk, and Broome, Botanical Journal of the Linnean Society 14:
120. 1873.
Cultural characters: Colonies growing up to ca. 4 cm diam. after one month,
white, cottony, becoming creamy towards margin (HKUCC 3137).
Known hosts: Arundinaria hindsii (this study), Berberis sp., Ficus sp. and
decorticated wood (Ju and Rogers, 1996).
Known distribution: Hong Kong (this study), India, Indonesia, Paraguay, South
Africa, Taiwan, USA, Venezuela and Zaire (Ju and Rogers, 1996).
Material examined: Hong Kong, The New Territories, Tai Shui Hang, Mu Tsz
Lam, on senescent culm of Arundinaria hindsii, 29 July 1998, Dequn Zhou (HKU(M)
9074).
Remarks: This collection is identical to H. subgilvum based on the description
by Ju and Rogers (1996). The stromata were covered by a layer of white filamentous
mass and Nodulisporium-like conidiophore but no conidia found.

KretzschmariellaViegas,Bragantia4: 105. 1944.

88

Chapter 2: Taxonomy

Kretzschmariella guaduae Viegas, the type species, was considered to be a


synonym of Hypoxylon culmorum Cooke by Petrak (Ju and Rogers, 1994). This
disposition was accepted by Miller (Ju and Rogers, 1994). Based upon cultural data,
including an anamorph that was unusual among the Xylariaceae, Ju and Rogers (1994)
reinstated Kretzschmariella. In Hypoxylon, the ascospores have a germ slit usually on
the convex side, the perispore of many taxa is dehiscent in KOH, and the
Nodulisporium-like anamorph occurs on wood. In Kretzschmariella, ascospores
usually have a germ slit on the flat or concave side, the perispore not dehiscent in
KOH and taxa generally occur on bamboo culms (Ju and Rogers, 1996).

Kretzschmariella culmorum (Cooke) Ju and Rogers. Mycotaxon 51: 242. 1994.


Known hosts: On the culms of Guardua, bamboo, Arundinaria sp. (Yu and
Roger, 1994) and Arundinaria amabilis (this study).
Known distribution: China (Hong Kong) (this study), Brazil, Grenada, Grenada,
Guadeloupe, Paraguay and USA (Ju and Rogers, 1994).
Material examined: Hong Kong, Hong Kong Island, Long Fu Shan, on culms
of Arundinaria amabilis, 3 June 1998, Dequn Zhou (HKU(M) 8337).
Rosellinia De Not., Giornale Botanico Italiano 1:1.1844.
Rosellinia was introduced by De Notaris based on Sphaeria aquila Fr. (Dargan
and Thind, 1979). It comprises about 100 species characterized by stromata with one
or few ascomata formed beneath a persistent or evanescent false subiculum (Lsessoe
and Spooner, 1994; Hawksworth et al., 1995). In the genus, asci are cylindrical to
clavate, with a barrel-shaped apical apparatus; ascospores are brown, fusiform or
ellipsoid to asymmetrically ellipsoidal, with a germ slit extending the full length, and
with sheath and/or appendages. Petrini (1992) monographed 20 species of Rosellinia
and also briefly described the distribution, host specificity and ecology in temperate
regions. A key to 20 species from temperate region was provided by Petrini (1992).
See under Astrocystis for comparison with this genus.

Rosellinia buxi H. Fabre, Annales des Sciences Naturelles Serie VI 9: 78. 1878.

89

Chapter 2: Taxonomy

Known hosts: Buxus sempervirens (Petrini, 1992) and Fargesia yunnanensis


(this study).
Known distribution: China (Kunming) (this study), Southern France (Petrini,
1994) and UK (Petrini, 1992).
Material examined: China, Yunnan, Kunming, West Hill, on senescent culm of
Fargesia yunnanensis, 4 July 1999, Dequn Zhou (HKU(M) 9263).
Remarks: Petrini (1992) did not describe asci, because at that time no recent
French material was available. Late Petrini (1993) designated a neotype for R. Buxi
collected from four new sites in Southern France. My collection resembles Rosellinia
buxi but differs in having slightly shorter ascospores (24-26 x 6-8 urn vs. 25-27.5 x 78 um) (Petrini, 1992). Based on my collection, Asci of/?, buxi are 206-216 x 10-12
um, tapering towards the stipitate end, stipe 38-64 um long, 8-spored, cylindrical,
apically rounded, with a massive, barrel-shaped subapical apparatus, 5-6 urn high, 4
um diam. This collection extends the host range and shows that this species has a
worldwide distribution from temperate (France) to subtropical regions (Kunming).
Rosellinia saccardii (Sacc.) L.E. Petrini, Sydowia 44: 226. 1992.
Known hosts: Fargesia yunnanensis (this study), Gleditschia, Robinia (Petrini,
1992) and Neosinocalamus affinis (this study).
Known distribution: China (Hong Kong) (this study), France, Germany and
Italy (Petrini, 1992).
Material examined: China, Yunnan, Kunming, Anlin, on senescent culm of
Neosinocalamus affinis, 20 June 1999, Dequn Zhou (HKU(M) 9257); 19 Aug. 1998,
Dequn Zhou (HKU(M) 9097); West Hill, on dead culm of Fargesia yunnanensis, 4
July 1999, Dequn Zhou (HKU(M) 9260).
Remarks: The dimension of ascospores in these collections overlap with those
of Rosellinia saccardii (Petrini, 1992).

90

Chapter 2: Taxonomy

Spirodecospora B.C. Lu, K.D. Hyde and W.H. Ho, Fungal Diversity 1: 169. 1998.
Spirodecospora is a monospecious genus and was established by Lu, Hyde and
Ho (1998) with S. bambusicola as the type species, described from Hong Kong.
Spirodecospora bambusicola B.C. Lu, K.D. Hyde and W.H. Ho, Fungal Diversity 1:
172. 1998.
Known hosts: On dead culms of Bambusa sp. (Lu et al, 1998), B. tuldoides,
Fargesia yunnanensis and Phyllostachys bambusoides (this study).
Known distribution: China (Hong Kong) (Lu et al, 1998).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of B. tuldoides, 19 Apr. 1999, Dequn Zhou (HKU(M) 9215); on
dead culm of Phyllostachys bambusoides, 8 July 1999, Dequn Zhou (HKU(M) 9278);
ibid, 5 Sept. 2000, Dequn Zhou (HKU(M) 9402); China, Yunnan, Kunming, West Hill,
on dead culm of Fargesia yunnanensis, 16 Sept. 1999, Dequn Zhou (HKU(M) 9356).
Remarks: The collections are identical to S. bambusicola with comparsion to
the type material and description (Lu et al., 1998).
Striatodecospora D.Q. Zhou, K.D. Hyde and B.S. Lu, Mycotaxon 76: 141-147 (2000)
See appendix.
Striatodecospora bambusae D.Q. Zhou, K.D. Hyde and B.S. Lu, Mycotaxon 76: 141147 (2000)
See appendix.

2.3.1.15 Familia Incertae Sedis


Bambicola D.Q. Zhou and K.D. Hyde, gen. nov.
Etymology: Referring to the bamboo host.
Ascomata immersed beneath a dome-shaped pseudostroma, visible as raised
blackened discs, cylindrical or ellipsoidal, soft, black, solitary or clustered. Ostiole
central, periphysate. Peridium thin, composed of a few layers of small, compressed,
brown-walled cells on the upper and lower wall. Paraphyses 5-10 um wide at the base,

91

Chapter 2: Taxonomy

filamentous, septate, hyaline, tapering distally (Fig 2.6.5). Asci 8-spored, broad
fusiform, short pedicellate, unitunicate, apically rounded, without an apical ring.
Ascospores multiseriate, filiform, hyaline, one-celled or multi-septate, without
refringent bands, straight or curved (Figs 2.6.6-2.6.7).
Bambicola is comparable with Pemphidium as it has ascospores without
refringent bands and asci without a refractive apical ring. In this way, it can be
distinguished from Linocarpon (Hyde, Taylor and Frohlich, 2000). Pemphidium
differs however, in having long cylindrical and long pedicellate asci, with a J-, wedgeshaped, subapical ring and ascospores with polar appendages. Bambicola can not be
confused with Oxydothis, as in Oxydothis, asci are cylindrical, with a J+ (occasionally
J-), wedge-shaped, or discoid, subapical ring, visible in size and shape, with a faint
canal leading to the apex (Hyde, Taylor and Frohlich, 2000).
Bambicola filiforma D.Q. Zhou and K.D. Hyde, sp. nov.

(Figs. 2.8.1-2.8.7)

Etymology: In reference to the filiform ascospores.


Ascomata immersed beneath a dome-shaped pseudostroma, visible as raised
blackened discs, cylindrical, soft, black, ca 1.5-2 mm diam. on the host, each
surrounded by a faint darkened zone after coalescing around groups of ascomata and
with central darkened ostiolar dot, mostly solitary or some gregarious; in section 325440 (am diam, 95-110 um high (Figs 2.8.1-2.8.2). Peridium 15-22 um wide,
comprising several layers of thin-walled, compressed, brown walled cells. Paraphyses
5-7.5 um wide at the base, hypha-like, filamentous, septate, hyaline, tapering distally
(Fig 2.8.5). Asci (87.5-) 95-105 x 20-22.5 um (3c= 101.5 x 21 um, n = 15), 8-spored,
broad fusiform, short pedicellate, apically rounded, without an apical ring (Figs 2.8.32.8.4). Ascospores 66-88 x 4-5 (am (3c = 77.4 x 4.3 um, n = 50), multiseriate, fusiform,
hyaline, 0-3 septate, straight or curved (Figs 2.8.6-2.8.7).
Material examined: Hong Kong, New Territories, Tai Shui Kang, Mui Tsz Lam,
on dead culm of Phyllostachys bambusoides, 29 July 1998, Dequn Zhou (HKU (M)
9068, holotype).
Other material examined: Hong Kong, New Territories, Tai Shui Kang, Mui
Tsz Lam, on dead culm of Phyllostachys bambusoides, 29 July 1998, Dequn Zhou

92

Figs. 2.8.1-2.8.7. Interference micrographs oi Bambicola filiforma (from (HKU(M) 9068,


holotype). 1. Appearance of ascomata on bamboo culm. 2. Section of ascoma. 3-4. Asci.
5. Paraphyses. 6.-7. Ascospores. Scale bars: 1 = 500 |j.m, 2 = 50 ^m, 3, 5 = 20 (am, 4, 6-7
= 10 urn.

Chapter 2: Taxonomy

(HKU (M) 9406).

Frondisphaeria K.D. Hyde, Mycoscience 37: 169. 1996.


The genus Frondisphaeria K.D. Hyde was introduced to accommodate fungi
with curved, long-fusiform ascospores with apiculate ends. The genus is most similar
to Linocarpon and Pemphidium, which also have filiform unicellular ascospores.
Linocarpon differs from Frondisphaeria, as ascomata in Linocarpon have conspicuous
ostioles, asci are cylindrical, and ascospores are filiform with polar appendages. In
Pemphidium, asci are cylindrical, with an indistinctive, subapical ring, and the
ascospores are cylindric-fusiform, with polar appendages comprising a hollow
cylinder containing mucilage (Hyde, Taylor and Frohlich, 2000). Frondisphaeria
comprises two species and is only known from palms (Frohlich and Hyde, 2000).
Frondisphaeria amplispora D.Q. Zhou and K.D. Hyde, sp. nov. (Figs 2.9.1-2.9.10)
Etymology: From Latin 'amplus' meaning 'broad' and ""spora1 meaning 'spore',
referring to the broad ascospores.
Ascomata developing beneath black, raised, dome-shaped slightly darkened,
irregular areas on the host surface, gregarious in groups of two or three, often with a
short ostiole; in section 240-310 um diam. (3c = 206 um, n = 10), 130-270 um high (x
= 267 um, n = 10), with a short periphysate ostiole (Figs. 2.9.1-2.9.2). Peridium 10-15
(am wide, comprising several layers of thin-walled, compressed brown walled cells
(Fig 2.9.3). Paraphyses 12-14 um wide at the base, hypha-like, filamentous, septate,
hyaline, tapering, 2 um wide at the apex (Fig 2.9.4). Asci 90-134 x (16-) 18-22 (-24)
um (x = 106 x 19.5 um, n = 20), 8-spored, broadly clavate, straight or flexible,
shortly pedicellate, apically rounded, with a J-, faint subapical ring (Figs 2.9.5-2.9.6).
Ascospores 100-110 (-114) x 4-6 um (3c = 102 x 5.5 um, n = 50), multiseriate,
relatively broadly filiform, straight or curved, hyaline, containing numerous refringent
septum-like bands, lacking a mucilaginous sheath or pads (Figs. 2.9.7-2.9.10).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa glaurescens cv. fernleaf, 18 May 1999, Dequn Zhou
(HKU(M) 9232, holotype).

93

Figs. 2.9.1-2.9.10. Interference micrographys of Frondisphaeria amplispora (from


(HKU(M) 9232, holotype). 1. Appearance of ascomata on bamboo culm (arrowed). 2.
Section of ascoma. 3. Section of peridium. 4. Paraphyses. 5-6. Asci. 7-10. Ascospores.
Scale bars: 1 = 200 urn, 2 = 50 urn, 3 = 20 urn, 5-6 = 10 urn, 4, 7-10 = 15 um.

Chapter 2: Taxonomy

Other material examined: Hong Kong, New Territories, Tai Po Kau Nature
Reserve, on senescent culm ofBambusa glaurescem cw.femleaf, 18 May 1999, Dequn
Zhou (HKU(M) 9405).
Remarks: Frondisphaeria amplispora is most similar to Linocarpon species in
that the ascomata of F. amplispora have inconspicuous ostioles and the ascospores
have refringent septum-like bands. They differ as asci of Linocarpon species are
cylindrical, and ascospores are filiform with polar appendages. Frondisphaeria
amplispora is similar to F. joanneii J. Frohl. and K.D. Hyde, but differs in asci and
ascospore dimensions (Hyde et al, 2000) (Table 2.5).
Table 2.5 Comparison of Frondisphaeria species
Taxon

Ascus length

Ascus width

Ascospore

Ascospore

Ascal ring

Ascal ring

(Hm)

(|im)

length (nm)

width (nm)

height (urn)

width (urn)

90-134

18-22

100-110

4-6

E joanneii

69-105

12-14

61-77.5

3-4

F. palmicola

158-225

22.5-30

81-115

8-10

F. amplispora

2.3.2 Basidiomycetes
2.3.2.1 Tricholomataceae
Marasmius Fr., Epicrisis Systematis Mycologici: 385. 1838.
Fries established Marasmius, which was typified by M. rotula (Scop.) Fr. In
1836 (Kiihner, 1926). A comprehensive theoretical framework for the taxonomic study
of Marasmius has been laid by Kiihner (1926, 1933, 1935), Kiihner and Romagnesi
(1953), and Singer (1958a, 1958b, 1965). Gilliam (1976) monographed Marasmius in
Northeastern United States and adjacent Canada and reviewed development of the
generic concept. Antonin and Noordeloos (1993) gave a historical survey of the genus.
About 350 species have been reported in this genus (Hawksworth, et al, 1995).
Marasmius rotula (Scop) Fr., Epicrisis Systematis Mycologici: 385 1838.
Known hosts: Abies borisiiregis, Abies cephalonica (Zervakis, Dimou and Balis,
1998), Acer negundo (Brenckle, 1918), Bambusa tuldoides (this study), Castanea
sativa (Zervakis et al, 1998), Fagus sylvatica (Zervakis et al, 1998), Quercus
conferta (Zervakis et al, 1998), on the rotten leaves in broadleaf forest (Wen and Sun,

94

Chapter 2: Taxonomy

1999).
Known distribution: Canada (Zervakis et al., 1998), China (Guangxi) (Wen and
Sun, 1999), China (Hong Kong) (Chang and Mao, 1995), Greece and USA (Zervakis
etal., 1998).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa tuldoides, 23 May 2000, Dequn Zhou (HKU(M) 9386).
Remarks: This collection is identical to M. rotula based on the description by
Zervakis et al. (1998).
2.3.3 Coelomycetes (mitosporic fungi)
Conidiocarpus Woron., Key to Fungi by Jaczewski Vol. 2 (Fungi imperfecti): 743.
1917.
Conidiocarpus was proposed by Woronichin (1917) for the single species
Conidiocarpus caucasicus Woron., which is the conidial state of Limacinula caucasica
Woron. (Hughes, 1976). The type collection was found on living leaves of Taxus and
Caucasus. Conidiocarpus pycnidia are commonly found in sooty mould colonies in
the tropics and subtropics (Hughes, 1976). They are readily distinguished from those
of Polychaeton (Pes.) Lev. by the stalk which is composed of closely adpressed
synnematous hyphae and by the conspicuous swelling which indicates the location of
the pycnidial cavity. The fructifications are essentially species of Conidiocarpus with
very robust pycnidia in which there have occurred one or more lateral proliferations of
the venter or neck of the lanceolate pycnidium, or a single, percurrent proliferation
through the ostiole to produce similar but smaller pedicellate pycnidia (Hughes, 1976).
Two species have been accepted (Hughes 1976; Hawksworth et al, 1995). Hughes
(1976) discussed the historical development of this genus.

Conidiocarpus caucasicus Woron., Key to Fungi by Jaczewski Vol. 2. (Fungi


Imperfecti): 743. 1917.
Teleomorph: Limacinula caucasica (Hughes, 1976).
Known distribution: China (Hong Kong) (this study) and USA (Hughes, 1976).
Known hosts: Bambusa beecheyana (this study) and Taxus (Hughes, 1976).

95

Chapter 2: Taxonomy

Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa beecheyana, 15 July 1998, Dequn Zhou (HKU(M) 9041).
Remarks: This collection is identified as Conidiocarpus caucasicus (Hughes,
1976), but the pycnidia are slightly shorter (520-580 vs. 540-650 urn) (Hughes, 1976).

Hyalopycnis Hohn., Hedwigia 60: 152. 1918.


Hyalopycnis Hohn. was introduced by Hohn (1918) and is typified by H.
blepharistoma (Berk.) Seeler. Seeler (1943) redescribed the genus and elaborated upon
the synonymy. The genus is monotypic (Sutton, 1980). Hyalopycnis is the anamorph
of the basidiomycete Heterogastridium, which is a monotypic genus in the
Heterogastridaceae (Hawksworth et al., 1995).
Hyalopycnis blepharistoma (Berk.) Seeler, Farlowia 1: 124. 1943.
Cultural characters: Colonies growing up to 6.5 cm diam. after one month,
brown, dusty, underside brown, the mycelia being sparse (HKUCC 3117).
Teleomorph: Heterogastridium (Hawksworth et al., 1995).
Known

distribution:

Australia,

China

(Hong

Kong)

(this

study),

Czechoslovakia (Sutton, 1980), Philippines (Dalisay, 1998) and USA (Sutton, 1980).
Known hosts: Aducus, Agaricus adustus, (Sutton, 1980), Dendrocalamus sp.
(Dalisay, 1998), D. pulverulentus (this study), Uritica and various agaric species
(Sutton, 1980).
Material examined: Hong Kong, New Territories, Sai Kung, on dead culm of
Dendrocalamus pulverulentus, 27 June 1998, Dequn Zhou (HKU(M) 8394).

Pestalotiopsis Steyaert, Bullletin Jardin Botanique de I'Etat Bruxelles 19: 300. 1949.
Pestalotiopsis was established by Steyaert (1949). The separation of
Pestalotiopsis and Truncatella Steyaert from Pestalotia Steyaert (1949) caused
considerable controversy (Sutton, 1969). Steyaert restricted Pestalotia to a single
species and reassigned several species formerly disposed in Pestalotia to
Pestalotiopsis and Truncatella (Raj Nag, 1985). Guba (1961) preferred to adopt a
broader generic concept. He reduced Pestalotiopsis, Truncatella and Labridella

96

Chapter 2: Taxonomy

Brenckle to synonymy with Pestalotia and accepted 220 species in that genus. Sutton
(1969) discussed in detail the arguments opposing or supporting either approach and
gave evidence favoring the re-arrangement proposed by Steyaert (1949). Sutton (1980)
and Raj Nag (1993) accepted Pestalotiopsis. The distinctions between the genera
among the Monochaetia (Sacc.) Allesch., Seiridium Nees and Pestalotia and
Pestalotalotiopsis complexes were discussed by Sutton (1969). Raj Nag (1985)
provided a key to Pestalotia-like genera. In Pestalotia, conidia are distoseptate while
in Pestalotiopsis and Monochaetia they are euseptate. In Monochaetia, the conidia
have a single and unbranched apical appendage, but in Pestalotiopsis the conidia have
several apical appendages in an apical crest or in several tiers on the wall of the apical
cell. There are 24 species in Pestalotiopsis (Raj Nag, 1993).

Pestalotiopsis uvicola (Speg.) Bissett, Canadian Journal of Botany 60: 2572. 1982.
Cultural characters: Colonies 4.5 cm. diam. after 38 days on PDA, cottony,
white, but no sexual sporulating structures produced after one month.
Known hosts: Bambusa tuldoides (this study), Gaura parviflora and Vitis
vinifera (Raj Nag, 1993).
Known distribution: China (Hong Kong) (this study), Italy and USA (Raj Nag,
1993).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa tuldoides, 14 Apr. 1999, Dequn Zhou (HKU(M) 9211).
Remarks: The collection has affinities wityh Pestalotiopsis uvicola following
the key and description given by Raj Nag (1993), but the conidia of this collection are
slightly smaller (16-20 * 4-6 urn vs. 18-25 x 5.5-7.5 urn).
2.3.4 Hyphomycetes (mitosporic fungi)
Acrodictys M.B. Ellis, Mycological Papers 79: 5. 1961.
Acrodictys was introduced by Ellis (1961) and typified by A. bambusicola M.B.
Ellis (Ellis, 1971). The genus is widespread and includes 26 species (Hawksworth et
al., 1995). Descriptions and illustrations with a key to the species were given by Ellis
(1971, 1976). Recently, Whitton (2000) updated the key with two new species.

97

Chapter 2: Taxonomy

Acrodictys bambusicola M.B. Ellis, Mycological Papers 79: 6. 1961.


Known hosts: Bambusa mutabilis (Dalisay, 1998), Pennisetum, Phyllostachys
glauca (this study) and unidentified bamboo (Ellis, 1971).
Known distribution: China (Hong Kong) (this study), Uganda and Venezuela
(Ellis, 1971).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa mutabilis, 17 July 1998, Dequn Zhou (HKU(M) 9030); on
dead culm of Bambusa tuldoides, 23 Dec. 1998, Dequn Zhou (HKU(M) 9208); ibid.
(HKU(M) 9210); on dead culm of Phyllostachys glauca, 6 June 1998, Dequn Zhou
(HKU(M) 8352).

Acrodicytys erecta (Ellis and Everh.) M.B. Ellis, Mycological Papers 79: 5. 1961.
Known hosts: Arundo (Ellis,

1971), Bambusa (Dalisay,

1998) and

Neosinocalamus qffinis (this study).


Known distribution: China (Kunming) (this study), Philippines (Dalisay, 1998),
USA and Venezuela (Ellis, 1971).
Material examined: China, Yunnan, Kunming, Anlin, Qiu Mu Yuan, on the
dead of Neosinocalamus qffinis, 19 Aug. 1998, Dequn Zhou (HKU(M) 9099).
Remarks: The collection is identified as Acrodictys erecta (Ellis, 1971), but the
conidia are slightly narrower (12-18 um vs. 15-22 um).
Acrodictys fuliginosa B. Sutton, Canadian Journal of Botany 47: 854. 1976.
Known hosts: Bambusa chungii (this study), Populus balsamifera and P.
tremuloides (Ellis, 1976).
Known distribution: Canada (Ellis, 1976) and China (Hong Kong) (this study).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa chungii, 9 Aug. 1998, Dequn Zhou (HKU(M) 9138).
Remarks: This collection resembles Acrodictys fuliginosa (Ellis, 1976), but the
conidia are slightly wider (12-22 \im vs. 12-16 pm).

98

Chapter 2: Taxonomy

AcrodictyssacchariM.B. Ellis, Mycological Papers 125: 7. 1971.


Cultural characters: Colonies growing fast up to 5 cm diam. after one week on
PDA, white, felty, effuse, with aerial mycelium, underside creamy green (HKUCC
3387).
Known hosts: Saccharum officinarum (Ellis, 1976) and Schizostachyum
dumetorum (this study).
Known distribution: China (Hong Kong) (this study) and Jamaica (Ellis, 1976).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Schizostachyum dumetorum, 8 Sept. 1998, Dequn Zhou (HKU(M)
9107).
Alternaria Nees, Das System der Pilze und Schwamme: 72. 1816.
Alternaria was introduced by Nees (1816) with A. alternata (FT.) Keissl. as the
type species. The genus includes about 50 species (Hawksworth et ah, 1995).
Descriptions of several species are provided by Joly (1964), Ellis (1971, 1976), and
Simmons (1990).
Alternaria alternata (Fr.) Keissl., Beihefte zum Botanischen Centralblatt 29: 434.
1912.
Cultural characters: Colonies quickly growing up to 5 cm diam. after one
week, grey, cottony, effuse, underside greenish grey (HKUCC 3458, 3459).
Known hosts: foodstuffs, soil, textile, seeds (Domsch et al., 1993; Ellis, 1976),
on various plants as parasites, as endophytes of Juniperus occidentalis, Calocedrus
decurrens (Petrini and Carroll, 1981), Bambusa tuldoides (Dalisay, 1998) and
Phyllostachys nigra (this study).
Known distribution: Cosmopolitan, China (Hong Kong) (Dalisay, 1998) and
China (Kunming) (this study).
Material examined: China, Yunnan, Kunming, International Horticultural
Exposition, on senescent culm oi Phyllostachys nigra, 28 Sept. 1998, Dequn Zhou
(HKU(M)9171).
Remarks: This fungus is very frequently mentioned in plant pathological

99

Chapter 2: Taxonomy

literature as causing leaf spots and fruit rots (Holliday, 1980). In nearly all cases it is
behaving as a wound parasite or invading a host that is physiologically or
pathologically weakened (Holliday, 1980). It is also a common saprobe in soil
(Domsch and Gams, 1993).

Arthrinium Kunze, Mykologische Hefte (Leipzig) 1:9. 1817.


Arthrinium was erected by Kunze (1817) and typified by A. caricicola Kunze
(Ellis, 1971; Domsch, Gams and Anderson, 1993; Hawksworth et al, 1995). Variation
within the genus is seen in three areas: (1) conidial shape, which is lenticular to
globose, but which may be short clavate, polygonal in face view, limoniform or curved,
sometimes with ends extended into horns which point outward in one species and
inward in another; (2) the germ slit, which extends halfway around the conidium in the
type species, forms a complete equatorial band in several species, is reduced to a pore
in some species, and is absent from at least one species; (3) the presence in some
species of a "sterile cell" formed at or near the top of the conidiophore, which may
represent aborted conidia; in several species these " sterile cells" are irregularly lobed
(Samuels, McKenzie and Buchanan, 1981). Samuels et al. (1981) compared this genus
with other related genera, such as Cordelia, Pteroconium and Scyphospora. A key to
species is provided by Ellis (1971) and Domsch et al. (1993).

Arthrinium state of Apiospora montagnei Sacc, Nuovo Giornale Botanico Italiano 7:


306. 1875.
Telemorph: Apiospora montagnei (Ellis, 1971).
Known hosts: Very common on bamboos and other substrata (Ellis, 1971) and
Phyllostachys nigra (this study).
Known distribution: Cosmopolitan (Ellis, 1971 and this study).
Material examined: China, Yunnan, Kunming, Kunming

International

Gardening Exposition, on senescent culm of Phyllostachys nigra, 18 Sept. 1998,


Dequn Zhou (HKU(M) 9172).

100

Chapter 2: Taxonomy

Arthrinium saccharicola Stev., Journal of Department of Agricultural of Puerto Rico


1:223. 1917.
Known hosts: Bambusa tuldoides (this study) and Saccharum officinarum (Ellis,
1971; Stevens, 1975).
Known distribution: Argentina (Ellis, 1971; Stevens, 1975), China (Hong Kong)
(this study), India, Pakistan, Puerto Rico and Venezuela (Ellis, 1971; Stevens, 1975).
Material examined: Hong Kong, the New Territories, Sun Leung Tam Country
Park, on senescent culm of Bambusa sinospinosa, 27 Jan. 1999, Dequn Zhou (HKU(M)
9207).
Arthriniumyunnanensis Zhou, Goh and K.D. Hyde, sp. nov.

(Figs 2.10.1-2.10.8)

Etymology: Referring to the location.


Colonies beneath the bamboo epidermis which split longitudinally to expose
the shiny black spores masses, 1-2 x 0.3-0.5 mm (3c = 1.4 x 0.4 mm, n = 10) (Figs
2.10.1-2.10.2). Conidiophores erect, simple, flexuous, cylindrical, verrucose, septate,
colorless (Figs 2.10.3, 2.10.5, 2.10.7, 2.10.8). Conidia 20-26 um diam. in face view,
16-18 um wide, lenticular, globose or subglobose, pale olivaceous brown to brown
(Figs 2.10.4-2.10.8).
Teleomorph: Unknown.
Known distribution: China (Kunming) (this study).
Known hosts: Neosinocalamus qffinis (this study).
Material examined: CHINA, Yunnan, Kunming, An Lin, Yu Long Wan, on
dead culm of Neosinocalamus afflnis, 19 Aug. 1998, Dequn Zhou (HKU(M) 9101,
holotype).
Other material examined: China, Yunnan, Kunming, West Hill, on senescent
culm of Neosinocalamus affinis, 4 Oct. 1998, Dequn Zhou (HKU(M) 9177); Hong
Kong, The New Territories, Tai Po Kau Nature Reserve, on senescent culm of
Bambusa chungii, 8 Sept. 1998. Dequn Zhou (HKU(M) 9141).
Remarks: Conidia of Arthrinium yunnanensis are much larger than those of all
other species in Arthrinium (Bhat and Kendrick, 1993; Ellis, 1971; Samuels et at.,
1981).

101

Figs. 2.10.1-2.10.8. Interference micrographs of Arthrinium yunnanensis (from (HKU(M) 9101, holotype).
1. Appearance of colonies beneath the bamboo epidermis which splits longitudinally to expose the shiny
black spore masses. 2-8. Conidiophores and conidia. Scale bars: 1 = 500 (am, 3 = 10 |im, 2,4-8 = 10 urn.

Chapter 2: Taxonomy

Brachysporiella Bat., Boletin da Secretaria de Agriculture 19: 109. 1952.


Brachysporiella was established by Batista with the type species B. gayana Bat.
in 1952 (Ellis, 1971). The genus comprises 7 species (Dalisay, 1998). Conidiophores
are macronematous, mononematous, with number of short branches near the apex,
brown or dark brown and smooth. Conidiogenous cells are integrated, terminal,
monoblastic, percurrent, cylindrical, doliiform or lageniform; conidia are solitary,
acrogenous, simple, 2- or pluri-septate, smooth, clavate, pyriform or turbinate (Ellis,
1971). Dalisay (1998) found Brachysporiella gayana Bat. in Hong Kong on the leaves
of Dendrocalamus sp. and senescent culm of Bambusa sp.

Brachysporiella pulchra (Subraman.) S. Hughes, New Zealand Journal of Botany 17:


184. 1979.
Cultural characteristics: Colony reaching ca 6 cm diam. on PDA after one
month, white, cottony, with copious aerial hyphae, no sexual sporulating structures
produced (HKUCC 3465).
Known hosts: Bambusa multiplex (Hughes, 1979), and Bambusa vulgaris var.
vittata (this study), Phyllostachys edulis, Psidium guajava and rotten wood (Hughes,
1979).
Known distribution: China (Hong Kong) (this study), India and Japan (Hughes,
1979).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa vulgaris var. vittata, 8 Sept. 1998, Dequn Zhou (HKU(M)
9111).
Remarks: Hughes (1979) cited several measurements for the conidia on
different hosts from different places, so conidial size is not a reliable character for
identification of this species. This collection is identical to Brachysporiella pulchra
based on conidial morphology, and conidiophore length as well as conidia dimensions
(Hughes, 1979).

102

Chapter 2: Taxonomy

Cordelia Speg., Anales de la Sociedad Cientifica Argentina 22: 210. 1886.


Spegazzini (1886) established Cordelia with C coniosporioides Speg. as the
type species (Samuels et al, 1981). Species of Cordelia resemble Arthrinium species
in their lenticular conidia with a hyaline rim, as well as in most other aspects. The only
distinguishing feature of Cordelia is the dark setae, which may accompany the
conidiophores. No species of Arthrinium sensu stricto has such setae. Arthrinium and
Cordelia are so similar that the presence or absence of setae does not constitute an
adequate genetic difference (Samuels et al., 1981). Only one teleomorph is known to
have a Cordelia phase and that is Apiospora setosa. Samuels et al. (1981) regarded
Cordelia as a synonym of Arthrinium. Their idea was not accepted by Hawksworth et
al. (1995) and Goos and Uecker (1992). The setae were very conspicuous in all
Cordelia species found on bamboo in this study, and thereafter Cordelia is treated as a
separate genus.

Cordelia coniosporioides Speg., Anales de la Sociedad Cientifica Argentina 22: 210.


1886.
Known hosts: On bamboo culms (Ellis, 1971) and Bambusa glaucescens cv.
fernleaf (this study).
Known distribution: China (Hong Kong) (this study), Paraguay and Venezuela
(Ellis, 1971).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa glaucescens cv. fernleaf, 18 May 1999, Dequn Zhou
(HKU(M)9231).
Remarks: This collection is identical to C coniosporioides based on the
description by Ellis (1971).
CordeliajohnstoniiMB. Ellis, Mycological Papers 103: 31. 1965.
Teleomorph: Apiospora setosa (Samuels et al, 1981).
Known hosts: On culms of Bambusa blumeana (Ellis, 1971) and B. tuldoides
(this study).
Known distribution: China (Hong Kong and Kunming) (this study), Malaysia

103

Chapter 2: Taxonomy

(Ellis, 1971) and USA (Goos and Uecker, 1992).


Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of B. tuldoides, 9 Mar. 1999, Dequn Zhou (HKU(M) 9239); China,
Yunnan, Kunming, Chengong, on senescent culm of Neosinocalamus affinis, 15 Sept.
1999, Dequn Zhou (HKU(M) 9357).
Remarks: The specimens resemble Cordelia johnstonii, although the conidia
are slightly larger (18-30 vs. 19-22 urn) (Ellis, 1971).

Dictyosporium

Corda,

Weitenweber's

Beitrage

zur

gesammten

Natur

und

Heiwissenschaften: 87. 1836.


Goh et al. (1999) monographed Dictyosporium with 21 accepted taxa including
three new species and excluded 16 unacceptable or doubtful species. They also
provided a key to the accepted species. Whitton (2000) added 4 more new species to
Dictyosporium, which are yet to be formally published, and gave a key to the species
occurring on the Pandanaceae.

Dictyosporium zeylanicum Petch, Annals of the Royal Botanic Gardens, Peradeniya 6:


252(1917).
Known hosts: Bambusa tuldoides (this study) and dead wood (Goh et al, 1999).
Known distribution: China (Hong Kong) (this study), Belize (Morris, 1978) and
Sri Lanka (Goh et al, 1999).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa tuldoides, 23 May 2000, Dequn Zhou (HKU(M) 9385).
Didymobotryum Sacc. Sylloge Fungorum 4: 626. 1886.
The genus was established by Saccardo (1886) with the type species of
Didymobotryum rigidum (Berk, and Broome) Sacc. The genus is commonly found on
woody debris, bamboo and palm spathes (Seifert, 1990). Seifert (1990) discussed the
history of Didymobotryum and Ellis (1971) discussed the taxonomy. This genus
comprises 6 species (Hawksworth et al., 1995).

104

Chapter 2: Taxonomy

Didymobotryum rigidum (Berk, and Brome) Sacc, Sylloge Fungorum 4: 627. 1886.
Cultural characters: Colonies growing relatively fast, ca. 5 cm diam. after one
week. The colonies being creamy, pink, felty, flat, underside dark brown, agar stained
brown (HKUCC 3002).
Known hosts: Arundinaria amabilis (this study), dead wood (Ellis, 1971),
Indocalamus sinicus and Sinobambusa tootsik (this study).
Known distribution: China (Hong Kong) (this study), Sri Lanka and Vietnam
(Ellis, 1971).
Material examined: Hong Kong, Hong Kong Island, Long Fu Shan Country
Park, on senescent culms of Arundinaria amabilis, 4 June 1998, Dequn Zhou
(HKU(M) 8336); New Territories, Tai Lam Country Park, on the dead culm of
Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9124); Tai Po Kau Nature
Reserve, on the dead culm of Bambusa textilis, 6 June 1998, Dequn Zhou (HKU(M)
8346); Yuan Long, on the dead culm of Sinobambusa tootsik, 8 June 1998, Dequn
Zhou (HKU(M) 8370); ibid. (HKU(M) 8375); Sai Kung, on the dead culm of
Indocalamus sinicus, 27 June 1998, Dequn Zhou (HKU(M) 8400).
Remarks: This species is quite common on dead bamboo culms during the rainy
season, but was scarcely found during the dry season.
Didymobotryum verrucosum Hino and Katumoto, Bulletin of the Faculty of
Agricultural, Yamaguti University 10: 1181. 1959.
Known hosts: Arundinaria hindsii (this study) and Sasa (Ellis, 1971).
Known distribution: China (Hong Kong) (this study) and Japan (Ellis, 1971).
Material examined: Hong Kong, Hong Kong Island, Victoria Peak, on the dead
culm of Bambusa sp., 8 June 1998, Dequn Zhou (HKU(M) 8343); Long Fu Shan
Country Park, on the dead culm of Arundinaria hindsii, 30 July 1998, Dequn Zhou
(HKU(M) 9086).
Remarks: The collection is similar to Didymobotryum verrucosum except that
the conidia are slightly shorter (11-19 um vs. 15-26 um) (Ellis, 1971).

105

Chapter 2: Taxonomy

Doratomyces Corda, Sturm's Deutschlands Flora, iii (Pilze), Band 3, Heft 7: 65. 1829.
Doratomyces was introduced by Corda (1829), which was typified by D.
stemonitis (Pers. ex. Fr.) Morton and Smith (Corda, 1937). Morton and Smith (1963)
amended the genus. Eight species, which are widespread in the world, are included in
the genus (Hawksworth et ai, 1995).

Doratomyces microsporus (Fr.) Morton and Smith, Mycological Papers 86: 77. 1963.
Cultural characters: Colonies on PDA growing slowly, about 1.5 cm after 7
days, white, cottony, producing conidiophores and conidia in the central colony.
Known hosts: On dead wood, dead leaves of grasses and sedges, herbaceous
stems, also isolated from feathers, dung and soil (Ellis, 1971) and on dead culms of
Bambusa tuldoides (this study).
Known distribution: China (Hong Kong) (this study) and Europe (Ellis, 1971).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa tuldoides, 19 Apr. 1999, Dequn Zhou (HKU(M) 9214).
Remarks: The conidia size in this collection is slightly larger (5-6 x 3-4 um vs.
3-5 x 2-3 urn) (Ellis, 1971).
Ellisembia Subram., Proceedings of Indian Natural Science Academy B38: 4:179.
1992.
Subramanian (1992) established Ellisembia to accommodate species previously
assigned to the genus Sporidesmium. The genus comprises 12 species having
proliferating, or percurrent and regular conidiophores and pseudoseptate conidia
(Subramanian, 1992). Descriptions and transfers of some species of Sporidesmium on
leaves of Freycinetia and Pandanus are provided by McKenzie (1995).

Ellisembia bambusicola (M.B. Ellis) D.Q. Zhou and K.D. Hyde, comb. nov.
Basionym: Sporidesmium bambusicola M.B. Ellis, Mycological Papers 70: 43.
1965.
Known hosts: Arundinaria hindsii (this study), Bambusa sp. (Ellis, 1976) and
Indocalamus sinicus (this study).

106

Chapter 2: Taxonomy

Known distribution: China (Hong Kong) (this study), Ghana and Sierra Leone
(Ellis, 1976).
Material examined: Hong Kong, Hong Kong Island, Victoria Peak, on the dead
culm of Arundinaria hindsii, 17 Sept. 1998, Dequn Zhou (HKU(M) 9199); New
Territories, Sai Kung, on the dead culm of Indocalamus sinicus, 27 June 1998, Dequn
Zhou (HKU(M) 9056); ibid. (HKU(M) 9057).
Remarks: This species is transferred from Sporidesmium bambusicola based on
the delimitation of Ellisembia (Subramanian, 1992). Ellisembia bambusicola is similar
to E. bambusae (= Sporidesmium bambusae), but in E. bambusae conidia are wider
with a mid or dark brown basal cell (Ellis, 1976). The conidia in these collections
agree with Sporidesmium bambusicola, although they are slightly shorter (56-96 urn
vs. 65-105 urn) (Ellis, 1976).

Ellisembia coronata (Fuckel) Subram., Proceedings of Indian Natural Science


Academy, B58: 183(1992)
Known hosts: Acer, Fagus, Helminthosporium velutinum, Philadelphus,
Sambucus, Sarothamnus (Ellis, 1976) and Sinobambusa tootsik (this study).
Known distribution: China (Hong Kong) (this study) and Europe, including UK
(Ellis, 1976).
Material examined: Hong Kong, Hong Kong Island, Long Fu Shan Country
Park, on the dead culm of Arundinaria hindsii, 7 June 1998, Dequn Zhou (HKU(M)
8341); New Territories, Tai Lam Country Park, on the dead culm of Sinobambusa
tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9121).
Remarks: These specimens are identical to Ellisembia coronata (Ellis, 1976),
but the conidia is slightly wider (10-14 urn vs. 9-12 urn) (Ellis, 1976).
Ellisembia pseudoseptata (M.B. Ellis) D.Q. Zhou and K.D. Hyde, comb. nov.
Basionym: Sporidesmium pseudoseptatum M.B. Ellis, Mycological Papers 103:
44. 1965.
Known hosts: On dead twigs (Ellis, 1976) and on senescent culm of
Phyllostachys heteroclata (this study).

107

Chapter 2: Taxonomy

Known distribution: China (Kunming) (this study) and Sierra Leone (Ellis,
1976).
Material examined: China, Yunnan, Kunming, West Hill, on senescent culm of
Phyllostachys heteroclata, 16 Sept. 1999, Dequn Zhou (HKU(M) 9361).
Remarks: According to the definition of Ellisembia, this species should be
transferred to Ellisembia, as the conidia are pseudoseptate (Ellis, 1976). This species
resembles E. coronata, however, in the latter species, conidia are much wider and pale
brown, while the conidia in E. pseudoseptata are pale straw-colored (Ellis, 1976). This
specimen is identified as Ellisembia pseudoseptatum, but conidia in these collections
are slightly longer (52-64 urn vs. 36-56 um) (Ellis, 1976).

Endophragmiella B. Sutton, Mycological Papers 132: 3. 1973.


Endophragmiella was proposed by Sutton (1973) for two species, E. pallescens
B. Sutton and E. candensis (Ellis and Everh.) B. Sutton. These species were described
as hyperparasites, with conidiophores, conidiogenous cells, and conidia typically pale
brown and thick-walled. Conidiophores are frequently branched, with conidiogenous
cells showing percurrent growth in E. pallescens. Ellis (1976) distinguished
Endophragmiella from Endophragmia primarily by conidiophores which branched
several times near the base and conidia that have a small protuberant thin-walled peg
at the base. The peg is evidently the frill derived from the distal end of the
conidiogenous cell, but Hughes (1979) found this feature in several species that Sutton
(1973) included in Endophragmia. Hughes (1979) transferred many species from
Endophragmia to Endophragmiella, emphasizing conidium succession and percurrent
proliferations, which arise from the penultimate cell of the conidiophore and grow
through the old conidiogenous cell apex.
In Endophragmiella, branching of conidiophores and conidium separation are
considered to have little or no generic value (Hughes 1979). Keys to species have been
provided by Kirk (1985) and Ellis (1976). There are 37 widespread species in the
genus (Hawksworth et al. 1995).

108

Chapter 2: Taxonomy

Endophragmiella oblonga (Matsush.) S. Hughes, New Zealand Journal of Botany 17:


139. 1979.
Known hosts: Bambusa sp., Dendrocalamus (Dalisay, 1998), Phyllostachys
bambusoides (this study) and P. edulis (Hughes, 1979),.
Known distribution: China (Hong Kong) (Dalisay, 1998), Japan (Hughes, 1979)
and the Philippines (Dalisay, 1998).
Material examined: Hong Kong, The New Territories, Tai Shui Kang, on
senescent culm of Phyllostachys bambusoides, 29 July 1998, Dequn Zhou (HKU(M)
9071); Tai Lam Country Park, on dead culm of Sinobambusa tootsik, 8 Sept. 1999,
Dequn Zhou (HKU(M) 9123).
Remarks: Matsushima (1975) collected this species in Japan but he identified it
as Endophragmia oblonga Matsush. Hughes (1979) treated it as a synonym of
Endophragmiella oblonga. This collection is consistent with E. oblonga, but conidia
are slightly wider (10-12 urn vs. 9.5-11.5 um).

Endophragmiella subolivacea (Ellis and Everh.) S. Hughes, Fungi Canadenses 129:


144. 1978.
Known hosts: Acer rubrum (Kirk 1981), Acer sp. (Ginns, 1980), and
Phyllostachys glauca (this study) and wood twigs (Castaneda et al. 1998).
Known distribution: Canada (Ginns, 1980), China (Hong Kong) (this study),
Cuba (Castaneda et al, 1998) and USA (Kirk 1981).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Phyllostachys glauca, 6 June 1998, Dequn Zhou (HKU(M) 8358).
Exserticlava (Matsush.) S. Hughes, New Zealand Journal of Botany 16: 333 (1978)
Exserticlava was introduced by Hughes (1978b) to accommodate E. triseptata
(Matsush.) S. Hughes as the type species and E. vasiformis (Matsush.) S. Hughes as
the second species. The genus was based, respectively, on Cordana vasiformis
Matsush. (isolated from wood) and C. triseptata Matsush. (isolated from bamboo
stems) (Matsushima, 1975). Exserticlava species have widespread distribution and
have been reported from Ethiopia (Bhat and Sutton, 1985), India (Rao and de Hoog,

109

Chapter 2: Taxonomy

1986), Japan (Matsushima, 1975), Kenya (Kirk, 1985), Malaysia (Kuthubutheen and
Nawawi, 1998), New Zealand (Hughes, 1978b) and North America (Crane and
Schoknecht, 1982). Whitton (2000) provided a key to 4 accepted species.
Exserticlava vasiformis (Matsush.) S. Hughes, New Zealand Journal of Botany 6: 311.
1978.
Known hosts: Bambusa tuldoides (Dalisay, 1998), Melocanna baccifera (this
study) and submerged twigs (Hughes, 1978b).
Known distribution: Hong Kong (Dalisay, 1998 and this study), Japan,
Malaysia (Kuthubutheen and Nawawi, 1998), New Zealand and Taiwan (Matsushima,
1980).
Material examined: Hong Kong, Hong Kong Island, No. 1 Pokfulam Reservoir,
on dead culm of Melocanna baccifera, 30 June 1998, Dequn Zhou (HKU(M) 9052);
New Territories, Tai Po Kau Nature Reserve, on senescent culm of Bambusa tuldoides,
June 1995, T.E. Dalisay and F. Layug, TD46SH (HKU(M) 9453); ibid. (HKU(M)
9454).
Remarks: The collections described here agrees well with the Malaysia
collection, although the conidia are slightly longer (20-25 urn vs. 18-24 um)
(Kuthubutheen and Nawawi, 1998).
Gilmaniella G.L. Barron, Mycologia 56: 514. 1964.
Barron (1964) introduced Gilmaniella with the type species G. humicola G.L.
Barron. The genus is saprobic in soil (Barron, 1964) or on dead plant substrata
(Dalisay, Goh and Hyde, 1998). Dalisay (1998) reviewed the genus and gave a key to
6 accepted species.
Gilmaniella bambusae Umali, Goh and K.D. Hyde, Mycological Research 102: 435.
1998.
Cultural characters: Colonies fast growing, ca. 5-6 cm diam. after two weeks,
white, velvety, underside orange in the centre and with a creamy margin (HKUCC
3018).
Known hosts: Bambusa mutabilis (this study), Bambusa tuldoides (Umali et al.,

no

Chapter 2: Taxonomy

1998); B. vulgaris var. vittata, Dendrocalamus bambusoides, Indosasa sinica,


Neosinocalamus affinis and Phyllostachys pubescens (this study).
Known distribution: China (Hong Kong and Kunming) (Umali et al., 1998 and
this study).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa vulgaris var. vittata, 8 Sept. 1998, Dequn Zhou (HKU(M)
9114); on the dead culm of Bambusa mutabilis, 15 July 1998, Dequn Zhou (HKU(M)
9034); on senescent culm of Phyllostachys bambusoides, 8 July 1999, Dequn Zhou
(HKU(M) 9371); 5 Aug. 1999, Dequn Zhou (HKU(M) 9330); ibid. (HKU(M) 9334);
Sai Kung, on the dead culm of Dendrocalamus pulverulentus, 27 June 1998, Dequn
Zhou (HKU(M) 9009); on senescent culm of Bambusa tuldoides, 19 Apr. 1999, Dequn
Zhou (HKU(M) 9217); China, Yunnan, Kunming, Anlin, Qiu Mu Yuan, on the dead
culm of Phyllostachys pubescens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9155); on dead
culm of Neosinocalamus affinis, 9 Aug. 1998, Dequn Zhou (HKU(M) 9094); 20 June
1999, Dequn Zhou (HKU(M) 9256); ibid. (HKU(M) 9267); ibid. (HKU(M) 9281); 17
June, 1999; Dequn Zhou (HKU(M) 92920; West Hill, on the dead culm of
Dendrocalamus bambusoides, 4 Oct. 1998, Dequn Zhou (HKU(M) 9183);
International Horticultural Exposition, on the dead culm of Indosasa sinica, 28 Sept.
1998, Dequn Zhou (HKU(M) 9173); West Hill, on dead culm of P. heteroclata, 6 Aug.
2000, Dequn Zhou (HKU(M) 9389).
Remarks: The collections are identified as Gilmaniella bambusae based on the
description given by Dalisay et al. (1998). The conidia are slightly larger than those of
G. bambusae (4.5-5 urn diam. vs. 3.5-4.5 um). The species is very common on
bamboo in different habitats and can exist in the rainy and dry seasons in Hong Kong
and Yunnan. This species shows host recurrence on bamboo in Hong Kong and
Kunming.
Graphium Corda, Icones Fungorum Hucusque Congnitorum 1:18. 1837.
Corda (1837) established Graphium for two species, G. penicillioides Corda
and G. tenuissimum Corda. The genus is the anamorph of Ophiostoma (Hawksworth et
al, 1995). There are 10 species in this genus (Hawksworth et al, 1995).

ill

Chapter 2: Taxonomy

Graphium putredinis (Corda) S. Hughes, Canadian Journal of Botany 36: 770. 1958.
Known hosts: On senescent culm of Bambusa tuldoides (this study) and dead
herbaceous stems, especially common on Brassica, also isolated from soil (Ellis,
1971).
Known distribution: Europe (Ellis, 1971) and Hong Kong (this study).
Material examined: Hong Kong, New Territories, Tai Kau Po Nature Reserve,
on senescent culm oi Bambusa tuldoides, 19 Apr. 1999, Dequn Zhou (HKU(M) 9216).
Remarks: This collection matches Graphium putredinis (Ellis, 1971), but
conidia are slightly shorter (6-8 (-10) urn vs. 5-1 lum) (Ellis, 1971).

Janetia M.B. Ellis, More Dematiaceous Hyphomycetes: 33. 1976.


Janetia was established by Ellis (1976) with J. euphorbiae M.B. Ellis as type
species and another species, J. faureae (Piroz.) M.B. Ellis. Hughes (1983) expanded
the concept of the genus with the addition of species with monoblastic conidiogensis
and Sutton and Pascoe (1988) developed further the concept and included species
having distoseptate conidia. Janetia M.B. Ellis (1976) was reviewed with an emended
description of the genus by Goh and Hyde (1996a), who accepted 16 species.
Janetia synnematosa A. Sivan. and W.H. Hsieh, Mycological Research 94: 566. 1990.
This species has been formally published as a new record to Hong Kong (see
Appendix).
Penzigomyces Subram., Proceedings of the Indian Natural Science Academy, B58:
186(1992)
Subramanian

(1992)

reassessed

Sporidesmium Link

and

established

Penzigomyces Subram. with type species P. nodipes (Penz. and Sacc.) Subram. for
fungi having simple conidiophores which are septate, brown with regular, successive,
doliiform, lageniform or nodose percurrent proliferations. Conidia are gangliar,
solitary, acrogenous, euseptate, brown and dry. Six new species oi Penzigomyces were
added to the genus by Subramanian (1997).

112

Chapter 2: Taxonomy

Penzigomyces flagellata (S. Hughes) Subram. Proceedings of Indian Natural


Scientific Academy, B58: 186 (1992)
This species has been formally published as a new record for Hong Kong (see
Appendix).

Penzigomyces uapacae (M.B. Ellis) Subram., Proceedings of the Indian Natural


Science Academy B58: 187. 1992.
Known hosts: overgrowing colonies of Echidnodes on leaves of seedling trees
(Ellis, 1976) and on the dead culm of Neosinocalamus affinis, Phyllostachyum
bambusoides and P. pubscens (this study).
Known distribution: China (Kunming) (this study) and UK (Ellis, 1976).
Material examined: China, Yunnan, Kunming, Anlin, Qiu Muyuan, on
senescent culm of Phyllostachys pubscens, 5 Oct. 1998, Dequn Zhou (HKU(M) 9159);
Lunan, on the dead culm of Phyllostachyum bambusoides, 30 Sept. 1998, Dequn Zhou
(HKU(M) 9181); Chengong, on the dead culm of Neosinocalamus affinis, 4 Oct. 1998,
Dequn Zhou (HKU(M) 9185); on the dead culm of Phyllostachyum bambusoides, 4
Oct. 1998, Dequn Zhou (HKU(M) 9186).
Remarks: Penzigomyces uapacae resembles Repetophragma aburiense (M.B.
Ellis) Subram., but in the latter species, conidia are 2-4-septate and trunctate at the
base (Ellis, 1976; Subramanian, 1992). The conidium size range of these collections is
slightly wider than those of P. uapacae (6-8 um vs. 6-7 um) (Ellis, 1976).

Periconia Tode, Fungi Mecklenburgenses Selecti 2: 2. 1791.


Periconia was introduced by Tode (1791) with P. lichenoides Tode ex Merat as
the type species (Ellis, 1971). It includes 30 species (Hawksworth et al., 1995) and a
key to some of the species is provided by Ellis (1971, 1976). Marson and Ellis (1953)
discuss the taxonomy and history of Periconia. The genus is characterized by mainly
macronematous, mononematous conidiophores, at the apex with a spherical head of
conidia produced by mono- or polyblastic conidiogenous cells. The conidia are
catenate, spherical, subspherical, ellipsoidal, oblong or broadly cylindrical, unicellular,
pale to dark brown, verruculose or echinulate (Ellis, 1971).

113

Chapter 2: Taxonomy

Periconia minutissima Corda, Icones Fungorium Malayensium 1:19. 1837.


Cultural characters: Colonies growing fast, up to 5 cm diam. after one week,
greyish, cottony, flat, grey in centre, with colourless dense mycelium (HKUCC 3455,
3563).
Known hosts: On dead stems, leaves and other plant parts, usually close to or
on the ground (Ellis, 1971), on the dead culm of Dendrocalamus brandisii,
Neosinocalamus affinis (this study) and Trachycarpus fortunei (Taylor, 1998),.
Known distribution: China (Kunming) (this study), Cuba, Europe, Ghana,
Kenya, Lebanon, New Zealand, Pakistan, Sierra Leone, Sudan (Ellis, 1971),
Switzerland (Taylor, 1998), Tanzania and Zambia (Ellis, 1971).
Material examined: China, Yunnan, Kunming, Anlin, Yu Long Wan, on dead
culm of Neosinocalamus affinis, 19 Aug. 1998 Dequn Zhou (HKU(M) 9102);
International Gardening Exposition, on the dead culm of Dendrocalamus brandish, 28
Sept. 1998, Dequn Zhou (HKU(M) 9175); West Hill, on the dead culm of
Neosinocalamus affinis, 4 Oct. 1998, Dequn Zhou (HKU(M) 9176).
Phaeoacremonium W. Gams, Crous and M.J. Wingf, Mycologia 88: 789. 1996.
Crous et al. (1996) erected Phaeoacremonium with the type species P.
parasiticum (Ajello, Georg and C.J.K. Wang) W. Gams, Crous and M. J. Wingf. The
genus is intermediate between Acremonium and Phialophora. It is distinguished from
Phialophora by its aculaete conidiogenus cells and inconspicuous collarette, and from
Acremonium by its pigmented vegetative hyphae and conidiophores (Crous et al.,
1996). Six species are included in the genus (Crous et al, 1996). Species of
Phaeoacremonium have been associated with human infections and disease symptoms
of several woody hosts, including grapevine decline (Crous et al., 1996; Scheck et al.,
1998). Dupont, Laloui and Roquebert (1998) found an important divergence between
Phaeoacremonium species isolated from Vitis vinifera when they analysed partial
ribosomal DNA sequences.

114

Chapter 2: Taxonomy

Phaeoacremonium aleophilium W. Gams, Crous M.J. Wingf. and L. Mugnei,


Mycologia88:791. 1996.
Known hosts: Phyllostachys bambusoides (this study) and Vitis vinifera (Crous
etal, 1996).
Known distribution: China (Kunming) (this study), Italy and Yugoslavia (Crous
etal, 1996).
Material examined: China, Yunnan, Kunming, Yilang, on senescent culm of
Phyllostachys bambusoides, 25 June 1999, Dequn Zhou (HKU(M) 9365).
Remarks: Phaeoacremonium aleophilum is distinguished from P. parasiticum
and P. inflatipes by its conidial dimensions and smaller conidiophores (Crous et al.,
1996). The collections here are identified as P. aleophilum based on the similar
morphology and dimensions.
Phaeoacremonium parasiticum (Ajello, Georg and C.J.K. Wang) W. Gams, Crous
and M.J. Wingf, Mycologia 88: 794. 1996.
Cultural characters: Colony ca 3.5cm diam. on PDA after one month,
developed poorly and slow growing, blackish grey, flat and cottony, conidiophores and
conidia covering the entire colony surface.
Known distribution: China (Kunming) (this study), Tunisia and USA (Crous et
al, 1996).
Known hosts: Humans, Prunus armeniaca (Crous et al., 1996) and
Neosinocalamus affinis (this study).
Material examined: China, Yunnan, Kunming, An Lin, Qiu Mu Yuan, on
senescent culm of Neosinocalamus affinis, 5 Oct. 1998, Dequn Zhou (HKU(M) 9162);
ibid. (HKU(M) 9193).
Remarks: The collection is similar to Phaeoacremonium parasiticum (Crous et
al, 1996), but the conidia are slightly larger (4-5 x 2-3 urn vs. 3-4 x 1.5-2 urn).

115

Chapter 2: Taxonomy

Phaeoisaria Hohn., Sitzungsberichte, der Kaiserlichen Akademie der Wissenschaften


Mathematisch-Naturwissenschaftliche Abteilung 1. 118: 329. 1909.
Phaeoisaria was erected by von Hohnel (1909a) and is typified by P. bambusae.
Phaeoisaria species typically form synnemata, where the conidiophores are profusely
branched and form tufts (De Hoog and Papendorf, 1976). The conidia are small,
aseptate, and ellipsoid. There are about 11 species in Phaeoisaria differentiated
primarily by synnemata, conidiogenous cells and conidial characters and dimensions
(Ellis, 1971; Deighton, 1974; De Hoog and Papendorf, 1976). For keys to some
species refer to Ellis (1971).

Phaeoisaria clematidis (Fuckel) S. Hughes, Mycological Papers 56: 42. 1953.


Cultural characters: Colony 5 cm diam on PDA within one month, brownish
grey, and dark brown (reverse side), flat and flutty, with sparse aerial hyphae. The
conidiophores and conidia cover all of the colony surface (HKUCC 3557).
Known hosts: Common and widespread on fallen branches and sometimes dead
stems and leaves of many different plants (Ellis, 1971, Whitton, 2000), Bambusa
chungii, B. vulgaris var. vittata, Dendrocalamus bambusoides, Neosinocalamus affinis
and Sinobambusa tootsik (this study).
Known distribution: Argentina (de Hoog and Papendorf, 1976); China (Hong
Kong and Kunming) (this study), Cuba, Germany (Deighton, 1974; De Hoog and
Papendorf, 1976), Ghana, Java, Malaya, New Guinea, Sabah and Sierra Leone (Ellis,
1971).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa chungii, 9 Aug. 1998, Dequn Zhou (HKU(M) 9136); on
senescent culm of B. tuldoides, 19 Apr. 1999, Dequn Zhou (HKU(M) 9218); 17 June
1999, Dequn Zhou (HKU(M) 9294); ibid. (HKU(M) 9303); 29 July 1999, Dequn
Zhou (HKU(M) 9314); on the dead culm of B. vulgaris var. vittata, 8 Sept. 1998,
Dequn Zhou (HKU(M) 9110); ibid., on the dead culm of B. textilis, 6 June 1998,
Dequn Zhou (HKU(M) 8350); on senescent culm of Phyllostachys bambusoides, on 5
Aug. 1999, Dequn Zhou (HKU(M) 9335); Tai Lam Country Park, on the dead culm of
Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9127); China, Yunnan,

116

Chapter 2: Taxonomy

Kunming, West Hill, on the dead culm of Dendrocalamus bambusoides, 4 Oct. 1998,
Dequn Zhou (HKU(M) 9182); on the dead culm of Neosinocalamus affinis, 4 Oct.
1998, Dequn Zhou (HKU(M) 9179); Aniin, Qiu Mu Yuan, on the dead culm of
Neosinocalamus affinis, 19 Aug. 1998, Dequn Zhou (HKU(M) 9098); 5 Oct. 1998,
Dequn Zhou (HKU(M) 9192); Hot Spring, 5 Oct. 1998, Dequn Zhou (HKU(M) 9195);
on dead culm of Phyllostachys bambusoides, 25 June 1999, Dequn Zhou (HKU(M)
9284).
Remarks: The collections are identical to P. clematidis (Ellis, 1971) and the
species is very common on the bamoo hosts collected in Hong Kong and Kunming.
Phialogeniculata Matsush., Microfungi of the Solomon Islands and Papua New
Guinea: 45. 1971.
Matsushima (1971) erected the genus Phialogeniculata Matsushima based on
Phialogeniculata guadalcanalensis Matsush. to include taxa lacking setae, with
geniculate conidiophores bearing funnel-shaped collarettes and conidiogenous cells
proliferating sympodially. The conidia are hyaline, obclavate, septate and non-setulate.
Phialogeniculata guadalcanalensis was transferred to Dictyochaeta guadalcanalensis
by Kuthubutheen and Nawawi (1991) who used Dictyochaeta Speg. in a wide sense.
Two more species were described by Matsushima (1993), namely P. dimorpha
Matsush. and P. multiseptata Matsush. Hyde, Goh and Steinke (1998) described P.
africana Goh, K.D. Hyde and T.D. Steinke and pointed out that species of
Phialogeniculata are distinct from Dictyochaeta at least in the combination of the
following salient characters: setae lacking, sympodially proliferating, geniculate
conidiophores, and obclavate, septate, nonsetulate conidia.

Phialogeniculata dimorpha Matsush., Matsushima Mycological Memoirs 7: 60. 1993.


Known hosts: On senescent culm of Sinobambusa tootsik (this study) and on
rotten petiole of unidentified palm (Matsushima, 1993).
Known distribution: China (Hong Kong) (this study) and Peru (Matsushima,
1993).
Material examined: Hong Kong, New Territories, Tai Lam Country Park, on

117

Chapter 2: Taxonomy

senescent culm of Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9126).
Remarks: This species is similar to Phialogeniculata guadalcanalensis
Matsush., but the latter is monomorphic and has 1-septate conidia. The latter species
was transferred to Dictyochaeta guadalcanalensis (Mats.) Kuthubutheen and Nawawi
(Kuthubutheen and Nawawi, 1991). The collection is similar to Phialogeniculata
dimorpha, but the macroconidia are slightly narrower 3-4 um vs 1 4-6 urn)
(Matsushima, 1993) and septa are inconspicuous, as septa and macroconidia are both
hyaline.

Pleurophragmium Costantin, Materiaux pour l'histoire des champignons, II. Les


Mucedinees Simples: 100. 1888.
Pleurophragmium was established by Constantin in 1888 and typified by P.
simplex (Berk, and Broome) Hughes (Ellis, 1971). The genus includes 15 species
(Ellis, 1971; Hawksworth et ah, 1995).
Pleurophragmium simplex (Berk, and Br.) Hughes, Canadian Journal of Botany 36:
798. 1958.
Known hosts: On Arctium, Bambusa mutabilis, B. cornigera (this study),
Brassica, Conium. Epilobium, Filipendula, Heracleum, Neosinocalamus affinis (this
study), Polygonum, Sambucus and Urtica dioica (Ellis, 1971).
Known distribution: China (Hong Kong and Kunming) (this study), European
countries (Ellis, 1971) and UK (Dennis, 1986).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa mutabilis, 15 July 1998, Dequn Zhou (HKU(M) 9036);
on dead culm of B. tuldoides, 17 June 1999, Dequn Zhou (HKU(M) 9302); ibid.
(HKU(M) 9296); Sai Kung, To Kwa Pin, on dead culm of Bambusa cornigera, 27
June 1998, Dequn Zhou (HKU(M) 8389); China, Yunnan, Kunming, An Lin, Hot
Spring, on dead culm of Neosinocalamus affinis, 5 Oct. 1998, Dequn Zhou (HKU(M)
9194).
Remarks: The collections is similar to P. simplex except that the conidia are
wider (6-8 um vs. 3.5-6 um).

118

Chapter 2: Taxonomy

Podosporium Schweinitz, Transactions of the American Philosophical Society 4: 278.


1832.
Podosporium was established by Schweinitz (1832) to accommodate a single
species. P. rigidum Schw (Ellis, 1971). Ellis (1971) lectotypified P. rigidum (Chen and
Tzean. 1993). The genus comprises 13 species (Ellis, 1976; Hawksworth et al, 1995).
Dalisay (1998) observed P. elongatum and P. nilgirense on Bambusa and
Dendrocalamus in Hong Kong.
Podosporium elongatum Chen and Tzean, Mycological Research 97: 637. 1993.
Cultural characters: Colonies growing fast, ca. 5 cm diam. after one week,
white, creamy, waxy, underside creamy and mycelium sparse (HKUCC 3118).
Known hosts: Arundinaria hindsii (Dalisay, 1998), bamboo (Chen and Tzean,
1993). Bambusa sp., Bambusa tuldoides (Dalisay, 1998), B. chungii, B. mutabilis, B.
vulgaris var. Vittata, D. pulverulentus (this study), Dendrocalamus sp. (Dalisay, 1998)
and Melocanna baccifera, (this study).
Known distribution: China (Hong Kong) (Dalisay, 1998) and Taiwan (Chen and
Tzean, 1993).
Material examined: Hong Kong, New Territories, Yuan Luang, on the dead
culm of Dendrocalamus asper, 18 June 1998, Dequn Zhou (HKU(M) 8367); Tai Po
Kau Nature Reserve, on the dead culm of Bambusa vulgaris var. vittata, 25 June 1998,
Dequn Zhou (HKU(M) 8383); ibid. (HKU(M) 8384); on the dead culm of Bambusa
sp., 15 July 1998, Dequn Zhou (HKU(M) 9023); on the dead culm of B. mutabilis, 15
July, 1998, Dequn Zhou (HKU(M) 9027); on the dead culm of Bambusa chungii, 8
Sept. 1998, Dequn Zhou (HKU(M) 9130); on senescent of B. tuldoides, 19 Apr. 1999,
Dequn Zhou (HK(M) 9213); Sai Kung, on the dead culm of Dendrocalamus
pulverulentus, 27 June 1998, Dequn Zhou (HKU(M) 8391); ibid. (HKU(M) 8396);
ibid. (HKU(M) 9006); on the dead culm of Arundinaria hindsii, 21 July 1998, Dequn
Zhou (HKU(M) 9035); Wu Tong Village, on senescent culm of B. chungii, 18 May
1999, Dequn Zhou (HKU(M) 9229); Hong Kong, Hong Kong Island, Victoria Cape,
on senescent culm of Bambusa sp., 5 June 1998, Dequn Zhou (HKU M) 8342); No. 1
Pokfulam Reservoir, on the dead culm of B. chungii, 30 June 1998, Dequn Zhou

119

Chapter 2: Taxonomy

(HKU(M) 9014); on the dead culm of Melocanna baccifera, 30 June 1998, Dequn
Zhou (HKU(M) 9053); Victoria Cape, on the dead culm of Melocanna baccifera, 30
June 1998, Dequn Zhou (HKU(M) 9018).
Remarks: As Chen and Tzean (1993) pointed out, morphologically and
ecologically, Podosporium elongatum is closely related to P. nilgirense. They share a
common habitat and substratum, bamboo, and are also similar in general morphology
and the development of synnematous conidiomata, and conidial ontogeny. However, P.
elongatum differs markedly from P. nilgirense in having slender, elongate,
multiseptate, obclavate conidia (62-188 x 6-10 um, 8-21 septate). In contrast conidia
of P. nilgirense are broader, shorter (32-50 x 7-9 um) and 4-6 septate (Chan and Tzean,
1993).

Podosporium nilgirense (Subram.) M.B. Ellis, More Dematiaceous Hyphomycetes:


383. 1976.
Cultural characters: Colonies 5.5 cm. diam. after two weeks and slow growing
on PDA, white, creamy, cottony, effuse, with dentate edge, with concentric rings
(HKUCC 3471).
Known hosts: On the dead culms of bamboo (Ellis, 1976), on senescent culm of
Bambusa, Dendrocalamus (Dalisay, 1998), on the dead culms of Schizostachyum
dumetorum, Indocalamus longiauritus and Phyllostachys nidularia (this study).
Known distribution: China (Hong Kong) (this study) and India (Ellis, 1976).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa tuldoides, 19 Apr. 1999, Dequn Zhou (HKU(M) 9213); 17
June 1999, Dequn Zhou (HKU(M) 9254); ibid. (HKU(M) 9249); ibid. (HKU(M)
9293); ibid. (HKU(M) 9300); on senescent culm of Schyzostachyum dumetorum, 8
Sept. 1998, Dequn Zhou (HKU(M) 9106); ibid., Bamboo Garden, on the dead culm of
Bambusa beecheyana, 15 July 1998, Dequn Zhou (HKU(M0 9040); Wu Kau Tang, on
senescent culm of Indocalamus longiauritus, 18 May 1999, Dequn Zhou (HKU(M)
9234); Dao Moushan, on dead culm of Phyllostachys nidularia, 19 May 1999, Dequn
Zhou (HKU(M) 9235); Hong Kong Island, Pokfulam Country Park, on dead culm of B.
tuldoides, 5 June 1999, Dequn Zhou (HKU(M) 9246); Lung Fu Shan Country Park, on

120

Chapter 2: Taxonomy

the dead culm of Schizostachyum dumetorum, 19 July 1998, Dequn Zhou (HKU(M)
9042); ibid. (HKU(M) 9043); on senescent culm of Arundinaria hindsii, 11 May 1999,
Dequn Zhou (HKU(M) 9219).
Remarks: The conidia dimension of these collections overlap, but are slightly
longer than those of P. nilgirense (30-56 (am vs. 32-50 urn) (Ellis, 1976). The
conidiogenous cells and conidium walls in this species resemble those of
Didymobotryum verrucosum, but in the latter species the conidia are catenate and 1septate.
Ramichloridium Stahel ex de Hoog, Studies in Mycology 15: 59. 1977.
Ellis (1976) enlarged the concepts of the genera Pleurophramium Cost, and
Yeronaea Cif. and Montemart. to include the species described in Ramichloridium.
Yeronaea botryosa Cif. and Montermart., the type species of Yeronaea, is
characterized by dark, thick-walled conidiophores with pale, two-celled conidia. de
Hoog (1977) thought that Ellis' concept was impracticable and re-defined the concept
of Ramichloridium species with erect, dark conidiophores and predominantly onecelled conidia. He provided a key to 15 accepted species of Ramichloridium.
Ramichloridium indicum (Subram.) de Hoog var. microsporum D.Q. Zhou and K.D.
Hyde, var. nov.

(Fig 2.11.1)
l

Etymology: 'micro ' means small and sporum' means spores.


Cultural characters: Colonies being up to ca. 4 cm diam. after two weeks,
cottony, with aerial mycelia, effuse, dark brown or greyish brown, underside creamy
white (HKUCC 3147).
Conidiophores up to 150 long, 3-5 urn wide, erect, straight or flexuous, brown,
cicatrized at the apex. Conidia 2.5-4 urn diam., acropleurogenous, verruculose,
spherical or subspherical, with a slightly protruding hilum, hyaline to pale brown (Fig
2.11.1).
Material examined: Hong Kong, Hong Kong Island, Lung Fu Shan Country
Park, on dead culms of Arundinaria hindsii, 30 July 1998, Dequn Zhou (HKU(M)
9084, holotype).

121

Figs. 2.11.1. Line Drawing of Ramichloridium indicum var. microsporum (from (HKU(M)
9084, holotype). Appearance of conidiophores and conidia. Scale bar: 1 = 10 jam.

Chapter 2: Taxonomy

Remarks: Ramichloridium indicum is similar to R. apiculatum, however, the


latter species has ellipsoidal or obovoid conidia with a papillate base (R. indicum has
spherical or subspherical conidia) (Ellis, 1976; de Hoog, 1977). This specimen is
similar to R. indicum, but the conidia are conspicuously smaller (2.5-4 urn vs. 5-7.5
um). Ramichloridium indicum occurs on dead leaves of Phoenix and other palms in
India (de Hoog, 1977) but this fungus occurs on dead bamboo culms in Hong Kong.
Ramichloridium musae (Stahel ex M.B. Ellis) de Hoog, Studies in Mycology 15: 62.
1977.
Cultural characters: Colonies growing up to 5 cm diam. after one week, grey,
woolly, low convex, with entire edge, underside dark grey (HKUCC 3452).
Known hosts: Bambusa chugnii, B. mutabilis, B. vulgaris var. vittata (this
study), Musa sapientum, Musa sp. (Ellis, 1976) and Neosinocalamus affinis (this
study).
Known distribution: Bolivia, Cameroons, China (Hong Kong and Kunming)
(this study), Ghana, Guinea, Jamaica, Papua New Guinea, Sarawak, Sierra Leone,
Uganda (Ellis, 1976).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HKU(M) 9137); on
dead culm of B. mutabilis, 15 July 1998, Dequn Zhou (HKU(M) 9038); on the dead
culm of B. vulgaris var. vittata, 10 Sept. 1998, Dequn Zhou (HKU(M) 9109); China,
Yunnan, Kunming, West Hill, on dead culm of Neosinocalamus affinis, 4 Oct. 1998,
Dequn Zhou (HKU(M) 9178); Yiliang, on dead culm of Phyllostachys bambusoides,
27 June 1999, Dequn Zhou (HKU (M) 9366); 26 June 1999, Dequn Zhou (HKU(M)
9368).
Remarks: The collections are similar to R. musae (de Hoog, 1977), but conidia
are slightly wider (2.5-3 urn vs. 2-2.6 um).

122

Chapter 2: Taxonomy

Repetophragma Subram., Proceedings of Indian Natural Science Academy B58: 185.


1992.
Repetophragma was introduced by Subramanan (1992) with the type species R.
biseptata (M.B. Ellis) Subram. Repetophragma species are characterized by
acrogenous, solitary, euseptate, dry conidia, which are truncate, apical annellate and
conidiogenous cells are integrated. The genus has 9 species, most of them are
transferred from Sporidesmium (Subramanan, 1992).
Repetophragma subulata (Cooke and M.B. Ellis) Subram., Proceedings of Indian
Natural Science Academy B58: 185. 1992.
Known hosts: On bark of Liquidambar (Ellis, 1976) and on the dead culm of
Phyllostachys glauca and P. basihirsuta (this study).
Known distribution: China (Hong Kong) (this study) and USA (Ellis, 1976).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the dead culm of Phyllostachys glauca, 6 June 1998, Dequn Zhou (HKU(M) 8361),
Hong Kong Island, No.l Pokfulam Reservoir, on the dead culm of P. basihirsuta, 30
June 1998, Dequn Zhou (HKU(M) 9010).
Remarks: Repetophragma subulata resembles Sporidesmium jasminicola,
however, the latter has much shorter conidia and conidiophores are annellate
(Subramanian, 1992). These collections are identical to Repetophragma subulata
(Ellis, 1976), but conidiophores are slightly narrower (13-18 urn vs. 15-19 um).

Spadicoides S. Hughes, Canadian Journal of Botany 36: 827. 1958.


Spadicoides S. Hughes was established in 1958 with the type species S. bina
(Corda) S. Hughes (Ellis, 1971). Goh and Hyde (1996b) reviewed the genus and
discussed the similarities and differences between Spadicoides and Diplococcium
Grove. They accepted 21 species in Spadicoides. Two further species, namely S.
palmicola Goh and K.D. Hyde and S. heterocolorata (R.F. Castaneda, Guarro and
Cano) Goh and K.D. Hyde were added by Goh and Hyde (1998). Recently a new
species, Spadicoides bambusicola Zhou, Goh and K.D. Hyde was reported (Zhou et ai,
1999).

123

Chapter 2: Taxonomy

Spadicoides bambusicola D.Q. Zhou, Goh and K.D. Hyde, Fungal Diversity 3: 179.
1999.
This species has been formally published (see Appendix).
Spadicoides cordanoides Goh and K.D. Hyde, Mycologia 88: 1023. 1996.
Known hosts: Phyllostachys glauca (this study) and submerged decaying wood
(Goh and Hyde, 1996b).
Known distribution: Australia (Goh and Hyde, 1996b) and China (Hong Kong)
(this study).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of Phyllostachys glauca, 6 June 1998, Dequn Zhou (HKU(M) 8356).
Spadicoides xylogena (A. L. Smith) G.C. Hughes, Canadian Journal of Botany 35:
806. 1958.
Known hosts: decaying wood, Hypochnicium eichleri (Holubova-Jechova, 1982)
and Phyllostachys bambusoides (this study).
Known distribution: China (Kunming) (this study), Czechoslovakia, Poland and
UK (Holubova-Jechova, 1982).
Material examined: China, Yunnan, Kunming, Chengong, on senescent culm of
Phyllostachys bambusoides, 4 Oct. 1998, Dequn Zhou (HKU(M) 9187).
Remarks: The collection is identified as S. xylogena based on the key provided
by Goh and Hyde (1996b) and description provided by Holubova-Jechova (1982). The
conidial size falls within the range of S. xylogena, however, they are slightly shorter
(16-20 vs. 17-26 um) (Holubova-Jechova, 1982). Spadicoides xylogena overgrows
living carpophores of Hypochnicium eichleri and is hyperparasitic (Holubova-Jechova,
1982), but Wang (1976) gave a wider size range of the conidia (16-34 x 7-10.5 (j.m)
and did not mention the fungus overgrowing on living carpophores of Hypochnicium
eichleri when he studied the holotype.

124

Chapter 2: Taxonomy

Sporidesmium Link, Magazin Berlinische Gesellschaft Naturforschender Freunde


(Berlin) 3: 41. 1809.
Sporidesmium was introduced by Link with the type species S. atrum Link
(Ellis, 1971). Subramanian (1992) reassessed Sporidesmium and proposed the
diagnostic features of Sporidesmium ehrengergii (the type material of A. atrum is
missing). Sporidesmium has simple, septate conidiophores and the solitary, gangliar,
thick-walled, euseptate conidia. The conidiophore may proliferate percurrently to
produce further solitary conidia. Based on the euseptate or pseudoseptate nature of the
conidium and distinctive features of conidiophore proliferation, Subramanian (1992)
divided the heterogenus Sporidesmium into 7 genera, namely Sporidesmium Link,
Polydesmus Mount., Sporidesmiella Kirk, Stanjehughesia Subram., Repetophragma
Subram., Penzigomyces Subram. and Ellisembia Subram.

Sporidesmium ehrengergii M.B. Ellis, Mycological Papers 70: 63. 1958.


Known hosts: Neosinocalamus affinis (this study) and Tilia sp. (Ellis, 1976).
Known distribution: China (Kunming) (this study), India and USA (Ellis, 1976).
Material examined: China, Yunnan, Kunming, Anlin, Yu Longwan, on the dead
culm of Neosinocalamus affinis, 9 Aug. 1998, Dequn Zhou (HKU(M) 9096).
Remarks: The conidia of this collection are slightly narrower (9-11 um vs. 1012 urn) (Ellis, 1971).
Sporidesmium eucalypti M.B. Ellis and D. Shaw, Mycological Papers 72: 74. 1959.
Known hosts: Arundinaria hindsii (this study) and Eucalyptus sp. (Ellis, 1976).
Known distribution: China (Hong Kong) (this study) and New Guinea (Ellis,
1976).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on the dead culm of Arundinaria hindsii, 6 June 1998, Dequn Zhou (HKU(M) 8347).
Remarks: The conidia in this collection are slightly narrower (5-7 urn vs. 68.5um) (Ellis, 1976).

125

Chapter 2: Taxonomy

Sporidesmium eupatoriicola M.B. Ellis, Mycological Papers 70: 67. 1958.


Known hosts: On dead stems of Betula, Eilipendula, Eupatorium, and
Ochthocosmus (Ellis, 1971) and senescent culm of Bambusa textilis (this study).
Known distribution: China (Hong Kong) (this study) and Sierra Leone (Ellis,
1971).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa textilis, 6 June 1998, Dequn Zhou (HKU(M) 9048).
Remarks: The collection is similar to Sporidesmium eupatoriicola, but conidia
are slightly wider (9.5-12.5 um vs. 8-11 um) (Ellis, 1971).

Sporidesmium fragilissimium (Berk, and Curt.) M.B. Ellis, Mycological Papers 70:
55. 1958.
Known hosts: Dendrocalamus bambusoides and Neosinocalamus affinis (this
study) and Smilax sp. (Ellis, 1976),
Known distribution: China (Kunming) (this study) and USA (Ellis, 1976).
Material examined: China, Yunnan, Kunming, International Gardening
Exposition, Bamboo Garden, on the dead culm of Dendrocalamus bambusoides, 28
Sept. 1998, Dequn Zhou (HKU(M) 9170); Lunan, on the dead culm of
Neosinocalamus affinis, 30 Sept. 1998, Dequn Zhou (HKU(M) 9191);
Remarks: These collections are similar to S. fragilissimum (Ellis, 1976), but
conidia are slightly narrower (6-8 um vs. 8-9 um) (Ellis, 1976).
Sporidesmiumpenzigii M.B. Ellis, Mycological Papers 82: 45. 1958.
Known hosts: On rotten wood (Ellis, 1976) and Sinobambusa tootsik (this
study).
Known distribution: China (Hong Kong) (this study) and Java (Ellis, 1971).
Material examined: Hong Kong, New Territories, Tai Lam Country Park, on
the dead culm of Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9120).

126

Chapter 2: Taxonomy

Sporidesmium verrucisporum M.B. Ellis, Mycological Papers, 70: 57. 1958.


Cultural characters: Colonies slow growing, ca 6.5 cm. diam. after two months
on PDA, flat, fluffy, brown, mycelium superficial, sparse, septate and branched
(HKUCC 3547).
Known hosts: Arundinaria hindsii (this study) and Uvaria sp. (Ellis, 1976).
Known distribution: China (Hong Kong) (this study) and Sierra Leone (Ellis,
1976).
Material examined: Hong Kong, Hong Kong Island, Lung Fu Shan Country
Park, on the dead culm of Arundinaria hindsii, 30 July 1998, Dequn Zhou (HKU(M)
9078).

Stachybotrys Corda, Icones Fungorum 1:21. 1837.


Stachybotrys Corda was established by Corda (1837) with S. atra Corda as the
type species. It comprises 10 species (Jong and Davis, 1976; Hawksworth et al., 1995).
Stachybotrys species are characterized by macronematous conidiophores, with dark
granules and are terminated by a single whorl of discrete, clavate phialidic
conidiogenous cells. The conidia are produced in slimy masses, are dark in colour but
sometimes can be hyaline (Jong and Davis, 1976). Descriptions, illustrations and a key
to species were provided by Jong and Davis (1976). Recently Whitton (2000) reported
three new species on the Pandanaceae.
Stachybotrys dichroa Grove, Journal of Botany, British and Foreign (London) 24: 201.
1886.
Known hosts: On dead stems of herbaceous plants and occasionally on
branches of trees, hosts include Acanthus. Angelica, Carduus, Carlina, Coprosma,
Dactylis, Dipsacus, Epilobium. Eupatorium, Filipendula, Gunnera, Heracleum, Iris,
Oenanthe, Petasites, Sambucus, Senecio (Ellis, 1976) and Sinobambusa tootsik (this
study).
Known distribution: China (Hong Kong) (this study) and New Zealand (Ellis,
1971).
Material examined: Hong Kong, New Territories, Tai Lam Country Park, on

127

Chapter 2: Taxonomy

the dead culm of Sinobambusa tootsik, 8 Sept. 1998, Dequn Zhou (HKU(M) 9122).
Stachylidium Link, Magazin Berlinische Gesellschaft Naturforschender Freunde
(Berlin) 3: 15. 1809.
Stachylidium was introduced with 5. bicolor and S. terrestre Link, neither of
which was assigned as the type species. Hughes (1951) reviewed the genus and S.
bicolor was chosen as the lectotype. Hughes (1958) placed S. bicolor in synonymy
with S. verticillatum (Pers.) S. Hughes, based on priority. There are two species of
Stachylidium reported from bamboo: S. bicolor on Phyllostachys sp. in Taiwan
(Matsushima, 1980) and S. verticillatum on Phyllostachys pubescens in the former
USSR (Melnik and Popushoi, 1992).
Stachylidium verticillatum (Pers.) S. Hughes, Canadian Journal of Botany 36: 813.
1958.
Known hosts: Allium, Ananas (Ellis, 1971), Bambusa textilis (this study),
Dioscorea, Gardenia, Heliconia, Heracleum, Hibiscus, Maninot, Musa, Oenanthe,
Petasites, Phoenix, Populus, Pteridium, Sambucus, solanum, Sporobolus, Theobroma
and Urtica, Zea (Ellis, 1971).
Known distribution: Brunei (Whitton, 2000), China (Hong Kong) (this study),
Ghana, Malaya, New Guinea, Rhodesia, Sabah, Sierra Leone, Uganda (Ellis, 1971)
and UK (Hughes, 1951).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on culm of Bambusa textilis McClure, 10 Apr. 1998, Dequn Zhou (HKU(M) 8335).
Stilbella Lindau, Die naturlichen Pflanzenfamilien 1: 489. 1900.
Lindau (1900) erected the genus Stilbella to accommodate hyphomycetes
formerly attributed to Stilbum and transferred many other species to Stilbella. Stilbella
erythrocephala (Ditmar) Lindau was selected as lectotype by Clements and Shear
(Seifert, 1985). Nineteen species were accepted in the genus by Seifert (1985) and two
species added by Matsushima (1980) and Seifert (1990). There are 40 species
(Hawksworth et al., 1995).

128

Chapter 2: Taxonomy

Stilbella bambusae (Pat. and Gaillard) Seifert, Studies in Mycology 27: 92. 1985.
Telemorph: Sphaerostilbe bambusae (Seifert, 1985).
Known distribution: China (Hong Kong and Kunming) (this study), India,
Indonesia, Paraguay (Seifert, 1985), Philippines (Dalisay, 1998), Puerto Rico,
Venezuela and Vietnam (Seifert, 1985).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on senescent culm of Bambusa chungii, 8 Sept. 1998, Dequn Zhou (HKU(M) 9147);
Tai Lam, on dead culm of B. mutalis, 21 June 2000, Dequn Zhou (HKU(M) 9387);
China, Yunnan, Kunming, International Gardening Exposition, on dead culm of
Dendrocalamus brandisii, 28 Sept. 1998, Dequn Zhou (HKU(M) 9174); An Lin Hot
Spring, on senescent culm of Neosinocalamus affinis, 5 Oct. 1998, Dequn Zhou
(HKU(M) 9196); ibid. (HKU(M) 9259).

Tetraploa Berk, and Broome, Annales and Magazine of Natural History Series 2. 5:
459.1875.
Tetraploa was introduced by Berkley and Broome (1875) and typified by T.
aristata Berk, and Broome (Ellis, 1971). The genus Tetraploa is characterised by
micronematous

conidiophores, which are branched,

septate and

anastomosing. The conidiogenus cells are holoblastic, integrated

frequently
into the

conidiophores and typically intercalary. The conidia are brown, muriform, and
typically composed of four columns of 2-6 cells (Ellis, 1971). The genus comprises 12
species (Rifai, Zainuddin and Chilil, 1988; Hawksworth et ah, 1995; Whitton, 2000).
Tetraploa aristata Berk, and Broome, Annales and Magazine of Natural History Series
2.5:459(1850)
Teleomorph: Massarina tetraploa (Scheuer, 1991).
Known hosts: Cosmopolitan on various plants (Ellis, 1971; Whitton, 2000) and
Indocalamus sinicus (this study).
Known distribution: China (Hong Kong) (this study) and widespread (Ellis,
1971; Whitton, 2000).
Material examined: Hong Kong, New Territories, Sai Kung, Tu Kwa Pin, on

129

Chapter 2: Taxonomy

senescent culm of Indocalamus sinicus, 26 July 1998, Dequn Zhou (HKU(M) 9059).
Remarks: The collection is identified as Tetraploa aristata, but the conidia are
slightly longer (17.5-30 urn vs. 15-29 um) (Ellis, 1971).

Veronaea Cif. and Montemart, Am' isitituto Botanico Universita Laboratorio


Crittogamico 15: 68. 1957.
Veronaea Cif. and Montemart. was established and typified by V. botryosa Cif.
and Montem. (Ellis, 1971). Nine species have been reported in the genus (Ellis, 1971,
1976; Hawksworth et al., 1995). Veronaea is characterized by conidiophores which are
macronematous, monomenatous, the conidiogenous cells are polyblastic, integrated,
terminal, sympodial, cicatrized, scars are usually small and flat and conidia are solitary,
acropleurogenous, ellipsoidal or fusiform.
Veronaea coprophila (Subram. and Lodha) M.B. Ellis, More Dematiaceous
Hyphomycetes: 210. 1976.
This species has been formally published as a new record for Hong Kong (see
Appendix).
Veronaea parvispora M.B. Ellis, More Dematiaceous Hyphomycetes: 210. 1976.
Known hosts: Bambusa vulgaris var. vittata (this study), Quercus and
Chaetosphaeria callimorpha (Ellis, 1976).
Known distribution: China (Hong Kong) (this study) and UK (Ellis, 1976).
Material examined: Hong Kong, New Territories, Tai Po Kau Nature Reserve,
on dead culm of on the dead culm of Bambua chungii, 8 Sept. 1998, Dequn Zhou
(HKU(M) 9139); B. vulgaris var. vittata, 8 Sept. 1998, Dequn Zhou (HKU(M) 9117).
Remarks: These specimens are identical to V. parvispora, but conidia are
slightly wider (2-2.5 um vs. 1.5-2 um) (Ellis, 1976).

130

Chapter 2: Taxonomy

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Chapter 2: Taxonomy

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Chapter 3: Biogeographical Study

CHAPTER 3: BIOGEOGRAPHICAL
STUDY
ABSTRACT
Decaying samples of Phyllostachys bambusoides were ramdomly collected in
Hong Kong and Kunming during 1999 and 2000 and examined for fungi following
incubation. Fifty-nine fungal taxa, including 37 ascomycetes and 22 mitosporic fungi
were identified from the 300 samples from Hong Kong and 41 fungal taxa, including
29 ascomycetes, 1 basidiomycete and 11 mitosporic fungi were identified from the 300
samples from Kunming. Similarity indices and species richness of the fungal
communities on Phyllostachys bambusoides from each of the three sampling sites are
compared. The similarity indices among the fungal communities from different
sampling sites in Hong Kong or Kunming are high, but those between different
regions are low. The similarity indices among the fungal communities from the same
site in different sampling periods are also low. Fungal diversity index (ShannonWiener Index) in Hong Kong is 2.65, richer than that in Kunming (2.11). The species
richness in Hong Kong is 29.5, and is higher than that in Kunming (20.5). The
possible factors influencing fungal distribution, the limitations of this study are
discussed.

Key words: Distribution, Bray-Curtis coefficient, Phyllostachys bambusoides,


similarity indices.
3.1 INTRODUCTION
It is not possible to give a simple definition of biogeography, for the word has
been used in a number of different ways (Simmons, 1982). Biogeography is
commonly used for the study of the geographical distribution of plants, animals and
other organisms. Biogeographical studies have two main phases: description, in which
meaningful patterns in distributions are identified; and explanation, in which patterns
are established (Simmons, 1982). Recently some authors have defined biogeography
as a study of the distribution and the patterns of distribution of plants, animals and

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Chapter 3: Biogeographical Study

other organisms across the globe, on land, in the sea or in the air (Spellerberg and
Sawyer, 1999). Typically biogeographical studies try to answer such questions as: why
are certain species and certain groups of organisms found in certain localities and
nowhere else? What has caused these patterns on a global scale? Why is it that for
many groups of organisms there are fewer species in temperate as compared to tropical
regions? Why is it that in some regions there are few species, but the abundance of
some individual species is very high? The answers will help to reduce human negative
impact on the environment and can help us to sustainably manage the environment.
The geographical distribution of certain fungal groups in restricted regions is
well discussed. Zang (1981) reported on the classification and distribution of
Termitomyces in Yunnan, China. He found that T. albuminosus is widely distributed in
different localities from the south of the Yangtze River, especially in Yunnan province.
T. robustus is a typical tropical species. Ryabova and Tomilin (1980) observed that
fungal distribution was linked to host plant distribution. Gray (1983) studied the
distribution and habitats of nematophagous fungi in Ireland. He pointed out that
Arthrobotrys oligospora and A. musiformis both had a highly restricted distribution.
Tan and Wu (1986) surveyed the geographical distribution of 108 macrofungi from
subtropical evergreen, broad-leaves forests across China. Of these fungi, 17 species
were distributed widely in at least 5 districts and 24 species had a narrow distribution
in only 1 or 2 districts. They thought that climatic factors impacted on the distribution
of fungi. Recently, van Maanen and Gourbiere (1997) found that dematiaceous
hyphomycetes, such as Verticicladium trifldum and Thysanophora penicillioides
shared the same ecological niche in fungal successions on different substrata. Some
researchers have also made comparative studies of the mycota among different regions
and have tried to answer some of the above questions (Ho, 1998; Dalisay, 1998;
Frohlich and Hyde, 1999; Whitton, 2000; Taylor et al., 2000). Unfortunately, much of
the data from these studies are not supported by statistics and the authors did not
further analyze the possible impact factors.
In this study, the difference in fungal communities on Phyllostachys
bambusoides is compared between Hong Kong and Kunming, China, with statistical
analysis, since P. bambusoides is the only one bamboo distributed both in Hong Kong

147

Chapter 3: Biogeographical Study

and Kunming. Through comparison between the fungal communities in these two
regions, fungal composition will be analyzed and the possible main factors which
impact on the difference between the fungal communities in different geographical
locations will also be addressed.
3.2 MATERIALS AND METHODS
3.2.1 Location description
3.2.1.1 Hong Kong
Hong Kong is situated on the southeast coast of China, east of Guangdong
Province. It lies between latitudes 22 9' and 22 37' and longitudes 113 52' and 114
30', less than 192 km south of the Tropic of Cancer. Hong Kong is subtropical or can
be regarded as an intermediate "transitional tropics" zone (Dudgeon and Corlett, 1994).
Hong Kong has a subtropical monsoonal climate and is dominated by two monsoons.
During April to September there is a wet and warmer season due to prevalent
southwestern monsoon, and from October to the following March there is the dry and
cooler season due to the prevalent northeastern monsoon. In general, January and
August are the driest and wettest months respectively. The mean daily temperature
ranges from 28.8C in July to 15.8C in January. The seasons are separated by two
shorter periods of transitional weather (Dudgeon and Corlett, 1994; Government of the
Hong Kong SAR, 2000). The flora of Hong Kong is tropical, but less so than places
nearer the equator, with some extra-tropical plant families or genera better represented
in Hong Kong than in lowland tropics further south (Dudgeon and Corlett, 1994). The
fungal taxa from Hong Kong are characteristic of both temperate and tropical regions
(Taylor et al, 2000).

3.2.1.2 Kunming
Kunming, the capital of Yunnan province, China, is located in the central part
of Yunnan and lies between latitudes 102 10'-10340' and longitudes 24 23'-26 22',
with the Tropic of Cancer traversing the south part of the Province. The average
altitude is 1889.3 m above sea level in the Kunming area. Kunming has a subtropical
monsoonal climate and is dominated by two monsoons. The rainy season is from May

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Chapter 3: Biogeographical Study

to August and the dry season is from September to April. The mean annual
temperature ranges from 14-16C. The annual mean rainfall is 1015.1 mm. In general,
the mean temperature in January is 15.1C; in April is 18.2 C and in October, 15.1 C.
The mean annual relative humidity is 74%. Rainfall in spring (March to May) is 132.4
mm, being 13% of the total yearly rainfall; in summer (rainy season, June to August) it
is 609.2 mm, which is 60% of the total yearly rainfall; in autumn (September to
November), 238 mm, which is 23% of the total yearly rainfall; and in winter
(December to February), 36.4 mm, which is 4% of the total yearly rainfall (Project
group of New Survey of Yunnan Province, 1996).

3.2.2 Study sites and sample collection


3.2.1 Tai Po Kau Nature Reserve, Hong Kong, SAR, China
Tai Po Kau has 460 hectares of reserved area incorporating forest plantations
extending from the eastern slopes of Tso Shan (Grassy Hill) down to Tai Po Road. The
area has more than 100 different tree species, comprising a mixture of recent and long
established plantings. The dominant tree is Pinus massoniana and Machilus velutina
(Dudgeon and Corlett, 1994). There are also many bamboo species in Tai Po Kau,
such as Bambusa basihirsuta, B. beecheyana, B. chungii, B. cornigera, B. duriuscula,
B. glaucescens, B. malingensis, B. mutabilis, B. pervariabilis, B. textilis, B. tuldoides
and Dendrocalamus pulverulentus (But et al., 1985).
3.2.2.2 Little White Dragon Forest Park, Kunming, Yunnan, China
Little White Dragon Forest Park, Kunming, Yunnan covers 2,000 hectares and
is located about 40 kilometres east of Kunming. The forest is mainly composed of
Cunninghamia lanceolata, Pinus yunnanensis, P. armandii and Rhododendron sp.
Phyllostachys bambusoides is mixed with P. armandii on the slopes of a small valley.
The bamboo forest is in the vicinity of a resort village and has been to subject of heavy
human disturbance, because it is near to a camping and barbecue site.

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Chapter 3: Biogeographical Study

3.2.3 Methods and materials


3.2.1 Field collection
Three sites were selected each in Tai Po Kau, Hong Kong and Little White
Dragon Forest Park, Kunming. At each sampling site (20 x 20 m), 50 individual dead
bamboo plants of Phyllostachys bambusoides were randomly collected. Each sampled
dead bamboo culm was cut into ca 25 cm long pieces. Collections were carried out in
Kunming and Hong Kong on 5 and 8 July 1999, 2 and 5 September 2000 respectively.
3.2.3.2 Processing of samples
Samples were placed in zip-lock polythene bags on site and returned to the
laboratory. They were then incubated in polythene bags lined with moistened tissue at
room temperature {ca 23-25 C). After three days, the material was examined for
fungal fruiting bodies under a microscope. Every fungus was recorded and identified.
Data analysis
Numbers of species and individuals and the frequency of occurrence of each
species were recorded and calculated for each sampling site. The equations used are as
follows.

Equitability: it is most commonly expressed as Pielou's evenness index ( J')


(Ludwig and Reynolds, 1988):

Where

H = Shannon-Wiener Index;
Hmax = Log2S'
J' = 1 when the community structure is perfectly even (i.e. all taxa are
found in an equal number of samples) and J' = 0 (or close to 0) when
the community structure is at the extreme of uneven (i.e. only one taxon
present).
S' = species richness

150

Chapter 3: Biogeographical Study

Percentage frequency of occurrence

Shannon-Wiener index (diversity index): This index was used to calculate


diversity (Shannon and Weaver, 1949):

Where H = the probability of finding each taxon in a collection, and P, = the number
of individuals in the rth species. This index can also be calculated using the program
developed in conjunction with Prof. Wang.

Species similarity matrix: a matching triangular array of similarities between


every pair of species, in terms of patterns of occurrence across the samples. The
two most useful in ecology are the Bray-Curtis coefficient (Bray and Curtis, 1957)
and Sorensen's index.
Bray-Curtis coefficient: the similarity between the y'th and th samples was
calculated as follows:

Where yr = score (count or biomass) for rth species iny'th sample (i -,l,2,..,p;
j = l,2,...,n). The Bray-Curtis Coefficient was calculated using a computer

151

Chapter 3: Biogeographical Study

program developed in conjunction with Prof. Wang.

Species-area curve: This was also plotted to estimate the minimum number of
bamboos that was necessary to establish the community in any habitat (Huston,
1994).

Species richness (S'): the number of species present in any given area
(Spellerberg and Sawyer, 1999).

3.3 RESULTS
3.3.1 Species-area curves
The species-area curves reached asymptote at about 23-25 bamboo samples
(Figures 3.1-3.2). The majority of fungi can be found after examining 25 bamboo
plants, although the minimum number of bamboo plants required might vary from site
to site.
3.3.2 Fungal diversity
The most common group of fungi on Phyllostachys bambusoides were
ascomycetes (66 spp.) followed by mitosporic fungi (33 spp.). Only one basidiomycete
was collected (Table 3.3 and Figure 3.3). A total of 59 fungal taxa were identified in
Hong Kong (Tables 3.1-3.3), including 37 ascomycetes (representing 61% of the all
taxa) and 22 mitosporic fungi (37.3%). Forty-one taxa were identified on
Phyllostachys bambusoides in Kunming, including 29 ascomycetes (70.7% of the total
taxa), 11 mitosporic fungi (26.8%) and one basidiomycete (2.4%) (Tables 3.3 and
Figure 3.3).
The mycota can be categorized into three groups: common species in both
Hong Kong and Kunming; common species in Hong Kong or Kunming; and rare
species (Tables 3.1-3.2). Fungal communities generally consisted of a large number of
rare species and a few common species (Tables 3.1-3.2). The most common species in
Hong

Kong

and

Kunming

was Roussoella

hysterioides

(Tables

3.1-3.2).

Didymosphaeria futilis was most common in Hong Kong (Tables 3.1-3.2). Linocarpon

152

Chapter 3: Biogeographical Study

pandanicola was also common in Hong Kong in July 1999 and Ellisembia bambusae
in September 2000 (Tables 3.1-3.2). Anthostomella contaminans and Apiospora
sinensis were the most common species in Kunming. Astrocystis palmarum was also
common in 1999. Most other fungal species occurred infrequently in the two regions.
3.3.3 Ecological indices
Table 3.3 summarizes the species diversity of sampling sites in Hong Kong and
Kunming. The greatest species richness was recorded in Hong Kong (36 taxa) in 2000,
followed by Kunming (24 taxa) in 2000. The highest individual number of fungi was
found in Kunming in 1999 (266 individual records), followed by Hong Kong in 1999
(245 individual records). The fungal diversity index (Shannon-Wiener Index) in Hong
Kong is 2.65, higher than that in Kunming (2.11).

3.3.4 Similarity indices


Similarity indices of the fungal communities collected from different sites but
in the same region were high, while those of the fungal communities collected in
different regions were low (Tables 3.4-3.7). Similarity indices of the fungal
communities in the same region but collected at different periods were also low
(Tables 3.4-3.7). These results were also supported by calculation using Sorensen's
indices (the results are not presented here).

153

Chapter 3: Biogeographical Study

Table 3.1 Frequency of occurrence of fungi in Hong Kong and Kunming (1999)
Region and collecting time
Site No.

Hong Kong (8 July 1999)


Sitel

Site 2

Taxa

Site 3

Kunming (5 July 1999)

X,

Sitel

Site 2

Site 3

x2

Frequency of occurrence

Anthostomella contaminans

24

22

20

22

A. Jlagellariae

10

12

9.3

14

16

12

14
2

A. irregularispora

Apiospora sinensis

4.7

12

12

26

17

Arthrinium sp.

Aslrocyslis cocoes

10

14

11

A. palmarum

14

18

12

15

Astrosphaeriella Irochus

Coelomycete sp.

Didymosphaeha futilis

18

16

14

D. massarioides

Eutypeila sp.

12

9.3

Gilmaniella bambusae

H\ phomycete sp. 1

2.7

Linocarpon pandanicola

46

24

28

33

L. verminosum

5.3

Massarina cisti

M. eburnea

2.7

3.3

M. immersa

Massarina sp.

S'eolinocarpon inconspicuus

10

Oxydothis oraniopsis

Phaeoisaria clematidis

Rosellinia aequatohalis

Rosellinia sp.

2.7

90

64

52

69

Roussoella hystehoides

22

12

12

15

R. intermedia

10

14

10

10

6.7

10

14

10

11

R. puslulans

Spadicoides bambusicola

12

Spirodecospora bambusicola

4
21

20

16

19

14

13

12

13

Evenness

0.50

0.65

0.67

0.63

0.48

0.58

0.57

0.54

Mean frequency of occurrence

6.3

4.8

5.2

5.4

6.3

6.3

5.3

6.0

Shannon-Wiener Index

2.61

2.80

2.66

2.69

1.83

2.14

2.06

2.01

Species richness

154

Chapter 3: Biogeographical Study

Table 3.2 Frequency of occurrence of fungi in Hong Kong and Kunming (2000)
Region and collecting time
Site No.
Taxa
Acremonium sp.
Acrodictys bambusicola
Alternaria ahernata
Anlhoslomella contaminans
A.flagellariae
A. lugubris
Anthostomella cf. Nypensis
A. tomicum
Apiospora sinensis
A. montagnei
Astrocystis cocoes
Astrosphaeriella floridana
A. mauritiae
A. minoensis
A. s leilata
A. tornata
Crandallia bambusae
Didymosphaeria futilis
Discomycete sp.
Ellisembia bambusae
E. coronatum
E. fusiforme
E. leonense
E. pseudoseptata
Fusarium sp.
Gilmaniella bambusae
Hyphomycete sp. 2
Linocarpon livistonae
Massarina balnei-ursi
M. talae
M. walkeri
Niesslia sp.
Phaeoisaria clemalidis
Phoma sp.
Pleurophragmium simplex
Ramichloridium musae
Roussoella hysterioides
R. intermedia
R. pustulans
Spirodecospora bambusicola
Spohdesmium baccharidis
S. bonarii
S. ehrenbergii
S.flagellatum
S. fragillissimum
S. penzigii
Submersisphaeria bambusicola
Thyridium chrysomallum
Ustilago sp.
Xylariaceae sp.
Species richness
Evenness
Mean frequency of occurrence
Shannon-Wiener Index

Hong Kong (5 Sept 2000)


Sitel

Site 2

2
4
2
0
6
0
0
0
6
2
0
2
0
0
4
0
0
54
2
34
0
0
2
0
4
6
4
0
2
0
0
0
0
0
4
0
14
0
0
2
2
0
2
0
0
2
4
0
0
0
23
0.57
3.1
2.6

2
10
0
4
2
8
0
0
4
2
0
4
0
0
4
0
0
24
0
22
2
0
2
2
0
2
4
4
0
4
0
0
0
0
2
0
8
0
0
0
6
0
10
4
0
8
0
0
0
0
24
0.59
2.7
2.7

Kunming (2 Sept. 2000)

Site 3
Sitel
Site 2
*,
Frequency of occurrence
6
3.3
0
0
8.7
0
12
0
0
2
1.3
0
2
10
2
22
2.7
2
0
0
2.7
0
0
0
4
0
4
0
0
0
2
0
4
18
2
6
4
2.7
2
0
0
0
0
4
2
2.7
0
0
0
0
6
8
0
0
4
0
4
2
6
4
2
0
0
0
4
6
0
0
40
39.3
0
0
2
1.3
0
0
28
28
4
4
0
0.7
0
0
0.7
0
0
2
0
1.3
0
0
4
2
0
0
2
2
0
0
6
4.7
10
6
6
4.7
0
0
2
2
0
0
0
0.7
0
0
0
1.3
0
0
6
2
0
0
2
0.7
0
0
0
0
4
4
0
0
4
2
2
2.7
0
0
0
0
2
6
12
11.3
44
38
2
0.7
14
10
2
0.7
8
8
4
2
0
0
6
4.7
0
0
0
0
0
6
0
4
0
0
4
2.7
0
0
2
0.7
0
0
2
4
0
0
0
1.3
0
0
0
0
0
2
0
0
0
2
0
0
2
0
28
24.7
20
17
0.54
0.57
0.59
0.54
3.2
3.0
2.8
3.0
2.6
2.6
2.1
2.3

155

Site 3

x.

0
0
0
18
2
0
0
2
10
0
2
0
8
0
2
0
4
0
0
12
0
0
0
0
0
6
0
0
0
0
0
0
6
0
0
2
48
8
2
0
0
0
0
0
0
0
0
0
0
0
15
0.56
2.6
2.2

0
0
0
16.7
1.3
0
2.7
1.3
11.3
0.7
2
0
7.3
1.3
3.3
0.7
4.7
0
0
6.7
0
0
0
0
0
6
0
0
0
0
0
0
4.7
2
0
3.3
43.3
10.7
6
0
0
2
0
0
0
0
0
0.7
0.7
0.7
12.7
0.56
2.8
2.2

Chapter 3: Biogeographical Study

156

Chapter 3: Biogeographical Study

Table 3.3 Fungal species diversity in Hong Kong and Kuming


Hong Kong
Collecting time

Kunming

8/7/1999

5/9/2000

Ascomycetes

18

19

37

Basidiomycetes

Mitosporic fungi

17

Fungal numbers

245

Species/per sample

1.63

Evenness

0.59

0.50

Shannon Index

2.69

2.60

5/7/1999

2/9/2000

X **

18

13

16

29

14.5

0.5

22

11

11

5.5

227

472

236

266

210

476

238

1.51

/
/
/

1.57

1.77

1.40

0.55

0.49

0.48

0.49

2.65

2.01

2.20

/
/
/

X*

1.59

2.11

: X l = mean of fungi collected on 8 July 1999 and 5 Sept. 2000; X 2 = mean of fungi collected on 5 July 1999 and 2 Sept. 2000.

Table 3.4 Similarity analysis (Bray-Curtis Coefficient) of mycota identified from


different sampling sites in 1999
Areas and Collecting time
Site No.
Site 1
Hong Kong
(8 July 1999)

Hong Kong (8 July 1999)


Site 1

Kunming (5 July 1999)

Site 2

Site 3

Site 4

Site 5

Site 6

73

68

32

36

34

100

68

28

33

33

100

20

26

24

100

80

74

100

74

100

Site 2
Site 3
Site 4

Kunming
(5 July 1999)

Site 5

100

Site 6

Table 3.5 Similarity analysis (Bray-Curtis Coefficient) of mycota identified from


different sampling sites in 2000
Areas and collecting time

Hong Kong (5 Sept. 2000)

Kunming (2 Sept. 2000)

Site No.

Site 7

Site 8

Site 9

Site 10

Site 11

Site 12

Site 7

100

56

70

15

16

22

100

63

14

13

13

100

14

17

22

69

72

100

68

Hong Kong

Site 8

(5 Sept. 2000)

Site 9
Site 10

Kunming

Site 11

(2 Sept. 2000)

Site 12

100

100

157

Chapter 3: Biogeographical Study

Table 3.6 Similarity analysis (Bray-Curtis Coefficient) of mycota identified in


Hong Kong at different sampling periods
Collecting time

8 July 1999

8 July 1999

5 Sept. 2000

Site No.

Site 1

Site 2

Site 3

Site 7

Site 8

Site 9

Site 1

100

73

68

22

17

26

100

68

19

12

22

17

15

21

56

71

100

62

Site 2
Site 3

100

Site 7
5 Sept. 2000

100

Site 8
Site 9

100

Table3.7 Similarity analysis (Bray-Curtis Coefficient) of mycota identified in


Kunming at different sampling periods
Collecting time
Site No.
Site 4
5 July 1999

Site 5

5 July 1999
Site 4

Site 5

Site 6

Site 10

Site 11

Site 12

100

80

74

55

46

56

100

74

56

49

56

100

57

56

62

Site 6
Site 10
2 Sept. 2000

2 Sept. 2000

100

Site 11

71

78

100

72

Site 12

100

Fig. 3.3 Fungal diversity on Phyllostachys bambusoides


In Hong Kong and Kunming

*: The samples of Hong Kong 1 and Kunming 1 were collected from Hong Kong and Kunming on 8
and 5 July 1999 respectively; the ones of Hong Kong 2 and Kunming 2 were collected from Hong Kong
and Kunming on 5 and 2 Sept. 2000 respectively.

158

Chapter 3: Biogeographical Study

3.4 DISCUSSION
3.4.1 Similarity indices
Bray-Curtis coefficients and Sorensen's index are generally used in ecological
analysis (Mumby et al, 1996; Scheltema, 1997; Ho, 1998; Yang and Chen, 1998;
Ayyappan and Parthasarathy, 1999; Starling, 2000). In this study, only the results
calcualated by Bray-Curtis coefficient was applied to analyze similarities between the
fungal communities (Tables 3.4-3.7), because Bray-Curtis coefficient is probably a
better ecological analysis, as it calculates the similarity among communities and does
not only compare the similarity in fungal composition. It also compares the frequency
of occurrence of every species in the community (Bray and Curtis, 1957). The results
were also supported by calculation made by Sorensen's index (the results are not
presented here).

3.4.2 Biogeography of fungi on bamboo


The overall species diversity of plants in the tropics is greater than that of
temperate and polar regions (Barry and Moore, 2000). Stevens (1989) concluded that,
as with most other organisms, there was a latitudinal species richness gradient of
fungal taxa increasing towards the tropics. With comparison to Kunming, Hong Kong
is nearer to the tropics and therefore more fungi should occur in Hong Kong. Taylor et
al. (2000) attempted to verify Stevens' hypothesis by studying palm microfungi in
several tropical, subtropical and temperate countries or regions. Unfortunately Taylor
et al. (2000) confessed that the results were not conclusive due to the difficulty in
interpreting the diversity indices. In this study, the mycota on Phyllostachys
bambusoides were collected at almost the same time in Hong Kong and Kunming over
2 years. A total of 600 bamboo samples were examined and 65 fungal species from
903 fungal collections were identified. The species richness, evenness and ShannonWiener index of the fungal communities in Hong Kong were all found to be greater
than those in Kunming (Table 3.3). This indicates that the fungal diversity on
Phyllostachys bambusoides in Hong Kong was richer than that in Kunming. The
results of this study support the hypothesis proposed by Stevens (1989).
Taylor et al. (2000) found different fungal assemblages in association with

159

Chapter 3: Biogeographical Study

palms in temperate as compared to tropical regions. The community difference was


more related to climatic influences than to the hosts sampled, as few of the fungi were
believed to be host-specific. Frohlich and Hyde (1999) have also observed that the
distribution of fungi in the Old World Tropics follow patterns consistent with climate,
rather than past and present biogeography or host distribution. In this present study,
the fungal communities in Hong Kong and Kunming were collected from the same
bamboo host, i.e., Phyllostachys bambusoides. The sampling size, techniques and
sampling time were similar. The results are therefore consistent with those of Taylor et
al. (2000) and Frohlich and Hyde (1999). The difference between the fungal
communities in Hong Kong and Kunming may most probably result from different
climatic conditions. Climatic factors recorded in Hong Kong and Kunming in June,
July 1999, August and September 2000 are listed in Table 3.8.

Table 3.8 Comparison of climatic factors in Hong Kong and Kunming in July
1999 and September 2000*
Area

Hong Kong

Kunming

Time

Mean air

Mean relative

Total rainfall (mm)

temperature (C)

humidity (%)

June 1999

28.9

79

197.4

July 1999

29.2

81

203.8

Aug. 2000

26.6

81

391.4

Sept. 2000

27.8

78

365.7

June 1999

19.6

76

173.2

July 1999

19.7

74

204.8

Aug. 2000

19.1

70

205.9

Sept. 2000

17.5

67

121.6

Source: Hong Kong Observatory, 2000 and Kunming Climatic Observatory, 2000.

Air temperature and rainfall are probably the major climatic factors that can
have an impact on fungal diversity, as the relative humidity in Hong Kong and
Kunming is not very different (Table 3.8). Most fungi are mesophilic, and grow at
temperatures in the range of 5-35C, with optimum temperatures between 25 and 30 C
(Dix and Webster, 1995). As the collections were carried out on 5 and 8 July 1999 and
2 and 5 September 2000, temperatures during June 1999 and August 2000 therefore,

160

Chapter 3: Biogeographical Study

are more meaningful for fungal occurrence. Temperatures of June and July 1999 and
August and September 2000 in Hong Kong were in the optimum range for fungal
growth (27.8-29.2C), but this was not so in Kunming (17.5-19.7C) (Table 3.8). This
may account for the higher fungal diversity on Phyllostachys bambusoides in Hong
Kong than in Kunming.
Water is of paramount importance to fungi, in terms of the intra- and extracellular chemical reactions and solute transportation, the volume increase responsible
for extension growth, reproduction propagation, spore germination, and ejection of
propagules (Ayres and Boddy, 1986). The development of fungal communities is
greatly influenced by water content and relatively small changes can have dramatic
effects on fungal development (Dix and Webster, 1995). Rainfall may therefore have a
direct and positive impact on fungal diversity.
The plant diversity in the vicinity of sampling habitats may also affect the
diversity of fungi. Pirozynski (1981) pointed out that an enormous diversity of plant
species in an area is likely to support an equally diverse mycota. At Tai Po Kau Nature
Reserve, Hong Kong, there are more plant diversity than in Yang Zhonghai Forest
Park, Kunming, perhaps it supports at Tai Po Kau more fungal diversity.
Human disturbance may also be one of the factors influencing fungal diversity
(Tsui et al, 1998). Taylor, Hyde and Jones (2000) concluded that the degree of human
disturbance of a habitat was a factor influencing fungal diversity. In this study, the
sampling sites of Kunming were heavily disturbed by people, because the sites were
near to a camp and barbecue site in a Forest Park, but the Tai Po Kau site is in a nature
reserve and receives few visitors.

3.4.3 Temporal change of fungal communities


The similarity indices among the mycota in different sampling periods over
two years were low (Tables 3.4-3.7). This may be because the fungal communities
were dynamic on bamboo hosts and fungi replace one another as their dynamic
communities alter in space and time (Frankland, 1998).

161

Chapter 3: Biogeographical Study

3.4.4 Core group of fungi


Spellerberg and Sawyer (1999) thought that each species requires certain
conditions for its own growth and survival at any particular place. The presence of
some species or groups of species may therefore, be used to indicate the state of the
environment in which they occur. Wong and Hyde (2001) discussed the " core fungal
group" that occurred on grasses. They defined a 'core group' as a group of fungi
commonly associated with various hosts. Hyde and Goh (1997, 1998) identified 37-42
fungi in stream and lake systems in Australia. Most of the species identified were rare
and a small 'core group' of less than 6-10 species formed the dominant species. They
tested several of the 'core species' for their ability to degrade lignocellulose and
concluded that no single species is capable of producing all of the necessary enzymes
to degrade woody tissue and in nature it is likely that a consortium of fungi work
together in the wood decomposition process. It is therefore, probable that a 'core
group of fungi' plays a pivotal role in the degradation of woody tissue in a local
stream system and can be regarded as keystone species.
Based on the above discussions and the results of this study, I define a 'core
group of fungi' as a group, which occurs on certain hosts, genera or families with a
relatively high frequency. This is not a taxonomic group, but an ecological group. The
loss of this 'core group of fungi' due to environmental change or human disturbance
could have a consequential effect on substrata decomposition (Wong et al., 1998).
Several authors have also discussed 'core group of fungi'. Saxena et al. (1985)
found that Cephalosporium acremonium, Cunningnghamella echinulata, Epicoccum
indicum, Ravenelia acaciae-caesiae and Septoria achyranthidi were a core group of
extra embryal contaminants of the seeds of Acacia nilotica. Ho (1998) listed a
potential 'core group of fungi' (he used the term keynote fungal taxa) on submerged
wood in Brunei, Hong Kong, Malaysia, South Africa and UK. Tsui (1999) also found
that fungal communities in freshwater of Hong Kong comprised a 'core group' of a
few species. Furthermore, Wong and Hyde (2001) found that Alternaria alternata,
Annulatascus triseptatus, Cladosporium cladosporioides, Fusarium senitectum,
Nigrospora state of Khuskia oryzae, Ornatispora taiwanensis, Periconia echinochloae,
Petrakia paracochinansis, Phomopsis sp., Stachybotrys kampalensis and Tetraploa

162

Chapter 3: Biogeographical Study

aristata were a 'core group of fungi' on six monocotyledonous grasses in Hong Kong.
In this study, a putative 'core group of fungi' on bamboo is listed in Table 3.9.
These fungi occurred on Phyllostachys bambusoides with relatively high frequency
(frequency of occurrence > 10%) in Hong Kong and/or Kunming. This group probably
plays a crucial role in the decay of bamboo. If some or all of these fungi disappear, this
may impact significantly on decomposition.

Table 3.9 Core group of fungi on Phyllostachys bambusoides in Hong Kong and
Kunming
Regions

Hong Kong

Kunming

Hong Kong and Kunming

Fungi

Frequency occurrence (%)*

Didymosphaeria futilis

26.7

Ellisembia bambusae

28.0

Linocarpon pandanicola

33.0

Anthostomella contaminans

15.4

Apiospora sinensis

14.2

Astrocystis palmarum

15.0

Roussoella hysterioides

34.7

*: Frequency of occurrence of fungi here are mean value (calculated from Tables 3.1-3.2).

3.5 LIMITATIONS TO THIS STUDY AND FUTURE WORK


The data on fungal biogeography is only from one bamboo host, Phyllostachys
bambusoides. Future work should survey more bamboo hosts in Hong Kong and
Kunming and probably other regions.
Data on fungal biogeography were collected twice over two years. It would be
better if the collections were carried out more often in order to confirm which is the
"core group of fungi'.
The bamboos in Hong Kong and Kunming naturally died in 1997 and 1992
respectively (according to the local people). Fungal succession on the dead bamboo
may therefore have affected the fungal communities in these two regions. Future work
should be conducted on bamboos which are all at the similar age of decomposition.

163

Chapter 3: Biogeographical Study

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2. Ayyappan, N. and Parthasarathy, N. (1999). Biodiversity inventory of trees in a
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3. Barry, C.C. and Moore, P.D. (2000). Biogeography: an Ecological and
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6. Dalisay, T.U. (1998). Biodiversity of Microfungi Associated with Species of
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global fungal diversity estimates realistic? Biodiversity and Conservation 8: 9771004.
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Fungi on Wood Submerged in Tropical Streams. Ph.D. thesis, The University of
Hong Kong, Hong Kong. 229 pp.
H.Huston, M.A. (1994). Biological Diversity: the Coexistence of Species on

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Chapter 3: Biogeographical Study

Changing Landscape. Cambridge University Press, Cambridge, UK. 681 p.


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Mt. Levis, North Queensland, Australia. Muelleria 10: 145-157.
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North Queensland, Australia. Mycological Research 102: 739-749.
17. Ludwig, J.A. and Reynolds, J.F. (1988). Statistical Ecology: A Primer on Methods
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18. Mumby, P.J., Clarke, K.R. and Harborne, A.R. (1996). Weighting species
abundance estimates for marine resource assessment. Aquatic Conservation 6:
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19. Pirozynski, K.A. (1981). Interactions between fungi and plants through the ages.
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Yunnan province. Yunnan People's Publishing House, Kunming, China. 467 pp.
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communities in the Central Chernozem Reserve, Russian SFSR, USSR.
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28. Stevens, G.C. (1989). The latitudinal gradient in geographical range: how so many
species coexist in the tropics. American Naturalist 133: 240-256.
29. Tan, H.C. and Wu, R.J. (1986). The ecological and geographical distribution of
108 species of macromycetes from the subtropical, evergreen, broad-leaves forests
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30. Taylor, J.E., Hyde, K.D. and Jones, E.B.G. (2000). The biogeographical
distribution of microfungi associated with three palm species from tropical and
temperate habitats. Journal of Biogeography 27: 297-310.
31. Tsui, K.M. (1999). Biodiversity and longitudinal distribution of fungi on
submerged wood, with reference to human disturbance. Ph.D. thesis, The
University of Hong Kong. 307 pp.
32. Tsui, K.M., Fryar, S.C., Hodgkiss, I.J., Hyde, K.D., Poonyth, A. and Taylor J.
(1998). The effect of human disturbance on fungal diversity in the tropics. Fungal
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34. Whitton, S.R. (2000). Biodiversity Studies of Microfungi on the Pandanaceae.
Ph.D. thesis. The University of Hong Kong, Hong Kong. 625 p.
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166

Chapter 4: Host Recurrence and Biodiversity

CHAPTER 4: HOST RECURRENCE AND


BIODIVERSITY
ABSTRACT
Fifty-eight taxa from 887 individual fungal identifications were made from
bamboo in Hong Kong, including 43 ascomycetes and 15 mitosporic fungi. Forty-six
taxa from 430 individual fungal identifications were made from bamboo in Kunming,
comprising 26 mitosporic fungi and 20 ascomycete species. There were only a few
dominant species on the bamboo hosts, the majority of species encountered being rare.
The species richness, number of identifications and Shannon-Wiener indices between
the fungi collected at different sites in Hong Kong was higher than in Kunming.
Species diversity varied in relation to the substratum. Bambusa shiuyingiana (in Hong
Kong) yielded the richest diversity (25 spp.). The lowest diversity (12 spp.) was found
in Kunming on Phyllostachys hetroclata.
Key words: Bamboo, diversity, estimates, host exclusivity.
4.1 INTRODUCTION
Estimates of fungal diversity are of paramount importance when placing value
on threatened habitats. Valuations are needed in order to prevent unsustainable
exploitation of habitats, which may be due to disturbances such as overlogging,
urbanization, or pollution (Cannon, 1997a). Estimates of fungal numbers rely heavily
on the ratio of the unique fungi which have host specificity to vascular plants. This
ratio in turn depends on the percentage of host specific fungi among all fungi. This
percentage has been hypothesized to be in the region of 6:1, but the figure has not
been statistically tested in field studies (Hawksworth, 1991; Hyde 1996; Frohlich and
Hyde, 1999). Using, amongst other data, a ratio of 6 fungi being unique to each
vascular plant species, Hawksworth (1991) estimated that there were 1.5 million
fungal species worldwide. This estimate has prompted discussion among mycologists,
who have tried to dispute or provide evidence for these figures (May, 1991; Hammond,
1992; Rossman, 1994; Hyde, 1996; Cannon, 1997a, b; Huhndorf and Lodge, 1997;

167

Chapter 4: Host Recurrence and Biodiversity

Lodge, 1997; Wildman, 1997; Frohlich and Hyde, 1999). Since data on the ratio of
host specific fungi to each host are extremely important in any estimates of fungal
numbers, host specificity of fungi on bamboo is therefore investigated in this study.
Host specificity is defined as the relationship between a living plant and a
fungus, which is restricted to the host plant, but not other plants in the same habitat
(Holliday, 1998). Zhou and Hyde (2001, in press) considered host specificity to be an
unsuitable term for saprobes and therefore suggested that the terms host recurrence
and host exclusivity were more suitable. Host recurrence was defined as the frequent
occurrence of a saprobic fungus on a particular host, but with an infrequent occurrence
on other hosts in the same habitat. Host exclusivity was defined as the exclusive
occurrence of a saprobic fungus on a particular host, when other hosts in the same
habitat are also studied and the fungus is absent. Discussion concerning these terms
was provided (Zhou and Hyde, 2001 in press). There is much literature dealing with
host specificity of pathogenic fungi, endophytes and mycorrhizal fungi (Zhou and
Hyde, 2001 in press), but associations between saprobic fungi and plant hosts have
received less attention.
Most reports on bamboo fungi have dealt with taxonomy or pathology
(Candoussau, Katumoto and Sherwood, 1985; Boidin, Candoussau and Gilles, 1986;
Spooner and Candoussau, 1988; Candoussau et al., 1996; Kuai, 1996; Umali, Goh and
Hyde, 1998). Dalisay (1998), however, compared bambusicolous fungi from Bambusa
spp. and Dendrocalamus spp. in Hong Kong and the Philippines.
The present study was initiated in order to examine the fungal diversity of six
bamboo hosts in Hong Kong and Kunming, China. Whether the saprobic bamboo
fungi are exclusive or recurrent on these hosts is tested. The results will provide much
needed data for biodiversity estimates.

168

Chapter 4: Host Recurrence and Biodiversity

4.2 MATERIALS AND METHODS


4.2.1 Location (Hong Kong and Kunming, refer to chapter three)
4.2.2 Study sites
4.2.2.1 Tai Po Kau Nature Reserve, Hong Kong, China (refer to chapter three)
4.2.2.2 Maomao Valley Village, Kunming, Yunnan, China
Maomao Valley Village, Kunming, Yunnan, China is located 25 km to the west
of the City of Kunming. Maomao Valley Village comprises 30-40 households and is
surrounded by artificial forest comprising Fargesia yunnanensis, Neosinocalamus
affinis, Phyllostachys hetroclata and Pinus armandii. The former three species are
bamboo and were selected for sampling.
4.2.3 Materials
Bamboo samples were randomly collected in Hong Kong and Kunming on 18
Sept. 1999 and 16 Sept. 1999 respectively. Each sample of bamboo culm was cut into
ca 25 cm long sections and the numbers of samples examined are provided in Table
4.1.
Table 4.1 Bamboo hosts sampled in Hong Kong and Kunming
Bamboo
Bambusa shiuyingiana

Location

Site size (m)

Site nuniber

Sample No./per site

Hong Kong

20x20

50

Bambusa tuldoides

Hong Kong

20x20

50

Bambusa text ills

Hong Kong

20x20

50

Fargesia yunnanensis

Kunming

20x20

50

Neosinocalamus affinis

Kunming

20x20

50

Phyllostachys hetroclata

Kunming

20x20

50

4.2.3 Processing of samples


Bamboo samples were placed in zip-lock polythene bags on site and returned to
the laboratory where they were incubated individually in polythene bags lined with
moistened tissue at room temperature. After three days, the samples were examined
under a microscope for fungal fruiting bodies. Each fungus was recorded and
identified.

169

Chapter 4: Host Recurrence and Biodiversity

4.2.5 Data analysis


Numbers of species and individuals, and frequencies of occurrence of each
species were recorded and calculated for each sampling site. In this study, percentage
frequency of occurrence, Pielous' evenness, Bray-Curtis coefficients, species-curves,
species richness and Shannon-Wiener Indices were used to analyze the data (details in
Chapter three). Correlation coefficients were also used in this chapter to examine the
correlation between fungi at the two sites.

Correlation coefficient: similarity between jth and Mi samples is:

Where yy = score (count or biomass) for /th species iny'th samples (I =,1,2,..,p;
j = l,2,...,n). The correlation coefficient was calculated by a computer program
developed by Prof. Wang and me.

4.3 RESULTS
4.3.1 Species-area curves
The species-area curves of all sampling sites indicated that asymptote was
reached at between 19-25 bamboo samples. Although the minimum number of
bamboo samples required varied from site to site, the majority of fungi could be
identified after examining 25 bamboo culms (Figures 4.1^.6).

4.3.2 Fungal diversity


4.3.2.1 Hong Kong
Thirty-five taxa from 887 individual identifications of fungi were found on
Bambusa shiuyingiana, B. tuldoides and B. textilis in Hong Kong. Ascomycetes (24
spp.) were dominant over mitosporic fungi (11 spp.) (Figures 4.7 and Tables 4.2 and
4.4). Species richness was the highest on B. shiuyingiana, with 25 taxa identified,
followed by 19 species on B. tuldoides and 14 species on B. textilis. The number of

170

Chapter 4: Host Recurrence and Biodiversity

identifications of fungi on B. tuldoides (330) was the highest, followed by B.


shiuyingiana (322) and B. textilis (235) (Table 4.4).
Most of the bambusicolous fungi collected in Hong Kong were rarely
encountered species. Astrosphaeriella stellata was common on all three bamboo hosts.
Roussoella hysterioides and R. pustulans were also common to all three bamboo hosts,
but their frequencies of occurrence were relatively lower. Astrosphaeriella
fissuristoma frequently occurred on Bambusa shiuyingiana, Anthostomella rehmii and
Podosporium nilgirense were common on Bambusa tuldoides. Arecophila bambusae,
Gilmaniella bambusae and Podosporium elongatum were common on Bambusa
textilis (Table 4.2).

4.3.2.2 Kunming
Twenty-nine taxa from 430 individual identifications of fungi were identified
from Fargesia yunnanensis, Neosinocalamus qffinis and Phyllostachys hetroclata. The
species diversity for each host is listed in Tables 4.3 and 4.5. Mitosporic fungi (18 spp.)
were generally dominant over ascomycetes (11 spp.). Species richness was highest on
F. yunnanensis and N. qffinis, each with 17 taxa, followed by 12 species on
Phyllostachys hetroclata. The number of identifications of fungi on F. yunnanensis
(292) was the highest, followed by N. affinis (74) and P. hetroclata (64) (Table 4.5).
The fungi identified on these three bamboo hosts were mostly rarely
encountered species. Gilmaniella bambusae commonly occurred on all three bamboo
hosts (Table 4.3). Astrocystis cocoes, Pestalotiopsis uvicola and Roussoella
hysterioides were dominant on F. yunnanensis and P. hetroclata was dominated by
Apiospora sinensis. Phaeoacremonium parasiticum was dominant on N. affinis (Table
4.3).

171

Chapter 4: Host Recurrence and Biodiversity

172

Chapter 4: Host Recurrence and Biodiversity

173

Chapter 4: Host Recurrence and Biodiversity

Table 4.2 Frequency of occurrence (%) of fungi on Bambusa in Hong Kong


Host name

Bambusa
shiuyingiana

Bambusa
textilis

Bambusa
tuldoides

Sampling sites
Fungi

Silel 1 Site2 1 Sitel

Site2 1 Site3 1 Site}

Silel 1 Site2 1 Site3

No.

Frequency of occurrence

/
/

Ascomycetes

Anthostomella belalongensis

3.0

11.6

12.2

14.1

4.5

A- contaminans

5.1

5.5

1.1

5.3

1.9

A. Jlagellariae

2.3

2.9

3.8

2.4

1.3

A. rehmii

1.0

55

30.4

27.8

31.3

1.1

108

A. sepelibilis

7.1

5.0

106

2.5

Apiospora sinensis

1.6

2.2

0.4

Arecophila bambusae

18.6

27.5

20.0

7.3

Astrocystis cocoes

3.0

3.1

2.1

0.9

Aslrosphaeriella bakeriana

1.4

2.4

0.4

10

A. desmonci

2.8

5.0

09

11

A. fissuristoma

35.4

32.0

2.7

1.0

29.5

6.3

11.9

12

A cf immersa

2.3

0.8

0.3

13

A. maculans

2.0

2.7

1.1

1.1

0.8

14

A stellala

10.1

63

12.5

23.3

13.3

9.5

20.0

15.0

20.0

14.4

15

A. uberina

2.0

0.8

2.3

1.1

0.7

16

Massarina desmonci

1.3

0.1

17

M. eburnea

2.2

1.6

0.4

18

Massarina sp.

1.0

1.6

2.1

0.5

19

Niesslia sp.

0.9

2.2

7.8

1.2

20

Rosellinia sp.

3.0

0.8

4.2

09

21

Roussoella hysterioides

30

2.3

1.6

10.1

6.3

6.3

5.7

88

35

5.3

22

R. pusmlans

14.1

26.6

36

2.2

1.6

22.1

4.3

6.3

2.4

9.2

23

Thyridium chrysomallum

1.4

1.3

2.4

0.7

24

Discomycetes sp. 1

2.7

0.3

25

Discomycetes sp.2

4.5

1.1

0.6

Coelomycetes (mitosporic fungi)

26

Hyalopycnis blepharistoma

1.4

2.5

0.4

Hyphomycetes (mitosporic fungi)

27

Acrodictys bambusicola

2.0

3.1

5.4

3.3

1.6

1.7

28

Arthrinium phaeospermum

2.7

1.1

3.9

0.9

29

GUmaniella bambusae

1.0

1.6

15.7

10.0

16.5

5.0

30

Janetia synnematosa

1.0

0.8

2.1

0.4

31

Pleurophragmium complex

3.0

0.3

32

Podosporium elongatum

7.1

3.9

3.3

95

18.6

12.5

20.0

8.3

33

P nilgirense

3.0

23

17.0

17.8

16.4

2.1

1.3

6.7

34

Sporidesmium bonarii

2.3

03

35

5. penzigii

36

1.1

0.8

0.6

/
/
/

Mean Frequency of occurrence

5.5

5.9

7.3

7.3

6.5

6.7

8.3

7.6

10.0

7.2

Species richness

18

17

14

15

16

15

12

14

10

146

Evenness

0.54

0.53

0.59

0.54

0.56

0.55

0.58

0.61

0.59

0.57

Shannon-Wiener Index of mycota

2.26

2.17

2.25

2.11

2.25

2.13

2.08

2.31

1.95

2.20

174

Chapter 4: Host Recurrence and Biodiversity

Table 4.3 Frequency of occurrence (%) of fungi on bamboo in Kunming


Host name

Fargesia

Neosinocalamus

Phyllostachys

yunnanensis

afflnis

hetroclata

Sitel | Site2 | Sitel

Site2 | Site3 | Site3

Sitel 1 Site2 1 Site3

No
Sampling sites

Fungi

Frequency of occurrence

Ascomycetes

Anthostomella contaminans

2.2

2.8

4.7

2.0

1.1

1.0

A. Jlagellariae

1.1

1.8

3.2

0.7

Apiospora sinensis

0.9

286

30.2

45.1

5.9

12.3

Aslrocystis cocoes

16.0

101

2.0

24.0

58

Astrosphaeriella venezuelensis

7.3

7.1

1.6

Eurypella gliciridiae

2.2

09

0.3

Roussoella hysterioides

24.0

32.1

7.1

7.0

5.9

23.0

11.0

R. intermedia

3.3

6.4

2.3

2.0

7.5

2.4

R. pusrulans

6.7

4.5

1.1

2.9

1.7

10

R. saccardii

3.6

2.9

0.7

11

Spirodecospora bambusicola

0.9

2.2

10.7

11.8

5.7

3.5

Coelomycetes (mitosporic fungi)

12

Hyalopycnis btepharistoma

3.3

0.4

13

Pestalotiopsis uvicola

22.0

19.3

19.0

6.7

Hyphomycetes (mitosporic fungi)

14

Arthrinium phaeospermum

33

3.7

11.9

7.0

5.9

2.2

38

5.9

2.9

5.2

15

Arlhrinhim sp.

1.1

3.2

0.5

16

Cordelia johnstonii

2.3

4.7

1.1

10.7

5.9

8.6

3.7

17

Ellisembia bambusicola

2.3

2.3

2.0

2.2

1.0

18

Gilmaniella bambusae

14.0

9.1

28.6

34.9

21.6

11.0

14.3

20.6

14.3

18.7

19

Hysterographium mori

2.0

2.3

2.0

0.7

20

Pezigomycetes uapacae

7.1

29

1.1

21

Periconia minutissima

3.6

2.9

5.7

1.4

22

Phaeoacremonium parasiticum

14.3

20.6

17.1

5.8

23

Phaeoisaha clematidis

10.7

5.9

14.3

3.4

24

Pleurophragmium simplex

7.1

5.9

8.6

2.4

25

Ramichloridium musae

3.6

2.9

0.7

26

Sporidesmium ehrenbergii

2.9

0.3

27

S. fragillissimum

2.9

2.9

0.6

28

S. uapacae

7.1

4.7

11.8

2.6

29

Stilbella bambusae

7.1

8.8

2.9

2.1

Mean frequency occurrence

8.2

8.2

11.2

95

10.0

7.7

8.0

8.3

6.8

8.7

Species richness

12

12

10

10

14

12

12

14

11.6

Evenness

0.S6

0.S8

0.54

0.54

0.48

0.57

0.65

0.64

0.63

0.58

Shannon-Wiener Index ofmycota

1.99

2.08

1.63

1.79

1.60

2.16

2.34

2.29

2.39

2.03

175

Chapter 4: Host Recurrence and Biodiversity

Fig. 4.7 Taxonomic groups of fungi on Bambusa spp. in Hong Kong

Fig. 4.8 Taxonomic groups of fungi on bamboo hosts in Kunming

176

Chapter 4: Host Recurrence and Biodiversity

Table 4.4 Summary of species diversity on 3 hosts from Hong Kong


Bambusa
shiuyingiana
Ascomycetes
Mitosporic fungi
Species richness
Individual numbers
Species/per sample
Evenness
Shannon-Wiener Index

Bambusa textilis

20
5
25
322
0.46
0.47
2.16

Bambusa
tuldoides

10
4
14
235
0.28
0.56
2.13

13
6
19
330
0.32
0.48
2.02

13.7
5
19.3
295.7
0.35
0.50
2.10

43
15
/
887
/
/
/

Table 4.5 Summary of species diversity on 3 hosts from Kunming


Fargesia
yunnanensis
Ascomycetes
Mitosporic fungi
Species richness
Individual numbers
Species/per sample
Evenness
Shannon-Wiener Index
of the mycotas

Neosinocalamus

afflnis

Phyllostachys
helroclata

I
20
26

10
7
17
292
0.34
0.50

4
13
17
74
0.23
0.59

6
6
12
64
0.19
0.49

6.7
8.7
15.3
143
0.25
0.53

2.03

2.41

1.77

2.07

/
430

/
/
/

4.3.3 Similarity among the mycota between different bamboo hosts


Tables 4.6-4.9 show that the similarity indices between the mycota from each of
the three sites on the same bamboo host in Hong Kong or Kunming are high. The
Pearson coefficient between the mycota from each of the three sites is also high
(Tables 4.10-4.11). The similarity indices between fungal communities on different
bamboo hosts are however low (Tables 4.4-4.7). The Pearson correlation indices are
also low (Tables 4.8-4.9). The sites in Hong Kong or Kunming were located in the
same forest area and macro-ecological conditions were similar.

177

Chapter 4: Host Recurrence and Biodiversity

Table 4.6 Similarity analysis (Bray-Curtis coefficient) of mycota on bamboo in


Hong Kong
Bambusa

Bambusa textilis

Bambusa tuldoides

shiuyingiana
Sitel

Bambusa

Site 2

shiuyingiana

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

100

76

81

25

33

23

29

28

26

100

80

18

26

16

25

24

22

100

30

38

26

22

25

22

100

78

90

19

22

17

100

70

30

28

28

22

23

20

100

77

78

100

79

Sitel

Bambusa textilis

Site 2
Site 3

100

Sitel

Bambusa

Site 2

tuldoides

Site 3

100

Table 4.7 Similarity analysis (Bray-Curtis coefficient) of mycota on bamboo in


Kunming
Neosinocalamus

Fargesia

Sitel

Fargesia

Site 2

yunnanensis

Site 3

100

Site 2

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

78

76

17

18

23

26

28

26

100

78

14

15

22

22

26

23

100

16

16

24

23

25

23

100

71

74

21

38

20

100

65

23

39

24

100

24

36

16

100

87

76

100

73

Sitel

Neosinocalamus

Site 2

afflnis

Site 3
Sitel

Phyllostachys

Site 2

hetroclata

Site 3

hetroclata

afflnis

yunnanensis
Sitel

Phyllostachys

100

178

Chapter 4: Host Recurrence and Biodiversity

Table 4.8 Similarity analysis (correlation coefficient) of mycota on bamboo in


Hong Kong
Bambusa

Bambusa textilis

Bambusa tuldoides

shiuyingiana
Sitel

Bambusa

Site 2

shiuyingiana

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

.914

.949

.010

.213

.083

.099

.091

.027

.952

.040

.162

.003

.144

.080

.061

.168

.266

.132

.065

.084

-.001

.906

.984

-.004

-.039

-.017

.867

.204

.127

.174

.008

-.028

-.010

.920

.940

.933

Sitel

Bambusa textilis

Site2
Site 3
Sitel

Bambusa tuldoides

Site 2
Site 3

Table 4.9 Similarity analysis (correlation coefficient) of mycota on bamboo in


Kunming
Fargesia

Neosinocalamus
afflnis

yunnanensis
Sitel

Fargesia

Sitel

yunnanensis

Sitel

Sitel

afflnis

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

.920

.915

-.141

-.011

-.098

.211

.226

.085

.880

-.203

-.089

-.143

.167

.171

.066

-.186

-.073

-.125

.120

.136

.031

.831

.785

.164

.460

.134

.779

.207

.487

.191

.027

.294

-.036

.976

.912

.882

Sitel

Phyllostachys

Sitel

hetroclata

Site 1

hetroclata

Sitel

Sitel

Neosinocalamus

Phyllostachys

179

Chapter 4: Host Recurrence and Biodiversity

Table 4.10 Pearson correlation of saprobic fungi on bamboo in Hong Kong


Bambusa
Sitel

Bambusa

Site 2

shiuyingiana

Site 3

shiuyingiana

Bambusa

textilis

tuldoides

Site 2

Site 3

Sitel

Site 2

Site 3

Sitel

Site 2

Site 3

1.000

.922**

.948**

.169

.129

.144

.150

.149

.082

1.000

.959**

.110

.111

.072

.194

.147

.115

1.000

.226

.199

.183

.115

.122

.044

1.000

.919**

.988**

.062

.243

.066

1.000

.891**

.037

.169

.037

1.000

.054

.230

.059

1.000

.943**

.956**

1.000

.940**

Site 2
Site 3
Sitel

Bambusa

Bambusa

Sitel

Sitel

Bambusa

textilis

tuldoides Site 2
Site 3

1.000

Correlation is significant at the 0.01 level (2-tailed), n = 35.

Table 4.11 Pearson correlation indices of saprobic fungi on lamboo in Kunming


Fargesia
Sitel

Fargesia

Sitel

yunnanensis

Sitel
Sitel

Neosinocalamus

Sitel

afflnis

Sitel

yunnanensis

Sitel

Site 2

Site 3

1.000

.924**
1.000

1.000

Neosinocalamus
Sitel

Site2

.931**

.20

.894**

-.013

Phyllostachys
hetroclata

Sitel

hetroclata

Site 3

Sitel

Site 2

Site 3

.106

.034

.271

.288

.136

.054

-.009

.245

.251

.118

.013

.076

.001

.204

.222

.103

1.000

.867**

.913**

.259

.099

.093

1.000

.844**

058

.037

.176

1.000

.245

.085

.091

1.000

.976**

.870**

1.000

.960**

Sitel
Sitel

afflnis Phyllostachys

1.000

**: Correlation is sisni! icant at t he 0.01 1 :vel (2-ta iled); n == 29.

180

Chapter 4: Host Recurrence and Biodiversity

4.4 DISCUSSION
4.4.1 Fungal diversity
The fungal communities on bamboo hosts in Hong Kong were dominated by
ascomycetes, followed by mitosporic fungi. This was also apparent in the literature
review (chapter one), the biodiversity survey (chapter two) and biogeographical study
(chapter three) on bambusicolous fungi. Dalisay (1998) and Leung (1998) have also
reported that ascomycetes were dominant over other taxonomic groups on Bambusa
spp.
The fungal diversity (Shannon-Wiener Index) on different bamboo hosts varied
from 2.02 to 2.16 (Hong Kong) and from 1.77 to 2.41 (Kunming). The average of the
indices on each bamboo host in Hong Kong was 2.10 and in Kunming was 2.07.
Species richness on bamboo hosts also varied, from 14 to 25 (Hong Kong), and from
12 to 17 (Kunming). The average is 19.3 (Hong Kong) and 15.3 (Kunming).
Individual identifications of fungi in Hong Kong were much higher than in Kunming
(Tables 4.4-4.5). Bambusa shiuyingiana from Hong Kong yielded the richest fungal
diversity (25 spp.) and the poorest diversity (12 spp.) was found on Phyllostachys
hetroclata in Kunming. The highest fungal identifications were found on Bambusa
tuldoides (330 identifications) and the lowest on P. hetroclata (64 identifications).
The fungal diversity on the bamboo hosts in Hong Kong was richer than that in
Kunming. This may result from climatic conditions, especially air temperature and
rainfall. The Hong Kong climate may be more favorable for fungal development than
that of Kunming (for details refer to Chapter three). Stevens (1989) pointed out there
was a latitudinal species richness gradient of fungal taxa increasing towards the tropics
and Hong Kong is nearer to the tropics than Kunming. The difference in fungal
diversity and species richness may also be affected by the hosts examined and fungi
themselves, such as the fungal ability to utilize different nutrients in the hosts (Boyd
and Hess, 1970; Carroll and Petrini, 1983), differentiation of host nutrients (Mason
and Bryant, 1975), and physical structures of hosts (Mason and Bryant, 1975;
Michaelides and Kendrik, 1978; Akanil and Middleton, 1997). Wong (2000) found
that woody grasses supported a higher diversity of fungi than those with soft tissues,
perhaps due to the longer time needed for the woody tissues to decay.

181

Chapter 4: Host Recurrence and Biodiversity

A latitudinal species richness gradient of fungal diversity increasing towards


the tropics was also observed by Dalisay (1998) in Hong Kong and the Philippines.
She identified 50 fungal species, including 36 mitosporic fungi and 14 ascomycetes in
Hong Kong, and 56 fungal species, including 42 mitosporic fungi, 12 ascomycetes and
1 basidiomycete in the Philippines on equal numbers of Bambusa and Dendwcalamus
samples. Species richness of fungi on the bamboo hosts in the Philippines was slightly
higher than in Hong Kong, while the number of fungal collections in the Philippines
were much higher (1320) than in Hong Kong (777). Dalisay (1998) also pointed out
that the differences in diversity in the two regions might be influenced by climate. The
Philippines is in the tropics, while Hong Kong is in the intermediate "transitional
tropics" (Dudgeon and Corlett, 1994).
Other studies have also shown an increase in fungal diversity towards the
tropics. Taylor et al (2000) investigated biodiversity and distribution of fungi
associated with three ecologically different palms, Archontophoenix alexandrae,
Cocos nucifera and Trachycarpus fortunei in tropical and temperate countries/regions.
Fungal diversity (Shannon diversity index) in Hong Kong was much higher than in
Hubei, China and Western Europe. Shannon diversity index of the fungi on palms was
higher in the tropical region (46.7), as compared to the temperate regions (17.2) within
Australia (Taylor et al, 2000). The fungal diversity in aquatic environment also
increased nearer to the tropics (Ho, 1998; Tsui et al., 2000).
This present study revealed that the fungal communities on bamboo hosts in
Hong Kong and Kunming mainly comprised rare taxa (91) and a few dominant ones
(13) (Table 4.12). This result is consistent with previous investigations of biodiversity
of fungi on various substrates, such as on red spruce (Bills et ah, 1986), on leaf litter
(Bills, 1994; Polishook et al, 1996;), on submerged wood (Ho, 1998; Tsui et al, 2000)
and on palms (Frohlich and Hyde, 1999).
Bamboos appear to have a lower fungal diversity, as compared to some other
tropical or subtropical monocotyledonous plants. Dalisay (1998) collected 83 fungal
taxa on Bambusa sp. and Dendwcalamus sp. in Hong Kong and the Philippines. In
this thesis, I have identified 120 fungal taxa on 34 different bamboo hosts in Hong
Kong and Kunming. Poon and Hyde (1998) investigated vertical distribution of

182

Chapter 4: Host Recurrence and Biodiversity

saprobic fungi on senescent Phragmites australis culms at Mai Po Marshes, Hong


Kong. They identified 58 fungal taxa at only two sampling times. Frohlich and Hyde
(1999) identified 189 fungal species, including 49 ascomycetes, 3 basidiomycetes, 131
mitosporic

fungi,

zygomycetes

and

ascomycetes/mitosporic

fungi

(anamorph/teleomorph) connections, only on 6 palm trees in Australia and Brunei.


Taylor et a! (2000) found 287 fungal taxa on three palm hosts. Wong (2000) also
collected 215 fungal taxa, including 61 ascomycetes, 1 basidiomycete and 153
mitosporic fungi from 6 grass and 1 sedge hosts in Hong Kong.
4.4.2 Host exclusivity and recurrence
The fungi on the same bamboo host but from different sites were highly similar
(similarity indices ranged from 70 to 90) and correlated well(correlation coefficients
ranged from 0.891 to 0.988) (confidence 99%). The fungi from different bamboo hosts
at the same sites were highly dissimilar (similarity indices ranged from 14 to 38) and
not well correlated (correlation coefficients ranged from -0.01 to 0.213) (Tables 4.64.11). This indicates that some saprobic fungi have a significant association with some
bamboo hosts, i.e., they exhibit host exclusivity or host recurrence.
There are many examples of terrestrial saprobic fungi being recurrent on a
single host (Zhou and Hyde, 2001, in press). For example, Oxydothis alexandrarum
very commonly occurs on Archontophoenix alexandrae, but not on adjacent palms
(Taylor et al., 2000). In this study, some fungi were found with a high frequency on
one or more bamboo hosts. Host recurrent fungi are listed with their hosts in Table
4.12. It is interesting that some fungi, such as Astrosphaeriella fissuristoma and A.
stellata exclusively occurred on Bambusa, which indicates genus exclusivity, while
Astrocystis cocoes and Pestalotiopsis uvicola only occurred on Fargesia yunnanensis,
which may indicate species exclusivity (Tables 4.2-4.3). On the other hand,
Gilmaniella bambusae occurred on the four bamboo hosts of different genera. This
fungus may have family recurrence/exclusivity.

183

Chapter 4: Host Recurrence and Biodiversity

Table 4.12 Recurrent fungi on bamboo hosts in Hong Kong and Kunming
Species

Hosts

Frequency of
occurrence (%)*
29.8

Hong Kong

Location

Anthostomella rehmii

B. tuldoides

Apiospora sinensis

F. yunnanensis

34.6

Kunming

Arecophila bambusae

B. textilis

22.0

Hong Kong

Astrocystis cocoes

F. yunnanensis

16.7

Kunming

Astrosphaeriella fissuristoma

B. shiuyingiana

32.3

Hong Kong

Astrosphaeriella stellala

14.4

Hong Kong

17.6

Pestalotiopsis uvicola

B. shiuyingiana, B. textilis,
B. tuldoides
B. textilis, F. yunnanensis,
N. affinis, P. hetroclata
F. yunnanensis

20.10

Hong Kong,
Kunming
Kunming

Phaeoacremonium parasiticum

N. affinis

17.33

Kunming

Podosporium elongatum

B. textilis

17.03

Hong Kong

Podosporium nilgirense

B. tuldoides

17.07

Hong Kong

Roussoella hysterioides

F. yunnanensis

26.37

Kunming

Roussoella pustulans

B. shiuyingiana

20.93

Hong Kong

Gilmaniella bambusae

*: Frequency of occurrence is an average value among the different sampling sites.


4.4.3 Percentage of recurrent fungi
Thirteen of the 56 taxa (19.6%) identified had a high frequency of occurrence
and distinct host recurrences. In calculating fungal estimates, different percentages of
host specific fungi have been used. Hawksworth (1991) estimated that 2/3 of the fungi
were host specific, while Hyde (1996) gave a more conservative estimate, i.e., 25% of
fungi being host specific. These authors did not refer to host recurrence. In this study,
ca. 20% of saprobes were recurrent on certain hosts. This does not include pathogens
and other ecological groups of fungi and therefore a figure of between 30-40%
specific/recurrent fungi is probably realistic. Host specificity and recurrence, involve
'selectivity' or some method by which fungi, are restricted to their hosts. It is not clear
if this figure applies to other host families, but it would be interesting to reexamine
fungal numbers taking into consideration these new figures.

184

Chapter 4: Host Recurrence and Biodiversity

4.5 LIMITATIONS TO THIS STUDY


It would be better if hosts other than bamboo were also studied in the same
habitat. Comparison of the mycota from surrounding plants may further confirm host
recurrence or host exclusivity.
This field experiment was conducted at one sampling time and does not reflect
fungal succession or seasonality.
The habitats sampled in this study were within the subtropics (Kunming) or
intermediate transitional tropics (Hong Kong), which did not represent all global
climatic zones.
4.6 FUTURE STUDIES
Mycologists should internationally cooperate to survey certain hosts in all
climatic zones at the same time in order to get closer to a realistic percentage of host
specific/exclusive/recurrent fungi among the total fungal world.
Future studies with other plants are needed to confirm host exclusivity and recurrence.
This study provides a figure for host recurrence/exclusivity in saprobes.
Endophytes, parasites, mycorrhizal fungi and other ecological groups may also be
specific, exclusive or recurrent to their hosts. Mycologists should extend this study to
cover the whole range of ecological groups.

185

Chapter 4: Host Recurrence and Biodiversity

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Bills, G.F. (1994). Abundance and diversity of microfiingi in leaf letter of lowland
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3.

Bills, G.F., Holtzman, G.I. and Miller, O.K. (1986). Comparison of


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Boidin, J., Candoussau, F. and Gilles, G. (1986). Bambusicolous fungi from the
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Boyd, C.E. and Hess, L.W. (1970). Factors influencing shoot production and
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Candoussau, F., Katumoto, K. and Sherwood, RM. (1985). Bambusicolous fungi


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Cannon, P.F. (1997b). Strategies for rapid assessment of fungal diversity.


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Chapter 4: Host Recurrence and Biodiversity

13. Frohlich, J. and Hyde, K.D. (1999). Biodiversity of palm fungi in the tropics: are
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14. Hammond, P.M. (1992). Species inventory. In: Global biodiversity: Status of the
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15. Hawksworth, D.L. (1991). The fungal dimension of biodiversity: magnitude,
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17. Holliday, P. (1998). A dictionary of plant pathology. Cambridge University Press,
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18. Huhndorf, S.M. and Lodge, D.J. (1997). Host specificity among wood-inhabiting
pyrenomyctes (Fungi, Ascomycetes) in a wet tropical forest in Puerto Rico.
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19. Hyde, K.D. (1995). Measuring biodiversity: Diversity of microfungi in north
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23. Lodge, D.J. (1997). Factors related to diversity of decomposer fungi in tropical
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Chapter 4: Host Recurrence and Biodiversity

decomposition of Phragmites communis Trin. and Typha angustifolia. Journal of


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25. May, R.M. (1991). A fondness for fungi. Nature 352: 475-476.
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leaves of two rain forest trees in Puerto Rico. Journal of Industrial Microbiology
17: 284-294.
29. Poon, M.O.K. and Hyde, K.D. (1998). Evidence for the vertical distribution of
saprophytic fungi on senescent Phragmites australis culms at Mai Po Marshes,
Hong Kong. Botanica Marina 41: 285-292.
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31. Spooner, B.M. and Candoussau, F. (1988). Bambusicolous fungi from the
southwest of France: III. A new species of Encoelia. Transactions of the
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many species coexist in the tropics. America Naturalist 133: 240-256.
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distribution of microfungi associated with three palm species from tropical and
temperate habitats. Journal of Biogeography 27: 297-310.
34. Tsui, 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.
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from Hong Kong. Mycological Research 102: 435-438.
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industry. In: Biodiversity of Tropical Microfungi (Hyde, K.D. ed). Hong Kong
University Press, The University of Hong Kong, Hong Kong. pp. 29-46.
37. Wong, K.M. (2000). Diversity, Host Preference, and Vertical Distribution of
saprobic fungi on grasses and sedges on Hong Kong. Ph.D. Thesis, The
University of Hong Kong, Hong Kong. 243 p.
38. Zhou, D.Q. and Hyde, K.D. (2001). Host specificity, host exclusivity and host
recurrence in saprobic fungi. Mycological Research (in press).

189

Chapter 5: Fungal Succession

CHAPTER 5: FUNGAL SUCCESSION


ABSTRACT
Fungal succession on Bambusa tuldoides in Hong Kong was effected by
seasonality and fungal occurrence peaked in the wet seasons. Rainfall positively
impacted on fungal occurrence, but temperature and relative humidity appear to have
little influence. Fungal communities changed over time during the succession progress.
Based on temporal replacement, the fungal communities on bamboo baits can be
categorized into early colonizers, middle-stage colonizers, later colonizers, regular
inhabitants and sporadic inhabitants. Fungal communities on naturally dead bamboo
and baits mainly comprised rare species and middle-stage colonizers. Anthostomella
species are regular inhabitants being dominant throughout the observation period and
probably played a dominant role in bamboo decomposition.
Key words: colonizers, bamboo baits, fungal community, inhabitants, seasonality.
5.1 INTRODUCTION
Fungal succession is a directional change in the composition, relative
abundance and spatial patterns in species communities. Usually fungal succession is
the sequential occupation of the same site by mycelia either of different fungi, or of
different associations of fungi. Fungi replace one another as their dynamic
communities of mycelia alter in space and time, each species being adapted for
occupation of particular niches (Frankland, 1998). Frankland (1998) reviewed studies
of fungal succession and pointed out that each succession is unique, dependent on the
host material and its environment. Studies of fungal succession on plant substrates are
well documented (Wildman and Parkinson, 1979; Kuter, 1986; Frankland, 1992, 1998;
Tokumasu et al, 1994), but fungal succession on bamboo is poorly documented.
Leung (1998) placed several bamboo baits in terrestrial habitats in Hong Kong and
observed a succession of fungi occurring on these baits. Sixty fungal taxa (including
33 unidentified fungi), comprising 16 ascomycetes and 44 mitosporic fungi were
identified during a 48-week exposure period. The observation time, however, was

190

Chapter 5: Fungal Succession

relatively short, as in temperate regions fungal succession on bracken litter would


probably not cease until after 5-6 years, when bacteria are dominant and 95% dry mass
reduction is estimated to take 11-23 years (Frankland, 1976, 1998). Leung (1998)
observed fungal succession on bamboo baits over one year, when the fungal
succession was probably still at an initial stage. The size of bamboo baits were also
small (10 cm x 2 cm) so that the majority of fungi in the succession progress may not
have been sampled. A longer-time survey using larger bamboo baits is therefore
desirable. In this chapter, succession and diversity of fungi on naturally dead bamboo
samples and baits of Bambusa tuldoides at Tai Po Kau Nature Reserve, Hong Kong
was investigated. The relationships between fungal succession and climatic factors are
also analyzed.

5.2 SITE DESCRIPTION


5.2.1 General description of Hong Kong sites (refer to chapter three)
5.2.2 Study site
Tai Po Kau Nature Reserve (refer to chapter three).
5.3 MATERIALS AND METHODS
The study site covering about 0.3 hectares was selected in a bamboo forest,
which comprised two bamboo species, i.e., Bambusa tuldoides and B. shiuyingiana.
The bamboo forest lies on the bank of Tai Po Kau forest stream near the lookout post,
at the field office of Tai Po Kau Nature Reserve, Department of Agriculture and
Fishery, Hong Kong, SAR. Bambusa tuldoides was selected for the fungal succession
study. On 23 August 1998, 114 healthy bamboo plants were randomly cut, labeled and
left to rot naturally on the forest floor. Every two months, four whole bamboo plants
were randomly retrieved and cut into samples ca. 25 cm long. Fifty samples were
randomly selected for study each time. At the same time, one standing healthy bamboo
sample was also collected as a control. Fifty naturally dead bamboo culm samples (ca.
25 cm long) were also randomly collected from the same site at the same time, in
order to compare the fungal diversity with that of the succession samples. A total of
1100 naturally dead bamboo samples and bamboo baits were collected during the two-

191

Chapter 5: Fungal Succession

year observation period. The samples were returned to the laboratory, where they were
incubated in zip-lock polythene bags lined with moistened tissue for three days. The
succession samples were checked after three days, one week, one month and two
months for fungal fruiting bodies. Squash mounts and sections of fungal fruiting
bodies were mounted in water for observation and measurement. The healthy bamboo
was washed several times in water and treated in the same way as the succession study
samples. In order to compare the mycota on bamboo baits and from soil at the site, two
soil samples were collected from below the bamboo baits and two soil samples were
collected where there were no bamboo baits. The soil collections were carried out on
23 March (dry season) and 23 May 2000 (wet season).

5.4 RESULTS
5.4.1 Species-area curves
Two species-area curves for collections carried out in December 1998 (the
lowest species richness in the observation period) and June 2000 (the highest species
richness), are selected to represent the whole observation period (Figures 5.1-5.2). The
minimum number of bamboo samples required to identify the majority of fungal
species varied from 23 to 25. Examination of 30 bamboo samples was therefore
considered adequate to acquire the majority of fungi occurring on the bamboo.
5.4.2 Fungal diversity
Fifty-seven fungal taxa were found on the bamboo baits and this was higher
than that on naturally dead bamboo samples (38 species). The average number of fungi
occurring on each bamboo bait was 2.1, but on each naturally dead bamboo sample it
was 0.75. (Table 5.3). The diversity indices (Shannon-Wiener index) for fungal
communities on bamboo baits was 2.37, while that on naturally dead bamboos was 2.
The similarity indices between the fungal communities occurring on naturally dead
bamboo samples or bamboo baits during different periods was generally low (Table
5.5-5.6).

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Chapter 5: Fungal Succession

5.4.3 Fungal seasonality


Figure 5.3 indicates that the occurrence of fungi on both naturally dead bamboo
samples and baits were seasonal. In the dry seasons, species richness was the lowest
(e.g. December 1998), while in the wet seasons, species richness was highest (e.g.
June 2000). The frequency of occurrence on naturally dead bamboo samples at each
collecting time varied from 2 (in Dec. 1998) to 27 (in June 2000), with troughs in the
dry seasons and peaks in the wet seasons. The diversity indices (Shannon-Wiener
index) of fungal communities on naturally dead bamboo samples varied from 0.5
(December 1998) to 2.6 (June 2000) (Table 5.1), while on bamboo baits, the indices
ranged from 0 (Oct. 1998) to 3.06 (Sept. 1999) (Table 5.2). The evenness of fungal
communities on naturally dead bamboo samples changed from 0.5 (December 1998)
to 0.8 (June 2000) (Table 5.1), while on bamboo baits, evenness varied from 0 (Oct.
1998) to 0.7 (March 1999) (Table 5.1).

5.4.4 Similarity of the fungal communities


The similarity indices (Bray-Curtis coefficients) between fungal communities
occurring on naturally dead bamboo samples, varied from 5 (between mycota
collected in March 1999 and July 1999) to 61 (between mycota collected in September
1999 and June 2000). In most cases, the fungal communities collected at different
periods were dissimilar (Table 5.5).
Similarity indices (Bray-Curtis coefficients) between the fungal communities
occurring on bamboo baits varied from 3 (between mycota collected in December
1998 and May 1999) to 72 (between mycota in March 1999 and May 1999). Most
fungal communities collected at different periods were also dissimilar (Table 5.6).
5.4.5 Fungal succession
In general, most fungi on bamboo baits were rare species, which were
encountered once or twice. Several species occurred on bamboo baits for ca. 6 month
periods, while quite a few species were present on the baits throughout the observation
period (Table 5.2). The temporal replacement of fungi can be categorized into (1) early
colonizers, i.e., those fungi occurring during the first 4 months and disappearing

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Chapter 5: Fungal Succession

thereafter; (2) middle-stage colonizers, i.e., these dominant during 6-10 months and
appearing thereafter; (3) late colonizers, which became dominant after 10 months and
persisted until the end of the experiment; (4) regular inhabitants, which occurred
throughout or mostly throughout the experimental period, and (5) sporadic inhabitants,
which occurred on bamboo baits sporadically (Table 5.4).
The dominant early colonizers were Apiospora sinensis, Arthrinium luzulae, A.
phaeospermum, Coelomycetes sp., Fusarium sp., Petroconium sp., Roussoella
pustulans

and

Veronaea indica; while Acrodictys bambusicola, A. erecta,

Anthostomella sulcigena, Apiospora montagnei, Astrosphaeriella australiensis,


Cordelia johnstonii, Eutypella gliricidiae, Gilmaniella bambusae, Monochaetia
karstemi, Oxydothis oraniopsis, Ramichloridium musae, Roussoella hysterioides,
Spirodecospora bambusicola and Thyric/mm chrysomallum were middle-stage
colonizers. Late colonizers were Astrosphaeriella stellata, Ellisembia bambusae and E.
bambusicola, while Anthostomella contaminans, A. flagellariae, Podosporium
elongatwm, Astrosphaeriella fissuristoma, Phaeoisaria clematidis and Podosporium
nilgirense were regular inhabitants (Table 5.2). There were twenty-six sporadic
inhabitants, which occurred on bamboo baits once or twice. These occurrence of these
fungi during different periods are summarized in Table 5.4.
Mitosporic fungi were dominant during all observation periods, followed by
ascomycetes. Marasmius rotula was the only basidiomycete, and was identified during
the later stages of succession on baits (Table 5.2 and 5.4).
5.4.6 Soil fungi isolation
Soil fungi were surveyed twice at the fungal succession site on 30 March 2000
and 23 May 2000. Five fungal taxa were identified, i.e., Aspergillus glaucus, A. niger,
Penicillium chrysogenum, Ramichloridium musae and Verticillium chlamydosporium.
Only one of the soil fungi, i.e., Ramichloridium musae, was also found on both
naturally dead bamboo samples and bamboo baits (Tables 5.1 and 5.2).

194

Chapter 5: Fungal Succession

Table 5.1 Frequency of occurrence of fungi on naturally dead bamboo

195

Chapter 5: Fungal Succession

Table 5.2 Frequency of occurrence of fungi on bamboo baits

196

Chapter 5: Fungal Succession

Table 5.3 Fungal species diversity on naturally dead bamboo and bamboo baits
Naturally dead bamboo samples
Bamboo baits
20
22
0
1
18
34
38
57
413
1134
0.75
2.1
0.84*
2.0
2.37
*: Sorensen's index shows similarity between fungal communities on dead bamboo samples and

Ascomycetes
Basidiomycetes
Mitosporic fungi
Species richness
Individual collections
Species/per sample
Sorensen"s index
Shannon index
bamboo baits.

Table 5.4 Ecological classification of fungi on bamboo baits


Successional group
Earl\ colonizers
Middle-stage colonizers
Late colonizers
Regular inhabitants
Sporatic inhabitants

Ascomycetes
2
8
2
3
7
21

Basid iomycetes
0
0
0
0
1
1

Mitosporic fungi
6
6
2
3
18
35

Z
8
14
3
6
26
57

Fig. 5.1 Species-area curve of the fungi identified on naturally dead bamboo
samples in December 1998

197

Chapter 5: Fungal Succession

Fig.5.2 Species-area curve of the fungi identified on naturally dead bamboo


samples in June 2000

Fig. 5.3 Climatic factors and frequency of occurrence of fungi on bamboo baits
and naturally dead bamboo samples

198

Chapter 5: Fungal Succession

Table 5.5 Similariity analysis (Bray-Curtis coefficient) of fuiigal communi ties on


naturally dead bamboo samples
Date
10/98

10/98
100

12/98

12/98

03/99

05/99

07/99

09/99

11/99

02/00

46

11

16

16

12

10

100

13

14

12

100

14

17

100

23
100

03/99
05/99
07/99
09/99

04/00

06/00

09/00

40

17

14

20

16

36

24

12

15

14

30

11

23

19

51

17

13

12

47

38

22

18

28

39

38

43

100

40

15

29

62

47

16

33

29

32

100

27

27

23

100

49

51

100

57

100

11/99
02/00
04/00
06/00

100

09/00

Table 5.6 Similarity analysis (Bray-Curtis coefficient) of fungal communities on


bamboo baits
Date

10/98

12/98

03/99

05/99

07/99

09/99

11/99

02/00

04/00

06/00

09/00

100

100

23

18

24

14

13

12

100

72

42

23

21

19

100

47

28

30

23

15

14

50

39

42

36

30

100

47

56

57

49

100

48

30

40

100

52

47

*
10/98
12/98

03/99
05/99
07/99

100

09/99
11/99
02/00
04/00

100

06/00

71
100

09/00

*: 10/98's value is zero (no fungi occurred when the healthy bamboos was newly cut).
5.5 DISCUSSION
5.5.1 Where do the early colonizers come from?
The issue of where the early colonizers come from has yet to be clearly
established (Frankland, 1992, 1998, Leung, 1998; Norden et ai, 1999). The early
colonizers may be endophytes, as endophytes exist asymptomatically in all tissues of

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Chapter 5: Fungal Succession

living plants (Bacon and White, 2000). Umali et al. (1999) studied endophytes in
Bambusa tuldoides at Tai Po Kau, Hong Kong. They identified 23 species and 14
"morphospecies" of mycelia sterilia. Among the early colonizers in this study,
Arthrinium phaeospermum, Fusarium sp. and Podosporium elongatum were also
found as endophytes by Umali et al. (1999), while Apiospora sinensis, Arthrinium
phaeospermum, Coelomycete sp., Fusarium sp., Roussoella pustulans and Veronaea
indica were also found on naturally dead bamboo samples, which were endophytes
(Umali, et al., 1999). Fusarium spp. was also found on samples in this study, after they
were incubated in zip-lock polythene bags lined with moistened tissues. They were
also possibly endophytes.
One soil fungus, i.e., Ramichloridium musae, was found on bamboo baits as an
earlier colonizer and was also found on naturally dead bamboo samples (Tables 5.15.2). It is therefore unlikely that the majority of early colonizers directly came from
soil in situ. The similarity indices (Serensen's index) between fungal communities on
bamboo baits and naturally dead bamboo samples were high (0.84) (Table 5.3). The
two fungal communities were very similar with many fungal species common to both
substrates (Tables 5.1-5.2). The early colonizers may therefore arrive from other
nearby substrata, such as naturally decaying bamboos. Endophytes, however, can not
be excluded, as 4 species identified by Umali et al. (1999) were also found as early
colonizers on bamboo baits and many of the endophytes isolated in traditional studies
cannot be identified and remain as mycelia sterilia (Umali et al., 1999).
5.5.2 Successional replacement
Fungi replace one another at different stages of succession and this has been
observed by various authors (Frankland, 1976, 1998; Leong et al., 1991; Ho, 1998;
Leung, 1998; Tokumasu, 1998). Frankland (1976) found that weak parasites first
dominated on dead bracken, followed by primary saprobes, then secondary saprobes
and finally common soil fungi. Ho (1998) categorized the fungi on submerged wood
baits into 4 groups, i.e., early colonizers, late colonizers, regular inhabitants and
sporadic inhabitants. Leung (1998) also divided fungi identified on bamboo baits into
two groups, i.e., early colonizers and regular inhabitants. There were 5 groups in this

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Chapter 5: Fungal Succession

study: early colonizers, late colonizers, middle-stage colonizers, regular inhabitants


and sporadic inhabitants, although they mainly comprised sporadic inhabitants and
middle-stage colonizers (Table 5.2). Frankland (1998) reviewed previous studies and
explained the possible mechanism for successional replacement of fungi. She thought
there were three levels of causes: at the highest level were general causes, e.g.
availability of space and species of differential performance; at the intermediate level
were processes and phenomena conditioning the first, e.g. dispersal, competition
between species and animal grazing involving whole communities; and at the most
detailed level were inherent characters that defined the outcome of the whole
communities and operated between populations and individual mycelia, e.g.
differential rates of growth and nutrient uptake.
5.5.3 Fungal diversity
Mitosporic fungi (34 spp.) were the dominant group on the baits during the
observation period, followed by ascomycetes (22 spp.) and 1 basidiomycete (Tables
5.2-5.3). The fungal communities on naturally dead bamboo samples were dominated
by ascomycetes (Table 5.1 and 5.3). Marasmius rotula (a basidiomycete species)
appeared on the bamboo baits (as a sporadic inhabitant) after 18 months. Previous
authors have also reported that mitosporic fungi were dominant during succession
studies. Leung (1998) found that mitosporic fungi (44 spp.) were dominant over
ascomycetes (16 spp.) on the baits of Bambusa sp. in Hong Kong and he also
identified an unidentified basidiomycete occurring on the bamboo baits during the late
stage of succession. Tsuneda (1983) suggested that the reason why basidiomycetes
occurred late in fungal successions was because of their relatively low saprobic
competitive abilities for the water- or solvent-soluble constituents of plant materials.
Several authors have also reported that mitosporic fungi were dominant over other
taxonomic groups in the succession process. Tokumasu et al. (1994) observed fungal
succession on pine needles in Germany and identified 1 ascomycete, 1 basidiomycete,
62 mitosporic fungi and 9 zygomycetes. Similar results have also been reported by
Gamundi et al. (1987), Aoki et al. (1990) and Ho (1998).

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Chapter 5: Fungal Succession

5.5.4 Comparison of fungal diversity on bamboo baits and naturally dead


bamboo
Species richness and frequency of occurrence of fungi on naturally dead
bamboo samples were lower than that on bamboo baits (Tables 5.1-5.3). It is likely
that the bamboo sampled in this study died several years ago. According to the field
workers at Tai Po Kau Nature Reserve, many bamboo plants were blown over in 1996
by a typhoon. This may mean that fallen bamboo plants at the site had been dead for
about 4 years. Thus, the fungi on the bamboo may be at the late stages of succession,
when fungal species and total numbers begin to decline (Dix and Webster, 1995). The
naturally dead bamboo plants in situ looked much more decayed and easier to break
than the bamboo baits.
Fungal succession on various baits are well documented (Willoughby and
Archer, 1973; Sanders and Anderson, 1979; Shaerer and von Bodman, 1983; Shearer
and Webster, 1991; Gonczol and Revay, 1993; Ghawana et al. 1997; Ho, 1998; Leung,
1998). Shearer and Crane (1986) recorded 134 fungi on submerged, decayed plant
substrata and balsa wood baits from two cypress-tupelo swamps and a hardwood
swamp and its adjoining lake in southern Illinois, USA. Ho (1998) identified 157 taxa
from 350 naturally occurring submerged wood samples. At the same time, he
identified 59 fungi from 140 Machilus velutina baits, and 60 fungi from 140 Pinus
massoniana baits in a freshwater stream at Tai Po Kau, Hong Kong. The average
number of fungi on naturally occurring wood, and baits of M. velutina and P.
massoniana were 3.1, 4.3 and 3.8. The results showed that the average number of
fungi on baits were higher than that on naturally occurring wood. This is perhaps that
some simple sacharides were available at early decay stage of the bamboo baits, and
some weak parasitic fungi as well as "sugar fungi", such as some mitosporic fungi,
could grow on them to utilize some simple sacharrides (Dix and Webster, 1995;
Frankland, 1998), but quite a few fungi could grow on deep decay bamboo culms
(Table 5.3).
In this study, Anthostomella species were regular inhabitants on naturally dead
bamboo samples and bamboo baits throughout most of the observation period (Tables
5.1 and 5.2). Lu and Hyde (2000) considered that Anthostomella species were

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Chapter 5: Fungal Succession

common on bamboo. This genus may play a dominant role in bamboo decomposition.
5.5.5 Dynamics of fungal communities on bamboo baits
A quantitative similarity index (Bray-Curtis coefficient) was used to assess the
simultaneous fungal communities on samples and baits at different times during the
succession period (Tables 5.5-5.6). Since most similarity values between the fungal
communities were very low, this indicates that the fungal communities changed over
time following death of the bamboo. There were only a few species in common
between different communities at different stages of succession. These findings
confirm that fungi replace one another in a succession process (Frankland, 1998). The
similarity indices (Shannon-Wiener index) among most fungal communities recovered
on the baits during observation periods were low, ranging from 3 to 49. Fungal
communities recovered on baits in May and July 1999 and June and September 2000
had higher similarity values ranging from 71 to 72 (Tables 5.6). This indicates that the
fungal communities during these latter periods were relatively similar, probably
because the fungi were regular inhabitants or middle-stage colonizers, and occurred
more frequently during these periods.
5.5.6 Seasonality
Many authors have reported that seasonality effects fungal succession.
Tokumasu (1998) observed that the temperature at the surface of decaying needle litter
of Pinus densiflora was a major factor contributing to seasonal changes in interior
fungal communities. Pandey and Dwivedi (1984) also observed that Colletothchum
gloeosporioides and Fusarium oxysporum f. psidii on the leaves of Psidium guajava
were recorded more frequently in the rainy season. Aleem (1980) suggested that
mangrove fungi display a seasonal periodicity with greater fungal numbers and growth
intensity in the wet season. Lamore and Goos (1978) also noted that fungal species
richness on naturally occurring wood samples submerged in a temperate stream was
highest following a period of heavy rainfall. Kuthubutheen and Webster (1986)
pointed out that fruiting bodies of coprophilous fungi were abundant in the wet season
due to water availability. Leung (1998) also suggested that seasonal factors, especially

203

Chapter 5: Fungal Succession

air temperature and rainfall affected the development of the fungal communities on
bamboo baits. A similar phenomenon was also observed in this study. The frequency
of occurrence of fungi on both naturally dead bamboo samples and bamboo baits
peaked in the wet seasons, i.e., July 1999 and June 2000 (Figure 5.3). Rainfall is
therefore, an important factor, which has the effect of increasing fungal occurrence on
bamboo.

5.6 LIMITATIONS TO THIS STUDY

The two-year observation period is not long enough to observe complete fungal
succession. At the end of the observation period, the bamboo baits appeared to be
poorly decayed. Fungal succession may continue for up to 6-7 years in the UK
(Frankland, 1976), although in the tropics, the decay period may be somewhat
shorter.

No endophyte isolation was conducted in order to establish if endophytes could be


the early colonizers of bamboo baits.

Fungi that did not produce fruiting bodies would not been recorded. For example,
some mycelial rhizomorphs were often observed during the later stages of
succession, these were most likely basidiomycetes, but they could not be
identified, as they did not produce fruiting bodies.

204

Chapter 5: Fungal Succession

REFERENCES
1. Aleem, A.A. (1980). Distribution and ecology of marine fungi in Sierra Leone
(Tropical West Africa) Botanica Marina 23: 679-688.
2.

Aoki, T., Tokumasu, S. and Tubaki, K. (1990). Fungal succession on momi fir
needles. Transactions of the Mycological Society of Japan 31: 355-374.

3.

Bacon, C.W. and White, J.F. (2000). Microbial Endophytes. Marcel Dekker, Inc.,
New York, USA. 487 p.

4.

Dix, N.J. and Webster, J. (1995). Fungal Ecology. Champman and Hall, UK. 549
P-

5.

Frankland, J.C. (1976). Decomposition of bracken litter. Botanical Journal of the


Linnean Society 73: 133-143.

6.

Frankland, J.C. (1992). Mechanisms in fungal succession. In: The Fungal


Community, Its Organization and Role in the Ecosystem (Wicklow, D.T. and
Carroll, G.C. eds). Marcel Dekker, New York, USA. pp. 403-426.

7.

Frankland, J.C. (1998). Residential Address: Fungal succession - unraveling the


unpredictable. Mycological Research 102: 1-15.

8.

Gamundi, I.J. Arambarri, A.M. and Spinedi, H.A. (1987). Fungal succession in
Nothofagus dombeyi leaf litter. Revista del Museo de la Plata Seccion Botanica 14:
89-116.

9.

Ghawana, V.K., Shrivastava, J.N. and Kushwaha, R.K.S. (1997). Some


observations on fungal succession during decomposition of wool in soil.
Mycoscience 38: 79-81.

10. Gonczol, J. and Revay, A. (1993). Further studies on fungal colonizations of twigs
in the Morgo-stream, Hungary. Nova Hedwigia 56: 531-542.
11. Ho, W.H. (1998). Biodiversity, Ecological and Ultrastructural Observations of
Fungi on Wood submerged in Tropical Streams. Ph.D. Thesis, The University of
Hong Kong, Hong Kong. 229 p.
12. Kuter, G.A. (1986). Microfungal populations associated with the decomposition
of sugar maple (Acer saccharum) leaf litter. Mycologia 78: 114-126.
13. Kuthubutheen, A.J. and Webster, J. (1986). Water availability and the
coprophilous fungus succession. Transactions of the British Mycological Society

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Chapter 5: Fungal Succession

86: 63-76.
14. Lamore, B.J. and Goos, R.D. (1978). Wood-inhabiting fungi of a freshwater
stream in Rhode Island. Mycologia 70: 1025-1034.
15. Leong, W.F., Tan, T.K. and Jones, E.B.G. (1991). Fungal colonization of
submerged Bruguiera cylindrica and Rhizophora apiculata Wood. Botanica
Marina 34: 69-76.
16. Leung, S.S. (1998). A study of Saprophytic Fungi Associated with Bamboo
Culms in Terrestrial, Freshwater and Marine Habitats. Ph.D. thesis, The City
University of Hong Kong. 259 p.
17. Lu, B.S. and Hyde, K.D. (2000). A World Monograph of Anthostomella. Fungal
Diversity Series 7. Fungal Diversity Press, The University of Hong Kong, Hong
Kong.
18. Norden, B., Appelquist, T, Lindahl, B. and Henningsson, M. (1999). Cubic rot
fungi: corticioid fungi in highly brown rotted spruce stumps. Mycologia Helvetica
10: 13-24.
19. Pandey, R.R. and Dwivedi, R.S. (1984). Seasonal incidence of phylloplane
mycoflora of guava (Psidium guajava) with reference to fungal pathogens. Acta
Botanica Indica 12: 1-8.
20. Sanders, P.F. and Anderson, J.M. (1979). Colonization of wood blocks by aquatic
hyphomycetes. Transactions of the British Mycological Society 73: 103-107.
21. Shearer, C.A. and Crane, J.L. (1986). Illinois fungi XII. Fungi and mycxomycetes
from wood and leaves submerged in southern Illinois swamps. Mycotaxon 25:
527-538.
22. Shearer, C.A. and von Bodman, S.B. (1983). Patterns of occurrence of
ascomycetes associated with decomposing twigs in a midwestern stream.
Mycologia 75: 518-530
23. Shearer, C.A. and Webster, J. (1991). Aquatic hyphomycete communites in the
River Teign. IV. Twig colonization. Mycological Research 95: 413-420.
24. Tokumasu, S. (1998). Fungal succession on pine needles fallen at different
seasons: The succession of interior colonizers. Mycoscience 39: 409-416.
25. Tokumasu, S., Aoki, T. and Oberwinkler, F. (1994). Fungal succession on pine

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Chapter 5: Fungal Succession

needles in Germany. Mycoscience 35: 29-37.


26. Tsuneda, A. (1983). Fungal Morphology and Ecology: Mostly Scanning Electron
Microscopy. The Tottori Mycological Institute, Japan. 320 p.
27. Umali, T.E., Quimio, T.H. and Hyde, K.D. (1999). Endophytic fungi in leaves of
Bambusa tuldoides. Fungal Science 14: 11-18.
28. Wildman, H.G. and Parkinson, D. (1979). Microfungal succession on living leaves
ofPopulus tremuloides. Canadian Journal of Botany 57: 2800-2811.
29. Willoughby, L.G. and Archer, J.F. (1973). The fungal spora of a freshwater stream
and its colonization pattern on wood. Freshwater Biology 3: 219-239.

207

Chapter 6: General Conclusion

CHAPTER 6: GENERAL CONCLUSIONS


6.1 DOES DECAYING BAMBOO SUPPORT A HIGH FUNGAL DIVERSITY?
In the host recurrence and biodiversity study, 56 fungal species were identified
from 6 sampling bamboo species in Hong Kong and Kunming. Fungal communities
comprised mainly rare species. Approximately 9 species were identified from each
bamboo host in consideration of taking out the common species. Dalisay (1998)
identified 50 fungal species including 36 mitosporic fungi and 14 ascomycetes in
Hong Kong, and 56 fungal species including 42 mitosporic fungi, 12 ascomycetes and
1 basidiomycete in the Philippines. The fungi were identified from equal numbers of
Bambusa and Dendrocalamus samples in both regions. Bamboos therefore appear to
support a lower diversity of fungi, as compared to some other tropical or subtropical
monocotyledonous plants. Hyde (1995) calculated that there may be a ratio of 26
fungal taxa to each palm host species in Queensland, Australia. This calculation was
based on a belief that ca 3 plant pathogens, 100 endophytes and 10 saprobes could be
expected to develop on each palm species, and that 25% of these fungi might be host
specific. Frohlich and Hyde (1999) also suggested that 26:1 would be an appropriate
estimate for the ratio of host specific fungi to palm species in the tropics. They
identified 86 fertile fungal species, including 22 ascomycetes, 59 mitosporic fungi and
5 (anamorph/teleomorph) connections and 13 morphospecies (mycelia sterilia) from 3
individual trees of Licuala ramsayi in Australia. In Brunei they identified 132 fertile
fungi, including 30 ascomycetes, 3 basidiomycetes, 95 mitosporic fungi, 1
zygomycetes and 3 anamorph/teleomorph connections and 40 mycelia sterilia from 3
individual trees of Licuala sp. Poon and Hyde (1998) investigated vertical distribution
of saprobic fungi on senescent Phragmites australis culms at Mai Po Marshes, Hong
Kong. They identified 58 fungal taxa from at only two sampling times. Taylor et al.
(2000) also found 287 fungal taxa on three palm hosts in Australia, China (Hong Kong,
Hubei and Hunan), Malaysia, Singapore, Switzerland and UK. More recently Wong
(2000) also collected 215 fungal taxa, including 61 ascomycetes, 1 basidiomycete and
153 mitosporic fungi from 6 grass and 1 sedge hosts in Hong Kong. Based on the

208

Chapter 6: General Conclusion

above discussion, it is concluded that bamboo hosts appear to support a lower diversity
of fungi than other monocotyledonous hosts.
6.2 ARE FUNGAL COMMUNITIES GEOGRAPHICALLY DIFFERENT?
The regional distribution of certain fungal groups has been well documented
(Ryabova and Tomilin, 1980; Zang, 1981; Gray, 1983; Tan and Wu 1986; Van Maanen
and Gourbiere, 1997; Ho, 1998; Dalisay, 1998; Whitton, 1999; Frohlich and Hyde,
1999; Taylor et al., 2000). In this study the fungal communities on Phyllostachys
bambusoides in Hong Kong (transitional tropical region) and Kunming (subtropical
region) were compared and found to differ. Fungal diversity in Hong Kong was richer
than in Kunming. This result confirms, at least with bamboo fungi, that there is a
latitudinal species richness gradient of fungal taxa increasing towards the tropics
(Stevens, 1989). The main factors impacting on the differences between fungal
communities in these two regions may be climate. Temperature and rainfall in Hong
Kong better favor growth and development of fungi than in Kunming. Fungal
communities are strongly influenced by climate. Frohlich (1997) investigated
biodiversity of fungi on palms in Australia, Brunei, Ecuador and Hong Kong. She
found that similarities and differences between the mycotas were more consistent with
climatic than biogeographical factors or host distribution. Taylor et al. (2000) also
found different assemblages of fungi in association with palms in temperate regions as
compared to those in tropical regions. These differences were more related to climatic
influences than to the hosts sampled.

6.3 DOES A ' CORE GROUP OF FUNGI' EXIST ON BAMBOO?


A 'core group of fungi' is defined as those fungi that occur on certain host
species, genera or families with a relatively high frequency and do not constitute a
taxonomic group, but an ecological group. These 'core group of fungi' probably plays
a crucial role in processes of substrate decomposition (Wong et al., 1998). The 'core
group of fungi' occurring on Phyllostachys bambusoides comprised Didymosphaeria
futilis, Ellisembia bambusae, Linocarpon pandanicola in Kunming, while in Hong
Kong is comprised Anthostomella contaminans, Apiospora sinensis, Astrosphaeriella

209

Chapter 6: General Conclusion

palmar urn.
6.4 ARE SAPROBES ON BAMBOO RECURRENT OR EXCLUSIVE?
Thirteen of 56 (20%) fungal taxa found on the bamboo hosts occurred
frequently. Many of these taxa were exclusive to, or recurrent on bamboo or on
individual bamboo genera or species, indicating that there was some association
between the hosts and fungi. It would be interesting to consider fungal exclusivity and
recurrence in reexamining estimates of fungal numbers. In previous estimates,
mycologists have focussed to some extent, on host specificity of pathogenic fungi,
mycorrhizal fungi and endophytes, as compared to the recurrence of saprobes on hosts
(Singh et al, 1995; Guillot 1997; Newton and Haigh, 1998; Peter et al, 1998). This
result of the present study may therefore draw mycologists' attention to importance of
recurrence on hosts.
6.5 FUNGAL SUCCESSION
Fungal succession on decaying Bambusa tuldoides peaked during the wet
seasons. Rainfall is considered to be very important in determining species richness
and diversity of fungi on bamboo and accounted for these peaks. There was a
succession of fungal fruiting bodies developing on decaying bamboo, although
maximal diversity occurred during the wet seasons. The fungi developing on samples
were categorized into early colonizers, middle-stage colonizers, late colonizers,
regular inhabitants and sporadic inhabitants, although the fungi mainly comprise
sporadic inhabitants and middle-stage colonizers. Other researchers have found similar
successional trends (e.g. Frankland, 1998; Ho, 1998; Leung, 1998).

6.6 FUTURE STUDIES


In this study, 120 fungal taxa were identified on 34 bamboo species through
out 24 field forays during 1998-2000. Thirteen taxa, including 2 genera, and 1 variety
were new to science. On average, there were only 0.4 new taxa on each bamboo host
studied. In comparison with other monocotyledonous plants, such as palms (Frohlich
and Hyde, 1999; Taylor et al, 2000), the possibility of discovering new taxa on

210

Chapter 6: General Conclusion

bamboo was found to be low. Bamboo, however, has been poorly studied. For example,
in Yunnan, China, there are ca 170 species of bamboo with 23 genera that have had no
fungal records (Zhou et al., 2000). If we adopted the ratio found in this study, i.e., 0.4
new fungal taxa to each bamboo species studied, then we can expect to find ca 70 new
taxa on bamboo in Yunnan. This figure is probably an underestimate, as the present
study was conducted in the vicinity of Kunming, Yunnan, and no collection was
carried out in the tropical areas, such as Xishuangbanna, Yunnan, where more fungi
may occur.
It is still necessary to investigate biodiversity of microfungi on bamboo, as this
host has received relatively little attention from mycologists and some of bamboo
fungi are economically important (Kuai, 1996). Hyde (1995) considered microfungi to
present the biggest challenge in measuring biodiversity because of their abundance. In
China the lack of study of microfungi is acute, as most Chinese mycologists have
investigated biodiversity of macrofungi, paying little attention to microfungi (Zang,
1980, 1986; Huang, 1985; Zhuang, 1997; Wen and Sun, 1999). It would be desirable
to select small plots (10 m2) or individual host trees and sample within these elements
to establish microfungal diversity, both in the tropical and the subtropical regions
(Hyde, 1995).

211

Chapter 6: General Conclusion

REFERENCES
1. Dalisay, T.U. (1998). Biodiversity of microfungi associated with species of
Bambusa and Dendrocalamus. Ph.D. thesis, The University of Hong Kong, Hong
Kong. 345 p.
2.

Frankland, J.C. (1998). Residential Address: Fungal succession - unraveling the


unpredictable. Mycological Research 102: 1-15.

3.

Frbhlich, J. (1997). Biodiversity of microfungi associated with palms in the


tropics. Ph.D. thesis. The University of Hong Kong, Hong Kong.

4.

Frohlich, J. and Hyde, K.D. (1999). Biodiversity of palm fungi in the tropics: are
global fungal diversity estimates realistic? Biodiversity and Conservation 8: 9771004.

5.

Gray, N.F. (1983). Ecology of nematophagous fungi: Distribution and habitat.


Annals of Applied Biology 102: 501-510.

6.

Guillot, J. (1997). The bases for specificity of ectomycorrhizal fungi with respect
to their host. Revue Forestiere Francaise Nancy. 49 (special issue): 57-66.

7.

Ho, W.H. (1998). Biodiversity, ecological and ultrastructural observations of fungi


on wood submerged in tropical streams. Ph.D. thesis, The University of Hong
Kong, Hong Kong. 229 p.

8.

Huang, N.L. (1985). Notes on Phallales from Fujian, China. Wuyi Science Journal
5:211-218.

9.

Hyde, K.D. (1995). Measuring biodiversity: diversity of microfungi in north


Queensland. In: Measuring and Monitoring Biodiversity in Tropical and
Temperate Forests (Boyle T.J.B. and Boontawee, B. eds). CIFOR, Indonesia, pp.
271-286.

10. Kuai, S.Y. (1996). A check-list of pathogenic bambusicolous fungi of mainland


China and Taiwan. Journal of Forest Science and Technology, Subtropical
Forestry Institute, 4:64-71.
11. Leung, S.S. (1998). A study of Saprophytic Fungi Associated with Bamboo
Culms in Terrestrial, Freshwater and Marine Habitats. Ph.D. thesis, The City
University of Hong Kong. 259 p.
12. Newton, A.C. and Haigh, J.M. (1998). Diversity of ectomycorrhizal fungi in

212

Chapter 6: General Conclusion

Britain: A test of the species-area relationship, and the role of host specificity.
New Phytologist 138: 619-627.
13. Peters, S., Draeger, S. and Schulz, B. (1998). Interactions in dual cultures of
endophytic fungi with host and nonhost plant calli. Mycologia 90: 360-367.
14. Poon, M.O.K. and Hyde, K.D. (1998). Evidence for the vertical distribution of
saprophytic fungi on senescent Phragmites australis culms at Mai Po Marshes,
Hong Kong. Botanica Marina 41: 285-292.
15. Ryabova, V.P. and Tomilin, B. (1980). Micromycetes of the principal plant
communities in the Central Chernozem Reserve, Russian SFSR, USSR.
Mikologiya Fitopatologiya 14: 322-327.
16. Singh, U.S., Singh, R.P. and Kohmoto, K. (1995). Pathogenesis and host
specificity in plant diseases : histopathological, biochemical, genetic and
molecular bases. Elsevier Science, Oxford.
17. Stevens, G.C. (1989). The latitudinal gradient in geographical range: how so
many species coexist in the tropics. American Naturalist 133: 240-256.
18. Tan, H.C. and Wu, R.J. (1986). The ecological and geographical distribution of
108 species of macromycetes from the subtropical, evergreen, broad-leaves forests
in China. Mycotaxon: 25: 183-194.
19. Taylor, J.E., Hyde, K.D. and Jones, E.B.G. (2000). The biogeographical
distribution of microfungi associated with three palm species from tropical and
temperate habitats. Journal of Biogeography 27: 297-310.
20. van Maanen, A. and Gourbiere, F. (1997). Host and geographical distribution of
Verticicladium trifidum, Thysanophora penicillioides, and similar fungi on
decaying coniferous needles. Canadian Journal of Botany 75: 699-710.
21. Wen, H.A. and Sun, S.X. (1999). Fungal flora of tropical Guangxi, China:
Macrofungi. Mycotaxon 72: 359-369.
22. Whitton, S.R. (2000). Biodiversity studies of microfungi on Pandanaceae. Ph.D.
thesis. The University of Hong Kong, Hong Kong. 625 p.
23. Wong, K.M. (2000). Diversity, Host Preference, and Vertical Distribution of
saprobic fungi on grasses and sedges on Hong Kong. Ph.D. Thesis, The
University of Hong Kong, Hong Kong. 243 p.

213

Chapter 6: General Conclusion

24. Wong, M.K.M., Goh, T.K., Hodgkiss, I.J., Hyde, K.D., Ranghoo, V.M., Tsui,
C.K.M, Ho, W.H., Wong, WS.W and Yuen, T.K. (1998). Role of fungi in
freshwater ecosystem. Biodiversity and Conservation 7: 1187-1206.
25. Zang, M. (1980). The phytogeographical distribution of higher fungi and their
evaluation of natural resources in Yunnan and Xizang (Tibet, China). Acta
Botanica Yunnanica 2: 152-187.
26. Zang, M. (1981). Classification and distribution of Termitomyces from Yunnan,
China. Acta Botanica Yunnanica 3: 367-374.
27. Zang, M. (1986). Notes on the Boletales from eastern Himalayas and adjacent.
Acta Botanica Yunnanica 8: 1-22.
28. Zhou, D.Q. Hyde, K.D. and Vrijmoed, L.L.P. (2000). Resources and diversity of
bambusicolous fungi in China. Guizhou Science (China) 18: 92-70.
29. Zhuang, W.Y. (1997). Some new species and new records of Discomycetes in
China. VII. Mycotaxon 63: 307-321.

214

APPENDICES

APPENDIX 1:

REFERENCES
1. Boidin, J. 1998. Taxonomie moleculaire des Aphyllophorales. Mycotaxon 66:445-491.
2. Dalisay, T.D. (1998). Biodiversity of Microfungi Associated with Species of Bambusa and Dendrocalamus. Ph. D. thesis, The University of
Hong Kong, 345 p.
3. Eriksson, O.E. and Yue, J.Z. 1998. Bambusicolous pyrenomycetes, an annotated check-list. Myconet 1:25-78.
4. Leung S.S. (1998). A Study of Saprophytic Fungi Associated with Bamboo Culms in Terrestrial, Freshwater and Marine Habitats. Ph.D.
Thesis, The City University of Hong Kong. 259 p.
5. White, Jr., J.F., and P.V. Reddy. (1998). Examination of structure and molecular phylogenetic relationships of some graminicolous
symbionts in genera Epichloe and Parepichloe. Mycologia 90:226-234.

APPENDIX 2:
Table 2.1 Families/orders list of bambusicolous fungi of Mainland China*
Family/Order Name
Pucciniaceae
Phyllachoraceae
Xylariaceae
Dothideales
Lasiosphaeriaceae
Capnodiaceae
Meliolaceae
Hypocreaceae
Clavicipitaceae
Uredinales
Amphisphaeriaceae
Apiosporaceae
Corticiaceae
Dothioraceae
Phakosporaceae
Hyaloscyphaceae
Chaetothyriaceae
Diatrypaceae
Didymosphriaceae
Melanommataceae
Pseudoperisporaceae
Zipfiaceae
Coleosporiaceae
Fenestellaceae
Hemenochaetaceae
Hyponectriaceae
Lachnocladiaceae
Leotiales
Lentariaceae
Lophiostomataceae
Massarinaceae
Micropeltidaceae
Mycosphaerellaceae
Myriangiaceae
Myxotrichaceae
Phaeosphaeriaceae
Phallaceae
Polyporaceae
Schizophyllaceae
Septobasidiaceae
Tilletiaceae
Tricholomataceae
Thekephoraceae
Ustilaginales
Ustilaginaceae
45

Genera Name**
Puccinia{ 10), Stereostratum( 1)
Coccodiellai, 1), Coccostroma( 1), Phragmocarpella( 1), Phyllachora(J),
AnthostomellaQ), Engleromyces(l). Hypoxylon(2), RoselliniaQ),Xylaria(l)
DidymellaQ), Homoslegia{2), ShiraiaQ), Sinosphaeria(\),
Ceretosphaeria( 1), Lasiosphaeria( 1), Leptosphaerella(4), Miyoshia( 1)
Aithaloderma(\), Neocapnodium(\), Scorias(2), Triposporiopsis(2)
Meliola(6)
Calonectria( 1), Gibberella( 1), Lisea( 1), Loculistroma( 1), Shiraiella( 1)
Balansia(\), Epichloe(2), Konradia(\),
Aecidium( 1), Physopella( 1), Uredo(2)
Amphisphaeria(i),
ApiosporaO),
Corticium(2), Hypochnus( 1)
Metasphaeria(1)
DaslurellaQ)
Pseudolachneaii)
Chaetothyrium{ 1), Phaeosaccardinula( 1)
Eutypa{ 1), Eutypella{ 1),
Didymosphaeria(2)
Trematospaerella{ 1), Trematosphaeria( 1)
Acanthostigma{2),
Caryospora(2)
Chrysomyxa{ 1)
Fenestella{ 1)
Phellinus(\)
Physalospora{ 1)
Vararia{ 1)
Ascotremellopsis( 1)
Lentaria( 1)
Massariosphaeria( 1)
Massarina( 1)
Micropeltis{ t)
Mycosphaerella( 1)
Myriangium{ 1)
Campsotrichum{ 1)
Leptosphaerella{ 1),
Dicyophora{ 1)
Polyporus{ 1)
Schizophyllum( 1)
Septobasidium( 1)
Urocystis( 1)
Dictyopanus( 1)
Hyporeopsis( 1)
K\veilingia( 1),
Ustilago(\)
74

I
11
10
9
7
7
6
6
5
4
4
3
3
3
3
3
3
2
2
2
2
2
2

122

*: mitosporic fungi are not included in this table; **: the numbers in brackets are the
number of species.

Table 2.2 List of mitosporic fungi in Mainland of China


Generic Names

Number of Species
3
3
2
2
2
2
2
2
2
2

Coniosporium
Fusarium
Dinemasproum
Actinospora
Arthrobotryum
Cercospora
Cladosporium
Collectotrichum
Didymobotryum
Endothiella
Goniosporium
Helm inthosporium
Hemitrichia
Isaria
Leptostroma
Macrophoma
Afelanconium
Odocephalum glomerulosum var. cantonense
Papularia
Plectronidium
Prosthemiella
Sclecrotium
Scolicotrichum
Septocytella
Tetraploa
25

37

Table 2.3 Distribution of bambusicolous fungi species on bamboo genera


Hyphomycetes
Basiomycetes
Ascomycetes
Hosts
I
Bambusa

46

19

37

102

Phyllostachys

35

16

16

67

Arundinaria

Pseudosasa

Sasa

Dendrocalamus

Brachystachyum

Chimonobambusa

Pleioblastus

Yushania

88

43

58

189

10

PUBLICATIONS

The following publication contains material which is also included in this thesis and I
declare that I have been responsible for the share indicated for the work listed.

Signed
Dequn Zhou
Publication reference

Cryptophiale sphaerospora sp. nov. occurring


on Janetia synnematosa.
Mycoscience 40: 189-191,1999

Share of work by candidate

20%

Countersigned by co-workers
Name: Dr. T.E. Umali
Signature:
Name: Dr. T.K. Goh
Signature:
Name: Dr. K.D. Hyde
Signature:

189
Mycoscience 40: 189-191, 1999

Cryptophiale sphaerospora sp. nov. occurring on Janetia


synnematosa
Teresita E. Umali, Dequn Zhou, Teik-K. Goh and Kevin D. Hyde
Fungal Diversity Research Project, Department of Ecology and Biodiversity, The University of Hong Kong, Pokfulam
Road, Hong Kong
Accepted for publication 18 February 1999
Cryptophiale sphaerospora sp. nov. is described and illustrated based on a single collection attached to a synnematous
fungus, Janetia synnematosa. from a dead bamboo culm. It differs from other species of Cryptophiale in having spherical to subspherical conidia and a cerebroid layer of phialides. The overall morphology of this species is smaller than that
of previously described species.
Key Words

bamboo fungus; hyperparasitism; hyphomycetes; mycoparasites; taxonomy.

Cryptophiale Piroz. was established by Pirozynski (1968)


and a key to 16 accepted species is provided by Goh and
Hyde (1996). The genus is characterized by dematiaceous, setiform conidiophores, bearing a subapical to apical fertile region of phialidic conidiogenous cells, producing hyaline, unicellular to multiseptate conidia. The
conidia are of various shapes, e.g., clavate (Goh and
Hyde, 1996), uncinate (Matsushima, 1975), cylindricobclavate (McKenzie and Kuthubutheen, 1993), filiform
(McKenzie, 1993b), falcate (McKenzie, 1993a), fusiform
(Sutton et al., 1989). Cryptophiale species have been
recorded on various substrata. Most species were described from leaf litter, but C. enormis B. Sutton, Nawawi
& Kuthub. (Sutton et al., 1989) was found on roots,
whereas C. iriomoteana Matsush. (Matsushima, 1975)
and C. multiseptata Goh & K. D. Hyde (Goh and Hyde,
1996) were described from bark and submerged wood,
respectively.
In our study of fungi occurring on dead bamboo, an
undescribed species of Cryptophiale associated with the
synnemata of Janetia synnematosa Sivan. & Hsieh
(Sivanesan and Hsieh, 1990) was identified. This
Cryptophiale species is distinguished from other species
by its small dimensions, the presence of spherical to subspherical, unicellular conidia and a cerebroid layer of cells
clasping the conidiophore. It is therefore described as a
new species in this paper.
Taxonomy
Cryptophiale sphaerospora Umali & D. Q. Zhou, sp.
nov.
Figs. 1-9
Coloniae in synnematibus Janetiae synnematosae
surgentes. Conidiophora solitaria, disseminata, erecta,
recta vel leniter curvata vel flexuosa, non ramosa,
atrobrunnea, ad basim pallide brunnea vel hyalina, laevia,
crassitunicata, ad basim distante septata, apicem versus

dense septata, 69-100/im longa, ad basim 7.5-10/um


lata, super regionem fertilem 3-4 /jtm et ad apicem 2.53 /im lata. Regio fertilis subapicalis, 20-44 //m longa,
10-1 5 pm lata, convoluta, ex cellulis brunneis 2.5-4 //m
latis composita.
Cellulae conidiogenae phialidicae,
monoblasticae, ca 2.5 /im latae. Conidia hyalina, unicellularia, globosa vel subglobosa, laevia, 1.5-2 /im
diam.
Holotype: Hong Kong, New Territories, Tai Po
Kau, Tai Po Kau Country Park, on dead culm of Schizostachyum dumetorum, 8 September, 1998, D. Q. Zhou
(HKU(M)9103).
Etymology: The specific epithet, sphaerospora,
refers to the spherical shape of the conidia.
Colonies occurring in association with synnemata of
J. synnematosa.
Conidiophores solitary, scattered,
macronematous, mononematous, erect, straight to
slightly curved or flexuous, unbranched, dark brown,
pale brown to hyaline towards the apex, smooth, thickwalled, distantly septate at the base, becoming closely
septate towards the apex, 69-100/im long, 7.5-10//m
wide at the base, 3-4 pm wide above the fertile region,
2.5-3 /im at the tip. Fertile region subapical, 20-44 pm
long, 10-15//m wide, cerebroid, comprising brown cells,
2.5-4 ^m wide, coiling and clasping the shaft of the
conidiophore.
Conidiogenous cells phialidic, monoblastic, ca 2.5 //m wide, arising perpendicular to the axis
of the conidiophores. Conidia produced in slime, hyaline, unicellular, spherical to subspherical, smooth, 1.52 ftm in diam.
Most species of Cryptophiale have falcate, fusiform,
clavate to obclavate conidia, while those of C.
sphaerospora are spherical to subspherical. Cryptophiale sphaerospora also has shorter conidiophores and
narrower fertile region when compared to the morphology of other accepted species (conidiophore = 69-100 /im
vs 175-450//m; fertile region = 20-24 x 10-15 ^m vs

190

T. E. Umali et al.

Figs. 1-6. Cryptophiale sphaerospora.


Light micrographs from holotype. 1. A portion of the colony of Janetia synnematosa with C. sphaerospora on dead culm of bamboo. 2. Conidiophore of C. sphaerospora on synnema of J. synnematosa.
3, 4. Rows of brown cells clasping and coiling on the
shaft of conidiophores. 5, 6. Fertile region showing layers of brown cells. Note the mass of conidia (arrowed). Scale bars: 1,
2=100/<m; 3 - 6 = 1 0 / m .

A new species of Cryptophiale on bamboo

1 3 - 1 1 0 x 9 . 5 - 3 0 / / m ) (Goh and Hyde, 1 9 9 6 ) . The fertile region of C. sphaerospora comprises cerebroid b r o w n


cells, a character t h a t is distinct. This is the first report
of a species of Cryptophiale associated w i t h a fungus.
Acknowledgements
The authors would like to thank Ms.
Helen Leung for technical assistance and Mr. A. Y. P. Lee for
photographic assistance. The University of Hong Kong is
thanked for the award of Postgraduate studentship to T. E.
Umali and D. Q. Zhou and the award of a Postdoctoral fellowship
t o T . K. Goh.
Literature cited
Goh, T. K. and Hyde, K. D. 1996. Cryptophiale multiseptata,
sp. nov. from submerged wood in Australia, and keys to the
genus. Mycol. Res. 100: 999-1004.
Matsushima, T. 1975. Icones microfungorum a Matsushima

191

lectorum. Published by the author, Kobe, Japan.


McKenzie, E. H. C. 1993a. New hyphomycete species from
litter in the Chatham Islands, New Zealand. Mycotaxon
46: 291-297.
McKenzie, E. H. C. 1993b. New species of Cryptophiale from
New Zealand and New Caledonia. Mycotaxon 49: 3 0 7 312.
McKenzie, E. H.C. and Kuthubutheen, A . J . 1993. Dematiaceous hyphomycetes on Freycinetia (Pandanaceae). 4.
Cryptophiale. Mycotaxon 47: 87-92.
Pirozynski, K. A. 1968. Cryptophiale, a new genus of Hyphomycetes. Can. J. Bot. 46: 1123-1127.
Sivanesan, A. and Hsieh, W. H. 1990. A synnematous species
of Janetia from Taiwan. Mycol. Res. 94: 566-569.
Sutton, B.C., Nawawi, A. and Kuthubutheen, A. J. 1989. Additions to Belemnospora and Cryptophiale from Malaysia.
Mycol. Res. 92: 354-358.

The following publication contains material which is also included in this thesis and I
declare that I have been responsible for the share indicated for the work listed.

Signed
Dequn Zhou
Publication reference

A new species of Spadicoides and new records


of bambusicolous hyphomycetes from Hong
Kong.
Fungal Diversity 3: 179-185,1999.

Share of work by candidate

80%

Countersigned by co-workers
Name: Dr. T.K. Goh

Signature:

Name: Dr. K.D. Hyde


Signature:
Name: Dr. L.L.P. Vrijmoed
Signature:

Fungal Diversity

A new species of Spadicoides and new records of bambusicolous


hyphomycetes from Hong Kong

De Qun Zhou1*, Teik Khiang Goh1, Kevin D. Hyde1 and Lilian L.P.
Vrijmoed2
1

Centre for Research in Fungal Diversity, Department of Ecology and Biodiversity, The
University of Hong Kong, Pokfulam Road, Hong Kong; * e-mail: zhoudq@hkusua.hku.hk
:
Department of Biology and Chemistry, City University of Hong Kong, Tat Chee Avenue,
Kowloon, Hong Kong
Zhou, D.Q., Goh, T.K, Hyde, K.D and Vrijmoed, L.L.P (1999). A new species of Spadicoides
and other hyphomycetes on bamboo from Hong Kong. Fungal Diversity 3: 179-185.
Spadicoides bambusicola sp. nov. is described and illustrated from a dead culm of Bambusa
textilis collected in Hong Kong. Spadicoides bambusicola resembles S. palmicola in having
verruculose conidiophores producing verrucose, obclavate, multi-euseptate conidia, but differs
from S. bambusicola in its bigger, concolorous conidia having a non-rostrate apex.
Didymobotryum verrucosum, Janetia synnematosa, Penzigomyces flagellata, Veronaea
coprophila and V. indica are introduced as new records for Hong Kong.
Key words: bambusicolous fungi, Hong Kong, hyphomycetes, Spadicoides bambusicola,
taxonomy

Introduction
In our study of the fungal diversity on bamboo in Hong Kong, an
undescribed species of Spadicoides (Goh and Hyde, 1996a) and some other
hyphomycetes were collected. This undescribed species of Spadicoides
resembles S. palmicola Goh and K.D. Hyde (1998) in having verruculose
conidiophores, producing verrucose, obclavate, and multi-euseptate conidia.
Spadicoides palmicola, however, differs from this undescribed species in its
conidia with a rostrate, subhyaline apex and the conidia of S. palmicola are
smaller (x = 43 x 6 urn vs. 48 x 7 um, n = 50). Spadicoides bambusicola sp.
nov. is therefore described in this paper. Several other interesting
hyphomycetes were isolated and are introduced as new records for Hong Kong.
Taxonomy
Spadicoides bambusicola D.Q. Zhou, Goh and K.D. Hyde, sp. nov. (Figs. 1-7)
Etymology: bambusicola, referring to the bamboo host of this species.

179

Coloniae in substrato naturali pilosae, atrobrunneae. Mycelium plerumque in substrata


immersum. Conidiophora 150-350 x 4-5 um, macronematosa, mononematosa, non ramosa,
erecta, recta vel leniter flexuosa, multiseptata, crassitunicata, verruculosa, ad basim robusta et
5-7.5 um lata. Cellulae conidiogenae in conidiophoris incorporatae, terminates vel
intercalares, polytreticae, rectae. Conidia 30-72.5 x 5-7.5 um, recta vel curvata, obclavata, ad
basim obconico-truncata, verruculosa, 5-7-euseptata, ad septa plerumque constricta, viridulobrunnea vel olivace-obrunnea, crassitunicata.

Colonies on natural culms dense, hairy, dark brown. Mycelium mostly


immersed in the substratum. Conidiophores 150-350 x 4-5 um,
macronematous, mononematous, unbranched, erect, solitary, straight or slightly
flexuous, robust at the base (5-7.5 (am wide), multiseptate, verruculose, thickwalled, mid grayish brown to dark olivaceous brown. Conidiogenous cells
integrated, terminal and intercalary, polytretic (up to 3 pores per cell). Conidia
30-72.5 x 5-7.5 um (3c = 48 x 7 um, n = 50), acropleurogenous, solitary,
straight to slightly curved, obclavate, obconically truncate at the base, subacute
at the apex, verrucose, with 5-7 eusepta, usually slightly constricted at the
septa, thick-walled, olivaceous-brown or greenish brown.
Material examined: HONG KONG, New Territories, Tai Po Kau Nature Reserve, on
dead culms of Bambusa textilis, 10 Apr. 1998, D.Q. Zhou and T.E. Umali (HKU(M) 8333,
HOLOTYPE; 9165, isotype).

Notes: Spadicoides was established in 1958 by S. Hughes with the type


species S. bina (Corda) S. Hughes (Ellis, 1971). Goh and Hyde (1996a)
reviewed the genus and discussed the similarities and differences between
Spadicoides and Diplococcium. They accepted 21 species in Spadicoides. Two
further species, namely S. palmicola Goh and K.D. Hyde and S. heterocolorata
(R.F. Castaneda, Guarro and Cano) Goh and K.D. Hyde were added recently
(Goh and Hyde, 1998). Spadicoides bambusicola cannot be confused with the
species of Helminthosporium, although in the both genera, conidiogenus cells
are polytretic, integrated, terminal and intercalary, determinate, cylindrical. The
conidia in the species of Helminthosporium are pseudoseptate, frequently with
a prominent, dark brown or black scar at the base (Ellis, 1971).
New records of bambusicolous hyphomycetes for Hong Kong
1. Didymobotryum verrucosum Hino and Katumoto, Bulletin of the Faculty
of Agriculture, Yamaguti University, 10: 1181 (1959).
Colonies effuse dark olivaceous-brown. Synnemata straight, up to 1 mm
tall, 25-26 um thick; separating threads 2-3 um thick, septate, pale to mid
olivaceous-brown. Conidiogenous cells usually clavate, swollen at the apex to
4-5 um. Conidia 18-22 x 6-8 um, clavate, cylindrical or ellipsoidal, rounded at
the ends, pale to greyish olive.

180

Fungal Diversity

Figs. 1-7. Spadicoides hamhusicola, light micrographs. 1. A portion of the colonies on bamboo
culm. 2. Close-up of conidiophores with conidiogenous pores (arrowed). 3. Apex of a
conidiophore with a developing conidium. 4-7. Conidia. Bars: 1 = 100 urn, 2 = 5 urn, 3 = 50
Urn, 4-7 = 10 |im.
Material examined: HONG KONG, Hong Kong Island, Lung Fu Shan Country Park, on
dead culms of Arundinaria hindsii Munro, 10 Apr. 1998, D.Q. Zhou (HKU(M) 9086).

181

fungi

Notes: This collection resembles Didymobotryum verrucosum (Ellis,


1971), especially in the dimensions of the conidia (18-22 x 6-8 um vs. 15-26 x
5-7 um) on dead wood of Sasa in Japan (Ellis, 1971). This fungus has been
recorded also on a variety of bamboos, e.g. on Pseudosasa japonica and Sasa
borealis var. purpurascens in Japan (Hino, 1961) and an unknown palm in
Indonesia (Seifert, 1990).
2. Janetia synnematosa Sivan. and W.H. Hsieh, Mycological Research 94:
566(1990).
(Figs. 8-10)
Colonies effuse, brown to dark brown. Synnemata scattered, straight,
cylindrical, dark brown to black, up to 27 mm tall, 500 urn wide at the often
bulbous base composed of compactly arranged conidiophores. Conidiophores
2.5-4 um thick, septate, smooth, brown to dark brown. Conidiogenous cells 1014 x 4-8 um, integrated, denticulate, terminal, intercalary and lateral, mono- to
polyblastic, determinate, thick-walled, truncate and 2-3 um wide at the apex.
Conidia 78-92 x 14-16 um, holoblastic, solitary, dry, 9-22-distosepatate,
smooth, obclavate, rostrate, pale brown to brown, with a truncate and darkened
base.
Material examined: HONG KONG, New Territories. Tai Po Kau Nature Reserve, on
dead culms of Schizostachyum dumetorum (Hance) Munro, 8 Sep. 1998. D.Q. Zhou (HKU (M)
9103); ibid., on senescent stems of Thysanolaena maxima (Roxb.) Kuntze, 16 Sep. 1997,
Michelle K.M. Wong (HKU(M) 12504).'

Notes: Janetia was reviewed by Goh and Hyde (1996b) who accepted 16
species. Janetia synnematosa was first recorded on Miscanthus floridulus in
Taiwan (Sivanesan and Hsieh, 1990). It has also been recorded on the
senescent stems of Thysanolaena maxima (Roxb.) Kuntze (M.K.M. Wong,
pers. comm.). Whereas the present collection was from dead culms of
Schizostachyum dumetorum. Janetia synnematosa appears to have a preference
for the grass family (Poaceae), whereas other Janetia species were reported
from living or dead dicotyledonous leaves (Goh and Hyde, 1996b).
3 Penzigomyces flagellata (Hughes) Subram.. Proceedings of Indian Natural
Science Academy, B58: 186 (1992).
(Figs. 12-14)
For synonyms refer to Subramanian (1992).
Colonies effuse dark blackish brown. Mycelium partly superficial and
partly immersed. Conidiophores 50-180 x 5-7 um, straight, mid to dark brown,
with 1-3 annellations. Conidia 52.5-97.5 x 11-12.5 um, 4-5 urn wide at the
base, verrucose, pale to mid olivaceous-brown.
Material examined: HONG KONG, New Territories, Sai Kung, Tu Kwa Pin, on dead
culms of Indocalannis sinicus (Hance) Nakai, 30 July, 1998, D.Q. Zhou (HKU(M) 9058).

182

Fungal Diversity

Figs. 8-10. Janetia synnematosa. 8. Synnemata with conidia. Note pleurogenous conidia
(arrowed). 9. Conidium. 10. A conidiogenous cell (arrowed) on a synnema, bearing a
developing conidium. 11. Veronaea coprophila, conidiophores and conidia. Figs. 12-14.
Penzigomyces Jlagellata. 12. Conidiophore with a developing conidium. 13, 14. Conidia. Bars:
8 = 300 urn, 9-14 =10 urn.

183

Notes: In reassessing Sporidesmium Subramanian (1992) separated


Penzigomyces with the type species P. nodipes (Penz. and Sacc.) Subram. and
other 12 new combinations. Six new species of Penzigomyces were added by
Subramanian (1997). Sporidesmium flagellatum (Hughes) M B . Ellis was then
transferred in Penzigomyces. This specimen matches Ellis' description of
Penzigomyces flagellata, but the conidia are slightly wider (52.5-97.5 x 1112.5 urn vs. 55-105 x 10-11 urn) (Ellis, 1976). Penzigomyces flagellata is
known from dead branches of Citrus in Ghana (Ellis, 1976) and Ripogonum
scandens in New Zealand (Hughes, 1977). This is the first record from bamboo
and Hong Kong.
4. Veronaea coprophila (Subram. and Lodha) M.B. Ellis, More
Dematiaceous Hyphomycetes, CAB International: 210 (1976).
(Fig. 11)
Colonies effuse dark brown. Conidiophores up to 350 long and 3-4.5 (am
wide, straight or flexuous, septate, mid to dark brown, paler towards the apex,
cicatrized. Conidia (6-)7-8 x 4 um, solitary, dry. acropleurogenous, straight,
cylindrical or ellipsoidal, conico-truncate at the base. 1-2-septate. hyaline.
Material examined: HONG KONG. New Territories. Mu Tsz Lam Country Park, on dead
culms of Phyllostachys bambusoides, 29 July 1998, D.Q. Zhou (HKU(M) 9073).

Notes: This collection matches Veronaea coprophila. but the conidia are
slightly narrower (7-8 x 4 um vs. 6-12 x 3-5 urn) (Ellis. 1976). Veronaea
coprophila was originally described from goat dung in India (Ellis. 1976) and
has been recorded from Artemisia. Bamhusa. Celtis. Dendrocalamus. Grewia.
Hydrangea. Passiflora. Primus. Rumex. Solanum and Zea (Paul et al., 1990).
This is the first record from Phyllostachys hamhusoides.
5. Veronaea indica (Subram.) M.B. Ellis, More Dematiaceous
Hyphomycetes, CAB International: 209 (1976).
Colonies effuse, dark brown or greyish brown, hairy. Conidiophores up to
150 long, 3-5 um wide, erect, straight or flexuous, brown, cicatrized at the
apex. Conidia 2.5-4 urn diam., acropleurogenous, verruculose. spherical or
subspherical, with a slightly protruding hilum, hyaline to pale brown.
Material examined: HONG KONG, Hong Kong Island, Lung Fu Shan Country Park, on
dead culms of Arundinaria hindsii Munro, 30 July 1998, D.Q. Zhou (HKU(M) 9084).

Notes: This specimen matches Veronaea indica except the conidia that was
narrower (2.5-4 um vs. 5-7.5 um; Ellis, 1976). The species has been previously
reported on Phoenix (Ellis, 1976). This collection extends the range of the
hosts of a very terrestrial saprotrophic fungus.

184

Fungal Diversity
Acknowledgements
D.Q. Zhou would like to thank The University of Hong Kong for the award of a
Postgraduate Studentship. We are grateful to Helen Leung for technical and photographic
assistance, to T.E. Umali for assistance in the collection of specimens and photograph plates
help. Michelle K.M. Wong is thanked for allowing us to cite her collection of Janetia
synnematosa on Thysanolaena maxima.

References
Ellis, E.B. (1971). Dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew,
UK: 608.
Ellis, E.B. (1976). More Dematiaceous Hyphomycetes. Commonwealth Mycological Institute,
Kew, UK: 507.
Goh, T.K. and Hyde, K.D. (1996a). Spadicoides cordanoides sp. nov., a new dematiaceous
hyphomycete from submerged wood in Australia, with a taxonomic review of the genus.
Mycologia 88: 1022-1031.
Goh. T.K. and Hyde, K.D. (1996b). Janetia curviapicis, a new species, and an emended
description of the genus. Mycologia 88: 1014-1021.
Goh, T.K. and Hyde, K.D. (1998). A new species of Spadicoides on Licuala sp. from Brunei,
and a note on S. heterocolorata, comb. nov. Canadian Journal of Botany 76: 1698-1702.
Hino, I. (1961). Species of fungi parasitic on bamboos in Japan. Icones Fungorum
Bambusicolorum: 1-333.
Hughes, S.J. (1977). New Zealand Fungi 25. Miscellaneous species. New Zealand Journal of
Botany 16:311-370.
Paul, Y.S., Bhardwaj, L.N. and Sharma, R.C. (1990). Additions to fungi of Himachal Pradesh
III (Hyphomycetes). Indian Journal of Mycology and Plant Pathology 20: 196-199.
Seifert, K.A. (1990). Synnematous hyphomycetes. Memoirs of New York Botanical Garden
59: 109-154.
Sivanesan, A. and Hsieh, W.H. (1990). A synnematous species of Janetia from Taiwan.
Mycological Research 94: 566-569.
Subramanian, C.V. (1992). A reassessment of Sporidesmium (Hyphomycetes) and some
related taxa. Proceedings of Indian Natural Science Academy B58: 179-190.
Subramanian, C.V. (1997). Hyphomycetes from South East Asia-novelties from Singapore and
Malaysia. Kavaka 22/23: 52-76.

185

The following publication contains material which is also included in this thesis and I
declare that I have been responsible for the share indicated for the work listed.

Signed
Dequn Zhou
Publication reference

Resources and diversity of bambusicolous fungi


in China.
Guizhou Science 18: 62-70, 2000.

Share of work by candidate

80%

Countersigned by co-workers
Name : Dr. K.D. Hyde
Signature:_
Name: Dr. L.L.P. Vrijmoed

/
Signature:

Vol.l8,No.l-2

J""- ' 2 0 0 0

GUIZHOU SCIENCE

(Phyllachoraceae)

Resources and diversity of bambusicolous fungi in China


Z H O U De-Qun ', Kevin D Hyde 1 and Lilian L P Vrijmoed 2

(1. Centre for Research in Fungal Diversity,

Department of Ecology and Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong

SAR,

People's Republic of China, 2. Department of Biology &. Chemistry, City University of Hong Kong, Tat Chee Avenue ,.Kowloon, Hong Kong SAR, People's Republic of China)
Abstract: The authors reviewed bambusicolous fungi based on literature study. Research history, biodiversity,
lifestyle, geographical distribution, host specificity, tissue specificity and succession of bambusicolous fungi were
reviewed. To date 189 species in 75 genera of bamboo fungi from China and 79 bambusicolous species fungi in 58
genera from Hong Kong have been reported. Most of them are ascomycetes, followed by basidiomycetes and then
hyphomycetes. The family Pucciniaceae (rust fungi) has the richest species divesity (20 species), followed by the
Phyllachoraceae with 15 species, Xylariaceae with 9 species and Capnodiaceae with 8 species.
Key words: bambooi fungi; diversityi ChinaiHong Kong
1003-6563 (2000)01-2-0062-09

Q949. 325

Introduction
T h e term biodiversity, which simply means " d i v e r s i t y of species" (Barthlott & Winiger,

1 9 9 8 ) , was developed during the National F o r u m on Biodiversity, held in W a s h i n g t o n , D . C. , on


September 21-24, 1986, under the auspices of t h e National Academy of Sciences and the Smithsonian Institution (Reaka-Kudla et. al. , 1997). "Biodiversity is defined as all hereditarily based
variation at all levels of organization, form the genes within a single local population or species,
to the species composing all or part of a local c o m m u n i t y , and finally to the communities themselves that compose the living parts of the multifarious ecosystems of the world" (Reaka-Kudla et
al. , 1 9 9 7 ) . Based on this definition, the unit of m e a s u r e m e n t used in this review of biodiversity
is based on number of species (exceptional v a r i e t i e s ) . R o s s m a n ( 1 9 9 7 ) pointed out t h a t within

:2000-01-16

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63

the fungi, species concepts vary considerably and the genetic basis for these concepts is largely
unknown and 'species richness' is only one (and a scientifically problematical) approach to biodiversity issues. The main references used in this review are "Sylloge Fugorum Sinicorum" (Dai,
1979); "Fungi of China" (Teng, 1996) i "Check-list of pathogenic bambusicolous fungi in Mainland China and Taiwan" (Kuai, 1996) and "Bambusicolous pyrenomycetes, an annotated checklist" (Erriksson and Yue, 1998). All other bambusicolous records are retrieved from the 'Index
of Fungi' on Internet (http://ars-grin.gov/fugaldatabases/). Hino (1938) is the first author to
use the term 'fungorum bambusicolorum' (bambusicolous fungi) but he did not give a definition.
'Bambusicolous' means "something lives on bamboo". Bambusicolous includes any fungi growing
on any bamboo substrates, which include leaves, culms, culm leaves, branches and roots.' The
fungi living on bamboo products (such as moulds on bamboo products) and freshwater fungi, marine fungi and the hyperparasitic fungi are not discussed in this review. The bambusicolous fungi
reported from Mainland China are discussed and compared with Hong Kong. The bambusicolous
fungi of Taiwan are also compared.

2 Research history of bambusicolous fungi


The most important literature on Chinese mycology is "Fungi of China" (Teng, 1996. The
books was first published in 1963 and reprinted in 1964 in Chinese. An English version was published by the Department of Plant Pathology, Cornell University and edited by Richard P. Korf).
The second one is "Sylloge Fugorum Sinicorum" , compiled by Dai (1979). All fungi known from
China up to 1979 are included in this book. The book was written in Chinese. Recently all the
fungi recorded in "Sylloge Fugorum Sinicorum" are accessible on Internet (http://ars-grin.gov/
fugaldatabases/). Oedocephalum glomerulosum (Bull. ) Sacc. var. cantonense Teng was the first
species reported from bamboo on a culm of Phyllostachys in Guangdong in 1932 (Dai, 1979;
Teng, 1996). Between 1934 to 38, 82 species of bambusicolous fungi were reported by Teng and
Ou from Mainland China (Teng, 1996), especially in the papers published by Teng and Ou
(1936), where 38 bambusicolous fungi were recorded. The Japanese invaded China during 1937~
1940 and therefore Teng only published 16 species of bambusicolous fungi. Kuai (1996) listed
190 pathogenic bambusicolous fungi in Mainland China and Taiwan. Eriksson and Jue (1998)
listed the bambusicolous pyrenomycetes on the Internet (http ://www. ekbot. umu. se/pmg/bam/
index, html) and reported 25 species known from China.
The first bambusicolous fungus record from Hong Kong was Puccinia tenella (Cummins,
1971). Consequently, Rosellinia congesta I. Hino & Katum from Bambusa sp. (Laeessoe &
Spooner, 1994) and Sporoschisma mirabile Berk & Broome were reported, from submerged dead
culm of Bambusa sp. (Goh et. al. , 1997). Recently Umali (1998) collected 47 species fungi from
Bambusa spp. and Dendrocalamus spp. Leung (1998) also found 37 bambusicolous fungi on unidentified bamboo, including many unidentified species. Therefore 73 species of bambusicolous
fungi have been found from Hong Kong.

Biodiversity of bambusicolous fungi

3. 1 Assessment of bambusicolous fungi resources


Hawksworth (1991) estimated there are at least 6 unique fungi species for each plant

18 #

64

species. Based on the estimation, he estimated there are 1. 5 million fungi worldwide. The estimation has been universally accepted. To date 189 species of fungi in 75 genera have been recorded
on bamboo in Mainland China. Based the ratio of 6 : 1 , there should be 2,400~3,000 fungi occurring on bamboo in Mainland China, where there are 400 500 bamboo species (Hui et at. ,
1996). If this estimate is accepted, the only 6. 3 ~ 7 . 9% of bambusicolous fungi is presently
known in Mainland China. In Taiwan there are ca 50 bamboo species in 9 genera (Huang, 1993;
Huang, 1994). Using the ratio of 1:6, ca 300 bambusicolous fungi should occur in Taiwan. At
present, 149 species of bambusicolous fungi are known from Taiwan. Meaning that 49. 7% of the
total bambusicolous fungi has been reported in Taiwan. There are 57 bamboo taxa in Hong Kong
(But et al. , 1985). About 340 bambusicolous fungi should be available, based on the ratio.
Based upon above the assessments, as estimated 3,0403,640 bambusicolous fungi should occur
in Mainland China, Hong Kong and Taiwan. The ratios of the host and fungi species known from
Mainland China and Taiwan are shown in table 1.
Table 1 Comparison of bambusicolous fungi in Mainland China and Taiwan

3. 2

Place

No. of bamboo species

No. of fungi species

Ratio (host: fungi)

Mainland

30

34

1: 1.13

Taiwan

21

102

1: 4. 85

Richness of mycota from different bamboo hosts


In many cases the bambusicolous fungi have no host description. The authors often recorded

as "on leaves and culms of bamboo" (Teng, 1996). In this section we calculate fungal diversity
on the different hosts, which have been recorded with detailed kkhost description. The highest
number of species is known from Bambusia spp. with 73 bambusicolous fungi records.
lostachys with 67 species and then Arundinaria,

Phyl-

Yushania and Pseudosasa with 5 records (Table

2). Bambusa and Phyllostachys are the most common genera with a broader distribution in east,
south and southwest China than the other bamboo genera (Gen, et al. , 1996). The two bamboo
genera are also multipurpose crops for industry and handicrafts use (as raw materials) and have
been planted artificially on a large scale nearby human communities (Gen et al. , 1996). The
bambusicolous fungi are likely to be collected more really by mycologists from these areas, as opposed to the other bamboo genera growing naturally in forest. The two bamboo genera also have
the richest species diversity, Bambusa containing more than 60 species and Phyllostachys having
more than 50 species in Mainland China (Gen et al. , 1996).
3. 3

Diversity analysis of bambusicolous fungi

3. 3.1

Mainland China

Ascomycetes have been recorded most frequently from bamboos in Mainland China (80
species accounting for 47. 9%) following by basidiomycetes (46, accounting for 27. 5%) and hyphomycetes (40, accounting for 24. 6%) (Table 2). The richest diversity with 20 species occurs
in the Pucciniaceae; the Phyllachoraceae with 15 species; the Xylariaceae with 9 species; the

65

Capnodiaceae and the Lasiosphaeriaceae with 4 species (Table 3). They are all ascomycetes. The
genera which have been recorded most often are Puccinia (10 species), Phyllachora (7 species)
and Meliola (6 species). In the case of mitosporic fungi, Coniosporium and Fusarium have 3
species and Actinospora has 2 species. Mitosporic fungi have been recorded less often than ascomycetes and basidiomycetes. Only one species are known from most genera.
Table 2 Occurrence of Bambusicolous Fungi on Bamboo Genera in Mainland
China (In decreasing order)
No.

Hosts

Ascomycetes

Basidiomycetes

Mitosporic fungi

1.

Bambusa

36

20

17

73

2.

Phyllostachys

35

16

16

67

3.

Arundinaria

4.

Pseudosasa

3"

5.

Sasa

6.

Dendrocalamus

7.

Brachystachyum

8.

Chimonobambusa

9.

Pleioblastus

10.

Yushania

10

80

46

41

167

3. 3. 2

Hong Kong

As Umali (1998) studied bambusicolous fungi in Hong Kong from two bamboo genera,
namely, Bambusa and Dendrocalamus and Leung (1998) did not identify the hosts of bambusicolous fungi in his thesis, it is difficult to make richness analysis for the bambusicolous fungi of
Hong Kong. Umali collected 556 fungal speciemens from Bambusa and 221 collections from Dendrocalamus (Umali, 1998) from the same site. Bambusa had a richer fungal diversity than Dendrocalamus. The two bamboo genera have almost the same distribution areas but Bambusa has
more species diversity (26 taxa) than Dendrocalamus (3 taxa) (But et al. , 1985). The genera
which have been most often recorded are the Xylariaceae (5 species) ; the Amphisphaeriaceae (3
species) and the Lasiosphaeriaceae (3 species). Both of the families are ascomycetes, the same
with Mainland China. Annulatascus has the richest species diversity (3 species), followed by Anthostomella, Astrospheriella, Rosellinia and Roussoella (2 species respectively). Only one species
are known from most genera. Most of mitosporic genera have only one species, accounting for
82% among all mitosporic fungi (Table 6) except that Acrodictys (5 species).

4 Life style of bambusicolous fungi


In the phytopathological point view, fungi can be divided into two main groups: saprobes,
which can obtain their food by attacking dead organic matter; pathogens, which live on/in living
plant tissues (Lucas, 1998). The latter group fungi can be divided into two subgroups: fungi capable of growing parasitically and of living saprobically on dead organic matter, according to circumstances, referred to as either facultative parasites or facultative saprobes (Lucas, 1998). The

18

66

first subgroup are always parasitic in nature and are called obligate parasites or biotrophs (Alexopoulos et al. , 1996) or biochemically obligate parasites (Lucas, 1998). The bambusicolous fungi of China include pathogens and saprobes. In general the pathogens include species of Puccinia,
Stereostratum and Uredo. Some of these fungi have very narrow host range and may occur only or
a single variety (Shao et. al. , 1984). Phyllachora,

Fusarium,

Sclerotium species are faculative

parasites on bamboo. Among all 189 bambusicolous fungi known from Mainland China, 164 fungi
are pathogens, which make up 86. 7% of the total (Kuai, 1996). This is probably because the
early bambusicolous fungal researchers in China were plant pathologists.

5 Geographical distribution of bambusicolous fungi


Most bamboo fungi are known from Taiwan with 149 records, the next is Hong Kong with
73 records, Jiangsu with 36 records and Guangxi with 35 records. Bamboo is mainly distributed
in the following provinces in China : Yunnan, Sichuan, Guangdong, Guangxi, Zhejiang, Fujian,
Taiwan, Hong Kong, Guizhou, and Hunan (Hui et al. , 1996). Yunnan has the richest bamboo
diversity in Asia with 210 woody bamboo species in 28 genera (Xue et al. , 1995). Hong Kong also has high bamboo diversity with 57 species in 14 genera of bamboo (But et al. , 1985) while in
Taiwan there are about 50 bamboo species in 9 genera (Hung, 1993; Huang, 1994). Therefore it
is odd that more bambusicolous fungi diversity have been recorded from Taiwan. It probably depends on intensity of bambusicolous fungi study and availability of mycologists as well as accessibility of mycologists to bamboo fungi. For example, During his fungal study, Teng's institute
was located in Nanjing, the capital of Jiangsu province. Perhaps that is why Jiangsu has the highest records with bambobusicolous fungi.

6 Host specificity
Host specificity infers a relationship between hosts and fungi, and has especially been applied
to plant pathogens (Lucas, 1998). Host specificity is a phenomenon in which it is thought that
both parasite genotype and host genotype influence the outcome of the relationship (Vanderplank, 1975). The specific parasite gene is for low pathogenicity and the specific host gene for
low reaction. Joosten et al. (1997) found that the interaction between tomato and Cladosporium
fulvum

has proven to be an excellent model for a host-pathogen specificity at molecular level.

Belisario et al. (1997) found that Phytophthora cactorum isolated from walnut proved to be highly
pathogenic to seedlings of English walnut (Juglans
seedlings of eastern black walnut (Juglans nigra).

regia)

and moderately pathogenic for

Furthermore, these isolates did not rot apple

seedlings, which infered host specificity. Erysiphe cichoracearum mildew from cosmos, sunflower
and cucumber infects eleven out of thirty-three species in eight families studied by Husain and
Akram (1995). They pointed out that isolates of cosmos and sunflower are restricted to the
Asteraceae, none of these isolates cross-infect the species of other families. The obligate parasitic
fungi have a very narrow host range, such as Puccinia graminis, greater host specificity may be
detected in the interaction of particular pathogen isolates (often described as races) and specific
host lines (cultivars) (Lucas, 1998). Some saprobes also exhibit weak host specificity or preference. Taylor (1998) observed that Oxydothis was exclusive to and frequently occurred on Archontophoenix alexandrae.

She stressed, however, host specificity was difficult to demonstrate (Tay-

67

lor, 1998). Umali (1998) could not find any evidences of host specificity for the fungi on Dendrocalamus and Bambusa during her study. She found a high diversity of fungi developing on Bambusa (75 species) indicating that the fungi on bamboo are extremely diverse, and therefore specificity at the subfamily level (Bambusoidea) would still have a significant impact on species numbers (Umali, 1998). The fungi on palms, which also occur in the tropics, found to be extremely
diverse and there is minimal overlap with those species occurring on palms (Frohlich, 1997; Taylor, 1998). Based on our calculations, 63 and 55 bambusicolous fungi have been reported on
Bambusa and Phyllostachys respectively. There are only 11. 9% bambusicolous fungi of the total
118 fungi on the both bamboo genera are the same. Puccinia longicornis and P.

phyllostachydis,

are found on the both host genera, although the both fungi are obligate parasitic (Kuai, 1996) , it
is different from the case of Puccinia graminis, which has high host specificity (Lucas, 1998),
even they belong to the same genus. It indicates that host specificity is quite complicated and not
all obligate parasitic fungi always have greater host specificity. Perhaps some of the fungi have
host specificity at family or subfamily level, under family or subfamily level, the host specificity
is indistinct. Janetia synnematosa is restricted on Poaceae but on different host genera (Zhou et
al. , in press).

Tissue specificity
Tissue specificity infers that particularly fungus is restricted to particular host tissues or or-

gans (Lucas, 1998). Fisher et al. (1992) isolated ascomycetes, deuteromycetes and zygomycetes
from 6 mature trees of Olea europaea growing on the Island of Majorca, Spain. They found that
only Kabatina sp. showed complete tissue specificity and was entirely confined to the xylem in all
trees. Acremonium spp. and Fusarium spp. were almost exclusively confined to the roots, while
others like Pestalotia spp. and Collectotrichum spp. were isolated from aerial plant organs (Dreyfuss and Petrini, 1984). Fisher et al. (1995) observed that the colonization frequency of endophytes in petiole of Gynoxis oleifolia was higher than that of other leaf parts and a gradient could
be seen in which the proximal parts were colonized more readily than the distal ones. Umali
(1998) observed that tissue specificity would be higher with host tissue age. Bambusicolous fungi
appear to have tissue specificity. Phyllachora shiraiana can be found on Bambusa,

Phyllostachys

and other several bamboo genera but is restricted to leaves (Kuai, 1996). Of the fungi occurs on
Phyllostachys species there are only 4 fungi which can live on more than one kind of host tissues
(Table 3). Of the 113 bambusicolous fungi known from Mainland China, which have detailed
host descriptions, only 6 fungi are known to occur on two types of host tissues or organs (Table
3).

8 Succession of bambusicolous fungi


Succession of both plants and fungi can be defined as a directional change in the composition,
relative abundance and spatial pattern of species comprising communities (Frankland, 1992).
Fungal succession is the sequential occupation of the same site by thalli (normally mycelia) either
of different fungi, or of different associations of fungi. Fungi replace one another as their dynamic communities of mycelia alter in space and time, each species adapted for occupation of particular niches. Early workers concentrated on replacement of species in time rather than in space, as

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18

pointed out by Frankland (1998). Succession studies fungi on bamboo are few. Leung (1998)
placed severa bamboo baits in terrestrial, freshwater and marine conditions and observed fungal
succession. In one site, 60 species were found during 48-weeks exposure period. Periconia
minutissima, Apiopsora montagnei state of Arthrinium, Giliocladium roseum, Nectria sp, Stilbella
sp. Wiesneriomyces javanicus, Phomopsis sp. were the first batch fungi appeared on the baits.
Periconia minutissima and Apiopsora montagnei state of Arthrinium existed throughout period the
study (Leung, 1998).

Economic importance
In terms of biodiversity conservation, a known universe of fungi would provide the means to

predict which groups of fungi are most diverse and valuable (Rossman, 1997). Most bambusicolous fungi known from China are important pathogens, e. g. Ceratosphaeria phyllostachydis
causes deadback of Phyllostachys pubscens (Kuai, 1996), and is broadly distributed across China.
Stereostratum corticioides is a common rust on many of bamboo species (Kuai, 1996). These
pathogenic bambusicolous fungi may cause economic losses by damaging bamboos. Some bambusicolous fungi are also medicial. Engleromyces goetzii, Hypocrella bambusae and Shiraia bambusicola are traditional Chinese medicines used to cure various human diseases (Ying et. al. , 1987).
Dictyophora indusiata, which are often associated with bamboo, is well known for its great medical and edible values (Ying et al. , 1987).
Table 3 Bambusicolous fungi on Phyllostachys from different tissues
fungi

No.
1

Amphisphaearia

2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
X

Arthrobotryum
rimpactum
Campsotrichum elegans
Ceretosphaeria
phyllostachydis
Corticium bambusac
Fenestella bambusicola
Fusarium
moniliforme
Goniosporium bambusae
Hypochnus pallescons
Hyporeopsis
phyllostachydis
Leptosphaeria
tigrisoides
Meliola stomata
Metasphaeria
denata
Mycosphaerella
babusifolia
Phragmocarpella
japonica
Phyllachora
shiraiana
Phyllachora sinensis
Ramaria
polypus
Scolecotrichum
phyllostachydis
Shiraia
Shiraia
Vararia
22

10 Conclusion

stellata

bambusicola
phyllostachydis
pallescens

culm

leave

branch

root

yes

yes
yes
yes
yes

1
1
1
1
1
1
1
1
1
1

/
1
1

/
1
1
1
1
1
1

/
yes

1
yes
yes
yes

1
1
1
1
1
1

yes
yes
yes
yes
yes
yes

yes

/
yes

/
yes

yes -

1
1
1

/ '
/
/
/
/
/
/
/
/
/
/
/
/
/

yes

yes

/
/
/

yes
yes

/
/
/
/
/
/
/
/

yes

yes
11

yes
' 9

2
26

69

A total of 413 species of bambusicolous fungi are known from China, including 189 species
from Mainland china, 75 species from Hong Kong and 149 specie from Taiwan. This accounts for
11- 3 1 3 . 5% of the assessed total 3 , 0 4 0 3 , 6 4 0 bambusicolous fungi number, which is based on
the conservative assessed ratio,

namely one vascular plant with six fungi,

proposed

by

H a w k s w o r t h ( 1 9 9 2 ) . There is a big gap between known fungi and the probable fungal resource.
For e x a m p l e , Yunnan has 23 bamboo genera with no fungi records. In the future, to exploit
bambusicolous fungi diversity is still very important, which is the first step for establishment of
all-taxa biodiversity inventory ( A T B I ) . Researchers must also provide detailed host descriptions,
which is important for diversity analysis. In China, the earlier researchers paid more attention on
pathogenic bamboo fungi than saprobes, probably because their applied importance. In fact,
some saprobic bamboo fungi are important economically or ecologically, such as antibiotics production (Wildman, 1997). There are some evidences to indicate host specificity and tissue specificity available in bambusicolous fungi. More observations and tests are necessary for further
prove these. Succession of bambusicolous fungi is poorly documented, as it is very difficult to be
observed and monitored.

11 Acknowledgments
Dequn Zhou would like to thank The University of Hong Kong for the award of a Postgraduate Studentship. Our cordial appreciation is also extended to Prof. Zhou Tong-xing, Ms Chen
Y u h u i , Mr. Liu Liang-zhong, Prof. Hui Caomao, Mr. Du Fan and Ms Zhang Xiaoping, Southwest Forestry College, Kunming; Mr. Yan Donghui, associate research professor, Forestry A cademy of China, Beijing, for their generously providing references.

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Press, Beijing. 1987, 579.

Fungal Diversity

Submersisphaeria
Kong

bambusicola sp. nov. from bamboo in Hong

Dequn Zhou and Kevin D. Hyde*


Centre for Research in Fungal Diversity, Department of Ecology and Biodiversity, The
University of Hong Kong, Pokfulam Road, Hong Kong; * e-mail: kdhyde@hkucc.hku.hk
Zhou. D.Q. and Hyde, K.D. (2000). Submersisphaeria bambusicola sp. nov. from bamboo in
Hong Kong. Fungal Diversity 4: 181-186.
A new Submersisphaeria species found from collections of fungi on bamboo in Hong Kong is
described. This fiingus has affinities with Submersisphaeria aquatica, but differs in having
long-pedicellate asci and unicellular ascospores. This new taxon is compared with S. aquatica,
Annulatascus velatisporus and Ascotaiwania lignicola.
Key words: Annulatascaceae, taxonomy.

Introduction
During a study of the fungal diversity on bamboo in Hong Kong, a new
Submersisphaeria species on dead culms of Arundinaria hindsii, from Mt.
Lung Fu Shan Country Park, Hong Kong was collected and identified. Hyde
(1996) established a monotypic genus, Submersisphaeria, which is typified by
S. aquatica K.D. Hyde. Characteristics salient to Submersisphaeria include
immersed or erumpent ascomata, short pedicellate asci with a massive,
refractive apical ring, and brown, bicellular ascospores with granular contents
and a single central septum which is slightly constricted (Hyde, 1996). In the
specimens from Mt. Lung Fu Shan, apart from the long pedicellate asci and
unicellular ascospores, all other characters are consistent with those in
Submersisphaeria. It is, therefore, necessary to extend the generic concept of
Submersisphaeria, and a new species, S. bambusicola is introduced to
accommodate the specimen.
Materials and methods
Dead culm samples of Arundinaria hindsii were collected from Lung Fu
Shan Country Park, Hong Kong Island, and returned to the laboratory where
they were incubated in polythene bags lined with moistened tissue. Material
was examined for bambusicolous fungi after 3 days and 1 week. Single-spore
isolation was attempted, but the ascospores failed to germinate on PDA at
181

room temperate. All microscopic measurements were taken from specimen


mounted in water.
Taxonomy
Submersisphaeria bambusicola D.Q. Zhou and K.D. Hyde, sp. nov.
(Figs. 1-15)
Etymology: From bamboo and the Latin cola meaning living on bamboo.
Ascomata 420-580 urn diam., 350-540 um alta, globosa vel subglobosa, immersa.
ostiolata, periphysata, solitaria. Peridium 50-60 urn crassum, nigrum, e textura angulari
compositum. Paraphysibus 4-8 um, crassis, septatis, numerosis et angustatis. Asci 235-290 x
10-12 um, octospori, cylindrici, leptodermi, longum pedicellati, apparatu apicale praediti.
Ascosporae (26-)28-36 x 6-8(-10) urn, uniseriatae, unicellulae. fusiformes-ellipsoideae.
fuscusa.

Ascomata deeply immersed in the host tissue, visible as a blackened cirrus


of ascospores, 420-580 urn diam. (x = 506 urn. n = 15), 350-540 urn high (x =
424 urn, n = 15), globose or subglobose, solitary, with a central periphysate
ostiolar canal up to 60 urn diam., 210 um high (Figs. 1. 2). Peridium 50-60 u.m
thick, comprising a brown outer layer and a hyaline inner layer (Fig. 4).
Paraphyses 4-8 um diam. (x = 6 (am, n - 15). hypha-like. filamentous, septate,
tapering, numerous (Fig. 3). Asci 235-290 x 10-12 um (.v = 258 x 11.6 urn, n =
50), 8-spored, cylindrical, thin-walled, with a long tapering pedicel, and a
relatively massive, elongate, refractive, non-amyloid apical ring, 5 um diam., 4
Um high (n = 10) (Figs. 5-8). Ascospores (26-)28-36 x 6-8(-10) um (3c = 31 x
7.5 \xm, n = 50), uniseriate or partly overlapping uniseriate, unicellular, mostly
inequilaterally ellipsoidal-fusiform, containing one lipid globule, smooth, dark
brown, with hyaline germ pores at each end, and sometimes with thin, drawnout sheath at each end (Figs. 9-15).
Anamorph. Unknown, as single-spore isolation tailed to germinate in culture.
Material examined. HONG KONG, Hong Kong Island. Lung Fu Shan, on a dead culm of
Antndinaria hindsii, 19 July 1998. Dequn Zhou [HKU(M)^9045, HOLOTYPE]; ibid.
[HKU(M)9353],

Discussion
Wong et al. (1998) introduced the Anmdatascaceae to accommodate
Anmtlatascus and AnnulatascusAike species, as the apical ring in these species
are similar and bipartite. In Anmtlatascus, ascomata are immersed to superficial
and black, and paraphyses are septate and wide. Asci are cylindrical with a
relatively massive refractive apical ring (Hyde, 1992). These fungi are mostly
found in freshwater habitats, although terrestrial species have been collected on
bamboo and palms in the tropics (Wong et al., 1998). Anmdatascus,
Aquaticola, Cataractispora, Clohiesia, Diluviocola, Fluminicola, Frondicola,
182

The following publication contains material which is also included in this thesis and I
declare that I have been responsible for the share indicated for the work listed.

Fungal Diversity

Figs. 1-15. Photo-micrographs of Submersisphaeria bambusicola (from holotype). 1.


Ascomata immersed in host tissue, with black ascospore mass (arrowed). 2. Vertical section
through ostiolar canal. 3. Paraphyses. 4. Peridium. Note it comprises a brown outer layer and a
hyaline inner layer. 5. Ascus. 6-8. Massive refractive apical rings of asci. 9-15. Ascospores.
Note the germ pores at each end and thin polar mucilaginous sheaths (arrowed). Bars: 1 = 500
Urn; 3 = 50 urn; 4 = 40 urn; 5, 9-15 = 20 urn; 2, 6-8 = 10 urn.

183

Fungal Diversity
Pseudoproboscispora and Submersisphaeria are the currently accepted genera
in the Annulatascaceae (Hyde, et al., 1998; Wong et al., 1998, 1999; Ho, et al.,
1999; Wong and Hyde, 1999). Submersisphaeria has dark brown ascospores
and thus differs from other genera in the Annulatascaceae which all have
hyaline ascospores (Hyde et al., 1998; Wong et al, 1998; Ho et al, 1999;
Wong and Hyde, 1999). Ascotaiwania was recently excluded from the
Annulatascaceae based on molecular studies (Ho et al., 1999; Ranghoo et al.,
1999). Ascotaiwania resembles Submersisphaeria in several aspects, however,
in Ascotaiwania the ascospores lacks germ pores, and with hyaline end cells
(Sivanesan and Chang, 1992; Hyde, 1996).
Submersisphaeria bambusicola is consistent with the generic description
of Submersisphaeria, especially in having subglobose, immersed ascomata,
cylindrical, unitunicate and pedicellate asci, with a refractive, nom-amyloid
apical ring and brown ascospores, with hyaline germ pores at each end (Hyde,
1996). Submersisphaeria bambusicola has long pedicellate asci and unicellular
ascospores, which conspicuously differs from S. aquatica, the type species of
Submersisphaeria. We believe that the characters, namely long pedicellate asci
and unicellular ascospores, are not good enough to establish a new genus,
which can distinguishes from Submersisphaeria. So the generic concept of
Submersisphaeria should be widened to include S. bambusicola.
Submersisphaeria bambusicola is saprobic and found on decaying bamboo
culms in terrestrial habitat in Hong Kong whereas S. aquatica is saprobic on
wood submerged in freshwater in Australia (Hyde, 1996).
Submersisphaeria bambusicola is compared with S. aquatica,
Annulatascus velatisporus and Ascotaiwania lignicola in Table 1.
Acknowledgements
Dequn Zhou would like to thank The University of Hong Kong for the award of a
Postgraduate Studentship. We are grateful to T.K. Goh for his contribution to the latin
diagnosis, Eric Mckenzie and Bing-Sheng Lu are thanked for comments on the manuscript and
Helen Leung for technical assistance.

References
Ho, W.H., Tsui, K.M.. Hodgkiss, I.J. and Hyde, K.D. (1999). Aquaticola, a new genus of
Annulatascaceae from freshwater habitats. Fungal Diversity 3: 87-97.
Hyde, K.D. (1992). Tropical Australian freshwater fungi. II. Annulatascus velatispora gen. et
sp. nov. Australian Systematic Botany 5: 117-124.
Hyde, K.D. (1996). Tropical Australian freshwater fungi. X. Submersisphaeria aquatica gen. et
sp. nov. Nova Hedwigia62: 171-175.
Hyde, K.D., Wong, S.W. and Jones, E.B.G. (1998). Dihiviocola capensis gen. and sp. nov. a
freshwater ascomycete with unique polar caps on the ascospores. Fungal Diversity 1: 133146.

185

Ranghoo. V.M., Hyde, K.D., Liew, E.C.Y. and Spatafora, J.W. (1999). Family placement of
Ascoiaiwania and Ascolacicola based on DNA sequences from the large subunit rRNA
gene. Fungal Diversity 2: 161-174.
Sivanesan. A. and Chang, H.S. (1992). Ascoiaiwania, a new amphisphaeriaceous ascomycete
genus on wood from Taiwan. Mycological Research 96: 481-484.
Wong. S.W. and Hyde. K.D. (1999). Proboscispora aquatica gen. et sp. nov., from wooc
submerged in freshwater. Mycological Research 103: 81-87.
Wong. S.W., Hyde. K.D. and Jones, E.B.G. (1998). Annulatascaceae. a new ascomycete
family from the tropics. Systema Ascomycetum 16: 17-25.
Wong, S.W., Hyde, K.D. and Jones, E.B.G. (1999). Ultrastructural studies on freshwater
ascomycetes, Fluminicola bipolaris gen. et sp. nov. Fungal Diversity 2: 189-197.
(Received 20 Oct. 1999. accepted 2 Feb. 2000)

186

The following publication contains material which is also included in this thesis and I
declare that I have been responsible for the share indicated for the work listed.

Signed
^Dequn Zhou
Publication reference

Striatodecospora gen. nov. from bamboo in


Hong Kong.
Myxotaxon 76: 141-147, 2000.

Share of work by candidate

80%

Countersigned by co-workers

Name: Dr. B.S. Lu

Signature:

Volume LXXVI, pp. 141-147

October-December 2000

STRIATODECOSPORA GEN. NOV. FROM BAMBOO IN


HONG KONG
Dequn Zhou*, Kevin D. Hyde and Bing-Sheng Lu
Centre for Research in Fungal Diversity, Department of Ecology and
Biodiversity, The University of Hong Kong, Pokfulam Road, Hong
Kong SAR, People's Republic of China
*Email: zhoudq@hkusua.hku.hk
ABSTRACT
From collections of fungi on bamboo in Hong Kong, we identified an
interesting, new taxon. The fungus has affinities with Spirodecospora,
but differs in that it lacks the verrucose ascospores with spiral wall
ornamentation and the surrounding fibrillar sheath of Spirodecospora.
Striatodecospora gen. nov. is described and illustrated to accommodate
this taxon.
Key words: bamboo fungi, Bambusa chungi, Spirodecospora,
Xy lariaceae.
INTRODUCTION
During a study of fungi on bamboo in Hong Kong we collected an
Anthostomella-like species on dead bamboo culms in Tai Po Kau
Nature Reserve, The New Territories. The collections could not be
suitably placed in Anthostomella, or other related genera in the
Xylariaceae, because the ascospores are covered with conspicuous
striate ornamentation and lack germ slits or pores. Striatodecospora

142

gen. nov. is therefore introduced to accommodate this taxon, with S.


bambusae as the type of this monotypic genus.

MATERIALS AND METHODS


Dead culms of Bambusa chungii, collected from Tai Po Kau
Nature Reserve, The New Territories, Hong Kong, were returned to the
laboratory and incubated in polythene bags lined with moistened tissue.
Material was periodically examined for the presence of fungal fruiting
bodies. Single spore isolations of Striatodecospora barnbusae were
attempted, but the ascospores failed to germinate. All microscopic
measurements were taken from specimens mounted in water.

DESCRIPTIONS
Striatodecospora D. Q . Zhou, K. D. Hyde and B. S. Lu, gen. nov.
Ascomata immersa, obpyriformia. ostiolata, paraphysata. Asci 8spori, cylindrici, unitunicati, ad apicem rotundati, jodi ope azurescens
praediti. Ascosporae late ellipsoideae. brunneae, unicellulae, cum
striatiter ornamentae.

Ascomata deeply immersed in host. obpyriform, ostiolate. often


solitary, but sometimes in pairs or gregarious. Peridium comprising
two layers. an inner layer of hyaline or pale brown, flattened cells, and
an outer layer of dark brown, thick-walled, angular or compressed cells,
fusing with host cells at the periphery. Paraphyses filamentous,
numerous, septate, branched and longer than asci with free ends. Asci
8-spored, cylindrical. short-pedicellate, unitunicate, apically rounded,
with a J+. wedge-shaped, subapical ring. Ascospores uniseriate or
partly overlapping uniseriate, ellipsoidal, brown, unicellular, lacking
germ slits or germ pores, with conspicuously striate ornamentation.
Etymology: from the Latin striato meaning "striate" and deco
meaning "decoration" in reference to the striate ornamentation on the
ascospores.
Typis generis: Striarodecospora barnbusae D. Q. Zhou, K. D.
Hyde arid B. S. Lu.

Striatodecospora bambusae D. Q . Zhou, K. D. Hyde and B. S. Lu, sp.


nov.
(Figs. 1-12)
Ascomata (190-) 215-230 (-245) pm in diametro, (260-) 285305 (-320) pm alta, obpyriformia, immersa, solitaria. Asci (1 lo-) 118132 x 10-12 p,octospori, cylindrici, subapicale rotundati, cum
annulis subapicalis, J+ cuneatis 1.3-2.5 x 2.5 pm, praediti. Ascosporae
14-16 x 5-6 pm, ellipsoideae, brunneae, striatiter omamentae, fissura
et poro germinatio non praeditae.
Ascomata deeply immersed in host, visible as blackened
flattened dots, often surrounded by black cluster of ascospores, usually
solitary+in vertical section (190-) 215-230 (-245) pm diam.
= 220.5
pm, n = lo), (260-) 285-305 (-320) pm high ( x = 295 pm, n = lo),
obpyriform, periphysate, with a central, ostiolar canal, 60-85 pm diam.,
170-275 pm high (Figs. 1-3,s). Peridium 14-22 pm wide ( x = 17 pm,
n = 15), comprising two layers, an inner layer of hyaline or pale brown,
flattened cells, and an outer layer of dark brown, thick-walled, angular
or compressed cells, fusing with host cells at the periphery (Fig. 6).
Paraphyses 2-4 pm wide, filamentous, numerous, septate, branched,
longer than asci and embedded in a gelatinous matrix (Fig. 4). Asci
(110-) 118-132 x 10-12 pm ( x = 125 x 11 pm, n = 15), &spored,
cylindrical, short-pedicellate, unitunicate, apically rounded, with a J+,
wedge-shaped, subapical ring, 2.5 pm diam. ( x = 2.5 pm, n = lo), 1.32.5 pm high ( x = 2 pm, n = 10) (Figs. 7-8). Ascospores 14-16 x 5-6
pm ( x = 14.5 x 5.5 pm, n = 30), uniseriate or sometimes partly
overlapping uniseriate, ellipsoidal, brown, unicellular, containing one
lipid globule, with striate ornamentation covering full length, lacking
germ slits or germ pores (Figs. 9-12).

(x

Etymology: Referring to the host.


Holotype: HONG KONG, New Territories, Tai Po Kau Nature
Reserve, on a dead culm of Bambusa chungii, 8 Sep. 1998, Dequn
Zhou (HKU (M) 9143).
Other material examined: HONG KONG, New Territories, Tai
Po Kau Nature Reserve, on a dead culm of Bambusa chungii, 8 Sep.
1998, Dequn Zhou (HKU (M) 9373).

Figs. 1-12. Striatodecospora bambusae (from holotype).


1 . Appearance of ostiole of ascoma on host surface. 2. Vertical section
of ascorna. 3. Ostiole. 4. Paraphyses. 5. Section of ascoma. 6. Peridium.
Note the inner part with hyaline, flattened cells and the outer part with
dark brown, thick-walled, angular or compressed cells. 7-8. Ascus with
8 ascospores. Note the J+, subapical ring in fig. 8. 9-12. Ascospores
with conspicuous, striate wall ornamentation (mowed). Bar: 1 = 100
p , 2 = 100 pm,3 = 2 0 pm,4= 10 prn, 5 = 100 w,6=10 p,7 =
20 pm, 8 = lOpm,9-12= lOprn.

Deeply immersed in
host, obpyriform with
a central ostiole
Cylindrical, shortpedicellate, with a J+,
wedge-shaped,
subapical ring
One-celled, brown,
lacking a germ slit or
germ pores, without a
sheath

Conspicuously
covered by striate
ornamentation

Ascomata

Asci

Ascospores

Wall of
ascospores

Conspicuously spirally
arranged vermcose wall
ornamentation

One-celled, olivaceous or
brown, lacking a germ slit or
germ pore, surrounded by a
hyaline, fibrous mucilaginous
sheath

Cylindrical, pedicellate, with


a J+, wedge-shaped,
subapical ring

Deeply immersed in host,


obpyriform with an oblique
or central ostiole

Amphisphaerella

One-celled, brown,
with equatorial perm
pores, often
surrounded by a
sheath
Smooth-walled

Mostly smooth-walled

Cylindrical,
pedicellate, with a
J+/J-, discoid or
wedge-shaped,
subapical ring

(Kirschsten, 1934;
Hyde, 1995)
Superficial with
only the base
immersed, globose
with a central ostiole

Mostly one-celled, light


brown, olivaceous or
black, mostly with a germ
slit, surrounded by a
sheath or with
mucilaginous appendages

Broad cylindrical,
pedicellate, with a J+/J-,
discoid or wedge-shaped,
subapical ring

Immersed often under a


clypeus, globose,
subglobose, or conical,
with a central ostiole

Anthostomella

(Francis, 1975)

Spirodecospora

(Lu et al., 1998)

Table 1. Comparison of Striatodecospora with some related genera


Striatodecospora

P
V,

146

DISCUSSION
Striatodecospora has many characters which are consistent with
the Xylariaceae, and may be well placed here (Hawksworth et al.,
1995). Similar genera include Anthostomella, A m p h i s p b l l a and
Spirodecospora. Striatodecospora bamburae cannot be confused with
Spirodecospora, which has vermcose ascospores covered with spiral
wall ornamentation, and larger ascospores (28-45 x 11-15 pm vs. 1416 x 5-6 pm) surrounding by fibrillar sheath (Lu et al., 1998).
Striatodecospora bambusae differs from Anthostomella species as
the ascospores are covered by conspicuous, striate ornamentation and
lack germ slits, germ pores, sheaths, or mucilaginous appendages.
Anthostomella is a species-rich and heterogeneous genus with more
than 250 species (Francis, 1975; Lu et al., 1998). Hyde (1996) reported
42 species of Anthostomella from palms and PandamLF, including ten
new species. In our understanding of Anthostomella species,
ascospores lack any spiral or striate ornamentation, but often have a
longitudinal germ slit and are surrounded by a gelatinous sheath.
Striatodecospora is distinct from Amphisphaerella, where the
ascomata are initially immersed but become superficial with only the
base immersed. Ascospores of Amphisphaerella are smooth, often
surrounded by a sheath, and have equatorial germ pores (Kirschsten,
1934; Hyde 1995).
All genera mentioned above are compared in Table 1. A key to
Striatodecospora and similar genera is provided.

Key to Striatodecospora and similar genera


I.
2.

3.
4.

Ascomata superficial, with base only immersed, ascospores with


equatorial germ pores-------------------A mphisphaerella
Ascomata immersed-------------------2
Ascospores with ornamentation--------------------3
Ascospores lacking ornamentation---------------------5
Ascomata forming in a stroma, asci clavate, ascospores having
Co coicola
sparse irregular striations or vermcose----------Ascomata lacking a stroma, asci cylindrical-------------4
Ascospores covered by striate ornamentation, lacking a
lnucilaginoussheath--------------------------------- S t r iatodecospora
Ascospores covered by spiral ornamentation, with a mucilaginous
sheath....................................................
S p irodecospo~

5.

6.

7.

8.

Ascomata forming in a stroma, ascospores having a central pallid


band and polar appendage...............................
F asciatispora
Ascomata lacking a stroma, ascospores lacking a central pallid
band........................................................................
6
Ascospores lunate, surrounded by an elaborate, layered sheath----....................................................................
Nip icola
Ascospores cylindrical or ellipsoidal.................................
7
Ascospores hyaline or yellow, with a thin sheath and blunt
appendages....................................................
Sa b alicola
Ascospores brown.......................................................
8
Ascospores thick-walled, reddish brown, with two polar germ
pores--------------------------------------------------------Pa ndanicola
Ascospores thin-walled, brown, often surrounded by a slimy
sheath......................................................
A nthostomella

ACKNOWLEDGMENTS
Dequn Zhou and Bing-Sheng Lu would like to thank The
University of Hong Kong for the award of Postgraduate Studentships.
We are grateful to Ms Helen Leung for technical assistance. Dr. Eric
McKenzie is thanked for serving as pre-submission reviewer.
REFERENCES CITED
Francis, S.M. (1 975). Anthostomella Sacc. (Part I). Mycological Papers
139: 1-97.
Hawksworh, D.L., Kirk, P.M., Sutton, B.C. and Pegler, D.N. (1995).
Dictionary of the fungi. 8th ed. Wallingford, UK, CAB
International, 6 16p.
Hyde, K. D. (1995). Fungi from palms. XVII. The genus Fasciatispora,
with notes on Amphisphaerella.Nova Hedwigia 6 1: 249-268.
Hyde, K.D. (1996). Fungi from palms. XXVI. The genus
Anthostomella, with ten new species. Nova Hedwigia 62: 273340.
Kirschsten, W. (1934). Remarks on a collection of British species of
Rosellinia and a redistribution of the species of that genus.
Transactions of the British Mycological Society 18: 302-307.
Lu, B.S., Hyde, K.D. and Ho, W.H. (1998). Spirodecdospora gen. nov.
(Xylariaceae, Ascomycotina) from bamboo in Hong Kong.
Fungal Diversity 1: 169-177.

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