Thai Forest Bulletin

Thai Forest Bulletin 

(Botany) 

No. 40 November 2012 ISBN 0495-3843 

The Forest Herbarium 

Department of National Parks, Wildlife and Plant Conservation 

Bangkok, THAILAND

Thai Forest Bulletin (Botany) No. 40, 2012 

CONTENTS 

Professor Kai Larsen (1921–2012) 1 

Obituary: Kai Larsen (1921–2012) 2–3 

In Memorium: Cees (‘Kees’) Berg, July 2 nd 1934 – August 31 st , 2012 4–8 

David J. Middleton & Pramote Triboun. Somrania, a new genus of Gesneriaceae 

from Thailand 9–13 

Stuart Lindsay, Thamarat Phutthai, Kitichate Sridith, Sahut Chantanaorrapint 

& David J. Middleton. Actinostachys wagneri (Schizaeaceae), a new record for Thailand 14–16 

Wittaya Kaewsri. Amomum tomrey Gagnep. (Zingiberaceae), a new record for Thailand 17–19 

Willem J. J. O. De Wilde & Brigitta E. E. Duyfjes. Revision of Cyclocodon Griff. ex 

Hook.f. & Thomson (Campanulaceae) 20–25 

Yaowanit Tarachai, Pornwiwan Pothasin, Wattana Tanming & Stephen G. Compton. The 

distribution and ecology of the purple form of Ficus montana in western Thailand 26–30 

Chalermpol Suwanphakdee, David A. Simpson & Pranom Chantaranothai. Three new 

species of Piper (Piperaceae) from Thailand 31–37 

Willem J. J. O. De Wilde & Brigitta E. E. Duyfjes. The lesser-sized Lobelias of Asia 

and Malesia 38–56 

Manop Poopath, Duangchai Sookchaloem & Thawatchai Santisuk. The Dipterocarpaceae 

of Hala-Bala Forest Complex, Narathiwat and Yala Provinces, Peninsular Thailand 

Stuart Lindsay, David J. Middleton & Piyakaset Suksathan. A new species of 

57–101 

Rhachidosorus (Rhachidosoraceae), a genus new to Thailand 102–104 

Jarearnsak Sae Wai. Orchipedum Breda (Orchidaceae, subfam. Orchidoideae), a new 

generic record for Thailand 105–107 

Hubert Kurzweil & Saw Lwin. New records in the orchid flora of Myanmar 108–113 

Sahut Chantanaorrapint & Amonrat Chantanaorrapint. A new species record of Sciaphila 

(Triuridaceae) for Thailand 114–117 

Paul Ormerod, Sahut Chantanaorrapint & Hubert Kurzweil. Cephalantheropsis longipes 

(Orchidaceae), a new record for Peninsular Thailand 118–120 

James A. Wearn & Charan Leeratiwong. A reassessment of the identity and rarity of 

Clerodendrum chlorisepalum (Lamiaceae) in Thailand and Vietnam 121–124 

Pakorn Tippayasri & Chatchai Ngernsaengsaruay. Coelogyne phuhinrongklaensis 

(Orchidaceae), a new species for Thailand 125–129 

Atchara Teerawatananon & Sarawood Sungkaew. Tripogon purpurascens (Chloridoideae: 

Poaceae): a native Thai grass recently recognized 130–133 

Phongsak Phonsena, Pranom Chantaranothai & Amornrat Meesawat. Two new records 

of Xyris L. (Xyridaceae) for Thailand 134–140 

Kamolhathai Wangwasit, Khanit Wangwasit & Pranom Chantaranothai. Fimbristylis 

pubisquama (Cyperaceae), a new record for the Flora of Thailand 141–143 

Printed by: Office of National Buddishm Press 

314-316 Bamrungmung Rd., Pomprabsatrupai, Bangkok, Thailand 

Tel : 0 2223 3351, 0 2223 5548 Fax: 0 2621 2910 

Page

Prof. Kai Larsen, the Danish co-editor of the 

Flora of Thailand, established the Flora of Thailand 

project in 1965 after the fi rst joint Thai-Danish 

Botanical expedition in 1958, organized as a cooperative 

effort with Prof. Tem Smitinand, the then 

director of the Forest Herbarium (BKF). Prof. Larsen 

found that it was time for Thailand to have a contemporary 

Flora to update the only existing work 

covering Thailand’s plants - the ‘Flora Siamensis 

Enumeratio’ written by W.G. Craib and A.F.G. Kerr 

in 1931. He contacted renowned botanists, mainly 

from prestigious botanical institutes in Europe, i.e. 

Aarhus, Copenhagen, Edinburgh, Kew, Leiden, and 

Paris, whom he knew, to join the Flora of Thailand 

project on a voluntary basis to pursue the production 

of taxonomic treatments of Thai vascular plant 

families. The editors and editorial board members 

of the Flora of Thailand were comprised of K. 

Larsen and Tem Smitinand as co-editors, B. Hansen 

as assistant editor and Chamlong Phengklai, R.C. 

Bakhuizen van den Brink Jr., B.L. Burtt, L.L. 

Forman and J.E. Vidal as editorial members. Prof. 

Larsen dedicated himself to the Thai flora, and 

tirelessly conducted many Thai-Danish botanical 

expeditions in remote areas throughout Thailand. 

Despite the difficult and discouraging access to 

botanical sites, a great number of Thai plant collections 

were made and deposited in the herbaria of 

Aarhus University and the Forest Herbarium, 

Royal Forest Department in Bangkok. Together 

with BKF staff, Kai managed to conduct botanical 

collections in the once luxuriant forests, and he 

became aware that the Thai fl ora encompassed far 

more undescribed species than expected. When it 

became more and more diffi cult to recruit foreign 

specialists to work for the Flora of Thailand project, 

he therefore encouraged a new young generation of 

Thai botanists to continue their own Flora. The 

Flora of Thailand can be proud that a large number 

of families have been revised by local botanists - 

more than for any other tropical Flora. Prof. Larsen 

succeeded in securing substantial aid from the 

Carlsberg Foundation and DANIDA, the Danish 

Development Aid Programme, for botanical expeditions, 

printing costs, and, most importantly, 

fi nancial support for the higher education of promising 

young Thai botanists in Denmark. Some have 

already got their Ph.D. degrees, while many received 

support for short or long periods of study, 

and this lengthy academic collaboration has continued 

to the present day. It was encouraging to see 

that several young Thai botanists were successfully 

PROFESSOR KAI LARSEN (1921–2012) 

educated and trained abroad, and plant taxonomy 

in Thailand was recognized and promoted. Prof. 

Larsen knew the history, as well as the present and 

the future vision of the Flora of Thailand when he 

saw so many young Thai botanists working on the 

Thai fl ora. He encouraged them to assure that the 

Flora of Thailand Project would be best in the hands 

of the younger generation. Due to his initiatives, 

and his remarkable work on the Thai fl ora in collaboration 

with BKF staff, the Flora of Thailand 

has reached an international standard and recognition, 

and serves as the base for fl oristic study. The 

Forest Herbarium is also known as a national herbarium 

well equipped with knowledgeable and 

well-trained botanists, herbarium specimens and 

botanical references. 

Since the beginning of the Flora of Thailand 

Project, Prof. Larsen took 40 years to do this indispensable 

task - not only writing the Flora on his own 

but also recruiting family experts for contributing to 

the Flora. In the early years, the Flora of Thailand 

progressed slowly, but at present the treatments of 

the remaining families are not far from completion. 

Prof. Larsen had sat as Chair of the Flora of Thailand 

editorial board meeting since the fi rst meeting till 

the fourteenth meeting in Copenhagen, but we were 

very regretful that he could not join the latest meeting 

in Chiang Mai in 2011 because of his health 

(though he gave a videotaped presentation. At the 

Flora of Thailand Board meeting he always brought 

up questions to be discussed including the possibility 

of the next meeting. Among other future initiatives 

to be considered by the Board, was a closer collaboration 

between the organizers of the Flora of 

Thailand and the other projects such as the Flora 

Malesiana and Flora du Cambodge, du Laos et du 

Viêtnam; and through his knowledge of experts, the 

Flora of Thailand has become a strong team to work 

on the Thai Flora. 

Prof. Kai Larsen devoted his life to the work 

on the Flora of Thailand, particularly on his most 

favourite family the Zingiberaceae, until his last 

years. He served as co-editor of the Flora of 

Thailand from the very fi rst volume until he passed 

away in 2012. He will always be remembered by 

Thai colleagues, and without his initiative and help 

the Flora of Thailand would have not materialised. 

He has done a great job for the Thai fl ora, and his 

name will forever remain in the history of the Flora 

of Thailand as one of the great Danish botanical 

explorers of the Thai fl ora. 

Thawatchai Santisuk & Kongkanda Chayamarit 

1

2 

Kai Larsen was born on 15 th November 1926 

in Hillerød, a small town north of Copenhagen 

with some 30,000 inhabitants and mostly know for 

being the home of many Danish kings who inhabited 

the Frederiksborg Castle. His father, Axel Georg 

Larsen, owned a book-binding business and Kai 

Larsen was brought up in the proud tradition of 

Danish craftsmanship in which high quality and 

perfection are key-words. He graduated from 

Frederiksborg Gymnasium (Highschool) in 1946 

and initiated an academic career with studies at the 

University of Copenhagen from which he graduated 

as Candidatus magisterii six years later with a major 

in botany (1952). Following his graduation he 

continued pursuing his academic career as a research 

assistant and teacher at the University of 

Copenhagen, until he became associate professor 

at the Farmaceutical University in Copenhagen in 

1962. One year later, in 1963,he became the fi rst 

professor of botany at Aarhus University and he 

was charged with building up a Botanical Institute 

at the university. Until then the only 30 years old 

university had not trained biologists, so it was Kai 

Larsen’s duty to build up research and teaching in 

botany to underpin the biology study. The city of 

Aarhus supported the young university in many 

ways and in general the economic climate was favorable 

during the high times of the 1960s. In a few 

years Kai Larsen was able to build up an institute 

OBITUARY: KAI LARSEN (1921–2012) 

Henrik Balslev 

that encompassed 2000 m2 of very beautiful greenhouses 

in the municipal botanical garden, a new 

herbarium, and many laboratories for a wide range 

of botanical studies including limnology, cytology, 

anatomy etc. Since then these various sections of 

the institute have grown and become important internationally, 

nationally and locally. The herbarium 

of Aarhus University is now widely known internationally, 

not least for its collections of tropical 

plants including very important collections from 

Thailand. The green houses are a major attraction 

for the public and exhibits 8000 plants from all 

over the World, and next year a new tropical palm 

house will be inaugurated. 

Kai Larsen was a prolifi c teacher. During his 

many travels he lectured about his research all over 

the World. In SE Asia he tutored many young botanist, 

especially from Thailand. Many of them spent 

prolonged visits and some of them took their degrees 

at Aarhus University. For his Danish students 

he will mostly be remembered for his taxonomy 

classes which were always meticulously prepared 

and elegantly delivered. For lack of a good text Kai 

Larsen wrote a Danish language book ( (Kormfyternes 

Taxonomi, 1974) which precisely summarized the 

most recent classifi cation of vascular and especially 

fl owering plants at the time based on the systems of 

Takhtajan and Cronquist.

Kai Larsen was much respected for his contribution 

to tropical botany and taxonomy. He was 

elected member of the Royal Danish Academy of 

Sciences and Letters (1979) and The Royal 

Norwegian Society of Sciences and Letters. 

Kai Larsen’s scientifi c career was as a plant 

taxonomist. His fi rst expedition was to Greenland 

when he was still a student, and his earliest publications 

are about Greenland’s fl ora. During his 

early years at the University of Copenhagen he was 

asked to administrate and lead the First Thai- 

Danish Expedition which was held during the years 

1958-59. This expedition was the initiative of the 

then Danish Ambassador to Thailand, Gunnar 

Seidenfaden, who was a keen botanist in his spare 

time with an immense knowledge of the more than 

one thousand species of Thai Orchids. The expedition 

explored the Malay Peninsula, eastern 

Thailand, and the mountains in the north. During 

the expeditions Kai Larsen and the other Danish 

members of the expedition established close working 

relationships with the Thai botanists, not least 

the then director of the Forest Herbarium in 

Bangkok, Tem Smitinand. Two shorter expeditions 

(1961-62; 1963) followed and contributed to further 

tighten the bonds between Thai and Danish 

botanists. In 1965 Kai Larsen invited a group of 

leading European and Thai botanist to a meeting at 

Kew where it was decided to initiate the Flora of 

Thailand d project. This was to become Kai Larsen’s 

largest scientifi c undertaking. Since its initiation 

Kai Larsen has contributed treatments of 44 families 

to the fl ora and has participated in all biennial 

Flora of Thailand d meetings, except the last one 

which was held in Chiang Mai in November of 

2011. Kai Larsen was editor of the Flora of Thailand 

together with Tem Smitinand from 1970 when the 

fi rst volume appeared until 2009 when part 29 had 

been published. Since then Kai Larsen has been 

editor together with Thawatchai Santisuk. This 

Thai-Danish undertaking with collaboration from 

many international specialists, is now very fi rmly 

established and the termination of the project is 

within reach in the coming years. As mentioned, 

Kai Larsen published taxonomic accounts of a 

great number of plant families, but his favorite 

families were the legumes and the gingers. In these 

families he has contributed a vast number of publications, 

not only about their Thai species, but treating 

species from all over South East Asia. For instance, 

he has published many shorter journal 

papers about new gingers and also collective books 

on Malay gingers (1999) and a treatment of the 

Chinese gingers in Flora of China together with 

Wu (2000). For the legumes he wrote the 

Caesalpinoid genera for r Flora of Thailand d and also 

for Flora Malesiana (with Ding Hou 1996), and he 

co-authored the entire volume on legumes for the 

Flora of China (with Wu, Raven and Hong, 2010). 

Kai Larsen was honoured by his colleagues who 

named many plant genera after him. Kailarsenia 

and Larsenaikia are two genera in the Rubiaceae; 

Kaisupeea – a gesneriad – honours both Kai Larsen 

and his wife Supee S. Larsen; Larsenianthus is a 

genus in the ginger family that he loved so much. 

Many species were also named for him: Bauhinia 

larsenii is a fossil legume from China; Impatiens 

larsenii is in the Balsaminaceae; Cassytha larsenii 

in Lauraceae; Crotalaria larsenii is a legume; 

Primula larsenii in the Primulaceae; Lasianthus 

larsenii and Mouretia larsenii both in the 

Rubiaceae; and fi nally Caulokaempferia larsenii, 

Cornukaempferia larsenii, Curcuma larsenii, 

Hedychium larsenii, Kaempferia larsenii and 

Zingiber larsenii all honours his great contribution 

to the taxonomy of Zingiberaceae. 

Kai Larsen was the grand old man in tropical 

botany. His contributions were many and encompassed 

institution building, teaching, research, tutoring 

young botanists, and he also participated in 

the work of international organisations such as 

International Association of Botanical Gardens. He 

was a true leader and highly respected, not least in 

Thailand where he was always treated with great 

admiration. On the 1st t of December 1996 Kai 

Larsen retired from his professorship at Aarhus 

University. But he continued his work as professor 

emeritus and came in to work several days a week. 

He was involved in many activities, both his own 

research and also representing Aarhus University 

in various international projects such as Flora 

Nordica and Flora of China. And he remained 

close to his love for the Flora of Thailand d and 

made many journeys and participated in many 

meetings. He organized the 14th Flora of Thailand 

meeting in Copenhagen in August of 2008. After 

that his health prevented him from travelling more 

to Thailand but his interest was just as vivid as it 

had always been. He delivered his talk to the 15th Flora of Thailand d meeting in Chiang Mai as a vid- 

eotaped presentation, which was his last public engagement 

with the project. On 23rd d of August 2012 

he passed away surrounded by his wife and 

colleagues. 

3

4 

IN MEMORIAM: CEES (‘KEES’) BERG, JULY 2 nd 1934 – AUGUST 31 st , 2012 

Dr Cigaro collecting a Marcgraviaceae in Ecuador (1977) – Paul Maas 

Cornelis Christiaan Berg, better known as 

Cees (or ‘Kees’ for the non-Dutch) Berg, was born 

on July 2 nd , 1934 in the city of Bandung on Java, 

then still the Netherlands East Indies. Later, the 

family moved to Sumatra, close to the city of 

Medan. During the Second World War, when Japan 

attacked the Netherlands East Indies, Cees’ father 

was enlisted and, unfortunately, did not survive the 

war. Cees, his four brothers and their mother were 

interned in a women’s camp near Medan, but at the 

age of 10 Cees was moved to a men’s camp. This 

period must have been very traumatic as Cees, after 

his release, spoke little for a long time. All brothers 

and their mother survived the war, but shortly after 

the liberation their mother died of hunger oedema. 

The fi ve orphans came to the Netherlands, where 

they were split over two foster families. Cees, 

together with his youngest brother, came into a 

household with two daughters, and during holidays, 

both families joined so that the brothers were 

united. Together with the eldest daughter, Cees 

started to explore the forests close to his house. 

Both studied in Utrecht, Cees majoring in biology 

and his foster sister studying nursing. ― Peter van 

Welzen 

Utrecht, The Netherlands. As a student, Cees 

developed an interest in experimental taxonomy, 

more specifi cally in the study of polyploidy complexes. 

He focused his attention to the cytotaxonomic 

study of two intricate species complexes, 

Cardamine pratensis s.l. and Myosotis palustris s.l. 

and used the technique of experimental cultivation 

for the investigation of genetic differences among 

populations. He collected many samples of these 

two species complexes. Later he supervised a cytotaxonomic 

study of Dorstenia (Moraceae) by one 

of his students. ― Theo Gadella 

About 50 years ago, Cees Berg and I had in 

the Utrecht Herbarium (Netherlands) a meeting 

with its director Prof.dr. J. Lanjouw. We both very 

much wanted to do a PhD study in plant taxonomy 

and Lanjouw suggested to do it within the framework 

of the Flora Neotropica Project, which was 

just about to start at that time. Lanjouw proposed 

as subjects for our study the families Moraceae or 

Zingiberaceae. Cees choose Moraceae, and I myself 

Zingiberaceae. At that time, we had no idea 

that we would work on those plant families for the 

length of our whole career. Cees and I spent several 

years in completing our respective Flora Neotropica

treatments and, as was still customary at the time, 

we conducted our work using herbarium material 

only - fi eld work was not necessarily considered to 

form an essential part of taxonomic studies. We 

both defended our PhD thesis on the same day, the 

4th of May 1973. 

After that period we both worked as Staff 

members at the Utrecht Institute involving, next to 

continuing taxonomic study of “our” respective 

families, a great deal of teaching and training 

young students in taxonomy. I particularly remember 

the course on the Dutch Flora we were giving 

each year, culminating in the week in the southern 

Dutch province of Limburg, walking through the 

nice and varied landscapes with many students, 

and fi nishing the days in the evenings enjoying excellent 

Limburgian beer in bar “De Kroon” in 

Gulpen together with Frits Jonker, Ad de Roon, 

Lubbert Westra, Carolien de Wal, and many other 

Utrecht Staff members. 

During that period, Cees did not feel overly 

enthusiastic about going into the fi eld. I remember, 

however, that after spending a year in Amazonian 

Brazil (1971) and joining several expeditions led 

by G.T. Prance all over the Amazon region, I suggested 

to Cees to do the same. He then agreed and 

went to Brazil, only to become completely “lost” 

after that. Many visits to Tropical America followed 

in order to study Moraceae and Cecropiaceae 

(a family that Cees newly described) in the fi eld, 

and also combining this with visits to numerous 

Neotropical herbaria. 

In 1977 we went into the fi eld together, visiting 

Panama where we were guided by our fantastic 

guide and dear friend Dr. Bob Dressler. Then we 

continued our trip to Ecuador, a country with many 

interesting Moraceae, Cecropiaceae, Urticaceae 

(and Zingiberaceae). There we made some very 

nice fi eld trips into the Oriente. But not all went 

well, as on one day we suddenly came to a halt 

because of a landslide. With Cees driving that day, 

we were forced to continue through the rubble with 

our car, but we had to pay dearly when suddenly a 

large piece of rock came down on top of our roof. 

To make things worse, an axle on the left side of 

our car broke down. There we were stuck –and 

what to do? One of our team members, Ben ter 

Welle, arranged for a truck and managed to bring 

the car back to Quito. Of course, Cees and I 

continued collecting in the meantime! But then we 

got a big fi nancial problem as the insurance did not 

cover expenses made for trips to the Amazon. So 

an emergency call to the Netherlands was necessary, 

to see if someone (my wife) could send 1000 

dollars to us. Cees during his fi eld work always had 

one major problem: he could not live without coffee 

and….cigars. I remember that whenever he was 

out of cigars he used to ask everyone in the villages 

we came through to supply him with a few cigars 

(which was not always that easy…). One of his 

Ecuadorian colleagues, the late Dr. Jaime Jaramillo, 

very aptly nicknamed Cees Doctor Cigaro. 

With Cees one of the Last Mohicans passed 

away. Throughout his long career he gained a vast 

and unparalleled knowledge of the huge family of 

Moraceae, and of the genus Ficus in particular. He 

could identify even the tiniest leaf fragments. It is 

unlikely that this achievement is ever going to be 

equalled by anybody, and particularly so in a time 

that favours short-time projects rather than longterm 

work so very much essential for understanding 

large plant families. We all shall miss Cees very 

much. ― Paul Maas 

Projeto Flora Amazônica and The New York 

Botanical Garden. Kees Berg’s long relationship 

with The New York Botanical Garden (NYBG) 

began in 1977 when he participated in one of the 

two parallel expeditions that inaugurated Projeto 

Flora Amazônica (PFA), part of Brazil’s ambitious 

plan to document its vast plant diversity. Over 

more than ten years, most of the fi nancial support 

for PFA came from the U.S. National Science 

Foundation, and the non-Brazilian participation 

was coordinated by Ghillean Prance, curator and 

later Vice President for Botanical Science at 

NYBG. Kees’s expedition included Prance, Antôno 

Sérgio da Silva (Brazilian counterpart), Michael 

Balick (then a graduate student and now director of 

the Institute of Economic Botany at NYBG), Bruce 

W. Nelson (now a researcher at INPA in Manaus), 

and two tree-climbing mateiros or woodsmen, 

Mario R. dos Santos and Raimundo P. Bahia (wellknown 

as “Doca”). During October–December 1977, 

the expedition collected plants in the Serra dos 

Carajás, along the Transamazon Highway, around 

Tucuruí, in the Serra do Cachimbo, and along the 

Santarém-Cuiabá Highway. The expedition produced 

1921 numbers, including 100 collections of 

palms. 

5

6 

For more than 40 years, Kees was appreciated 

by his colleagues at NYBG for sharing his rich 

knowledge of botany by identifying specimens, 

training students, publishing monographs, and 

contributing treatments of his plant families to 

many fl oristic projects. Probably his most important 

contribution to NYBG and to the botanical 

community overall were his treatments in his 

groups of expertise for these monographic and fl oristic 

projects, among others: 

• Flora Neotropica Monographs for the 

Organization for Flora Neotropica, based at NYBG: 

Olmedieae and Brosimeae (Moraceae) in 1972; 

Coussapoa and Pourouma (Cecropiaceae) in 1990; 

Moreae, Artocarpeae, and Dorstenia (Moraceae) in 

2001; and Cecropia (Cecropiaceae) in 2005; he 

submitted his treatment of Ficus to Flora Neotropica 

just before his death. 

• The Moraceae, Cecropiaceae, and Urticaceae 

for the Guide to the Vascular Plants of Central 

French Guiana. 

• Those families plus the Ulmaceae for the 

First Catalogue of the Flora of Acre, Brazil. 

Another service that Kees provided to NYBG 

botanists was the identifi cation of nearly all of our 

collections of Moraceae (including Cecropiaceae), 

Urticaceae, and Ulmaceae, which has greatly increased 

the scientifi c value of our collections. 

NYBG’s institutional data-base has over 3200 

identifi cations made by Kees over the years, but 

unquestionably the NY herbarium contains many 

more. 

Curators at NYBG are grateful for the nearly 

40 years of collaboration with Cornelis Berg, not 

only for the improvements he made to our collections 

and the intellectual contributions he made to 

our publication program, but also because of the 

close friendships that he maintained with many of 

our staff. He will be sorely missed, but he will continue 

to inspire everyone here who had the privilege 

of knowing him. ― Douglas Daly and Scott 

Mori 

Bergen, Norway. (A second phase in Cees his 

career was his professorship in Norway.) One of 

the fi rst things Kees said to me in the process leading 

to his employment at Milde (Norway, near 

Bergen) in 1985 was: “I am an old tree, and they 

are diffi cult to transplant.” Nevertheless the transplantation 

took place, and as predicted, he had a 

rather diffi cult process adapting to the new environment 

and the new language, but he took that 

challenge with restraint. 

Frankly, the conditions at Milde were far 

from ideal for research of the tropical genus Ficus 

and its relatives. Nevertheless, Kees continued to 

work indefatigably on this enormous and very 

complicated group of plants, which are so important 

in the tropics, and managed by a generous gift 

from our benefactor, Bjarne Rieber, to establish a 

greenhouse to grow them. He had about 200 different 

species in cultivation. 

This being solved, a much more diffi cult task 

remained: to adapt to the rather complicated management 

of the organisation, one which was an enduring 

mystery to him, and where I was given the 

task to assist him. We had many long discussions, 

and somehow found ways through this jungle. I 

cannot claim they were easy talks, but Kees had an 

unusually friendly persistence, which I liked, so we 

never really clashed, even when we disagreed. 

I particularly remember that, to my surprise, 

he was very keen on establishing a collection of 

native Norwegian trees, an idea that had not previously 

crossed our minds - we were primarily engaged 

in fi nding foreign woody plants for 

Norwegian gardens. But certainly he was right: 

The Norwegian Arboretum should also take an interest 

in our native trees! 

He also engaged in the relationship with the 

Friends of the Arboretum and started nearly immediately 

to write a Newsletter which under his 

successor Per H. Salvesen has developed into the 

important journal Årringen, issued yearly, where 

we present results from our collections to the 

general public. This close relation to the Friends 

led to the establishment of a heather garden, which 

was donated in 1996, and which still is run by the 

Friends. This garden was an enjoyment to him, as 

well as to all of us. 

After his retirement in 2005 and return to the 

Netherlands, he was a frequent visitor to Bergen - 

surely the old tree had developed some Norwegian 

roots. He usually came about midsummer, when 

also visiting his daughter Hendrieke at Voss. He 

then fi lled our tables with specimens that are in the 

BG herbarium (2,000–3,000 specimens), which 

now houses a comprehensive, well-identifi ed collection 

preserved for future generations to study

- of particular importance since the tropical forests 

are disappearing quickly. 

But this summer in 2012 we missed him, he 

was too ill to travel, though he still steadily worked 

on the task to revise Ficus. In his last letter, which 

we received a few weeks before his death, he was 

concerned about a loan, which was needed quickly 

for drawings (by his daughter Hendrieke) for a paper 

he had nearly fi nished, since his days were numbered. 

He was certainly one of the most dedicated 

and industrious botanists I have known. 

The old tree has fallen, but the seeds he 

spread, will grow! ― Per M. Jørgensen 

Leiden, The Netherlands. Back in the Netherlands 

Cees, of course, continued his work on Moraceae. 

The fi rst time I met him in the Leiden herbarium, I 

vaguely remembered that I had seen him before, 

when I was still an M.Sc. student, working on the 

ecology of tsetse fl ies in Ivory Coast. My supervisor 

was Prof.dr. Koos Wiebes, a specialist in fi g 

wasps. Wiebes announced that he would come for 

fi eld work to the Ivory Coast and while driving to 

the expedition site he would visit me and he would 

be accompanied by a colleague, which indeed was 

Cees. Together they worked on the interaction 

between fi gs and wasps, resulting in a book about 

the African species (Berg & Wiebes, 1992). Leiden 

is for many taxonomists synonymous with Flora 

Malesiana. However, Asia had never been a focus 

for Cees, probably because another world expert 

on Moraceae had worked there, Corner in 

Singapore. Corner had produced a manuscript on 

the Malesian Moraceae, but disagreement with Van 

Steenis (editor of Flora Malesiana) about several 

species concepts stalled publication of the 

Moraceae. We were happy that we could interest 

Cees in revising the Malesian Moraceae, using 

Corner’s manuscript as a basis. He consulted thousands 

of specimens during a few sabbatical periods, 

and quickly produced two big volumes, one on 

Ficus (2005) and one on the remaining Moraceae 

(2006). The editors of local Asian Floras now 

became aware of Cees his knowledge and invited 

him to help with their Flora treatments as well. He 

quite liked working on the Thai Moraceae. This 

took quite some time, but it allowed him to visit the 

country several times and at the time of his death 

Cees was still supervisor of a Thai PhD student 

working on a group of fi gs. Cees only worked with 

a morphological species concept, but he was happy 

with the results of molecular work (though those 

results also synonymised his Cecropiaceae with the 

Urticaceae), and he collaborated closely with Finn 

Kjellberg’s group in Montpellier (France). Cees 

was certainly a person who liked to travel; he visited 

and did fi eld work in almost all the tropical 

countries of Africa and South America. In Asia, he 

visited southern China and Thailand. Only after 

retirement did he return once to his land of birth, 

now called Indonesia. ― Peter van Welzen 

Collecting in Acre, Brazil. The state of Acre, 

Brazil in Southwestern Amazonia is honoured to 

have been the site of Dr. Cornelis C. Berg’s last 

botanical expedition in April, 2010. Given the high 

diversity of Moraceae in Acre, he was one of the 

key botanists invited to participate in the Mobilizing 

Taxonomic Specialists for Acre project, developed 

by the collaborative research program between the 

New York Botanical Garden and the Universidade 

Federal do Acre (UFAC) to advance our knowledge 

of that region’s most important plant groups. 

He joined the fi eld team of UFAC’s 

Laboratório de Botânica e Ecologia Vegetal, which 

at the time consisted of Flávio Obermuller, Marcos 

Silveira, Herison Medeiros, Wendeson Castro, 

Edilson Consuelo de Oliveira, Lívia Souza and 

Heloisa Polary. The group visited diverse localities: 

the Riozinho do Andirá, Seringal Cachoeira, 

Fazenda Catuaba, Reserva Florestal Humaitá, Rio 

Iquirí, and the Vila do V in six municipalities (Rio 

Branco, Sena Madureira, Bujari, Porto Acre, 

Senador Guiomard and Xapuri) in the eastern part 

of the state. The expedition produced more than 

400 collections, the vast majority of them in the 

families of Dr. Berg’s expertise. Before and after 

the fi eld work, he annotated virtually all the UFAC 

herbarium’s specimens in those plant groups. 

His work contributed signifi cantly to our 

knowledge of the Acre fl ora, adding two new genus 

records and 25 new species records for the state, 

and among those eight new records for Brazil. 

Undoubtedly, if he had had more time, he would 

have added a number of species new to science 

from the Acre fl ora, considering that 18 or nearly 

half of his Ficus collections from that trip remained 

undetermined. 

During his brief visit to Acre, ”Berg” (as he 

was dubbed there) made a lasting impression on 

everyone he met. He will be remembered in Acre 

with fondness and great respect. ― Flávio 

Obermüller and Douglas Daly 

7

8 

Molecular work? Cees was sceptical about the 

ability of DNA-sequence based phylogenies to 

uncover the true relationships, but - as always - welcoming 

anybody who wanted to contribute to our 

understanding about any aspect of Ficus. Embarking 

on a quest for discovering the global phylogeny of 

Ficus, I fi rst contacted Cees in 2002 to ask if he 

would support my funding applications as an 

expert of the classifi cation of Ficus. He kindly 

wrote back to me on the same day and offered me 

his assistance, as well as access to his extensive 

living collections in Milde, which I visited for a 

pleasant couple of days in 2003. I since met him 

several times when he visited the herbarium in 

Kew and Minnesota where I worked during my 

postdoctoral years. He was always very helpful on 

checking my identifi cations and commenting on 

the results of the phylogenetic analyses, and he 

always had time for a nice meal, and a good beer 

and a chat about future research needs. He was particularly 

happy about the possibility of the DNAbased 

work to enlighten the large and diffi cult 

Neotropical section Americana, which he was in 

the process of revising until his death. However, he 

was very sceptical about some of the infrageneric 

relationships suggested by the molecular data, and 

wondered about the limitations and methodological 

errors. When offered co-authorships for his 

help and comments on the fi rst global phylogeny 

published in 2005, he kindly refused with a smile 

and a twinkle of his eye, because he would rather 

keep the right to criticise it afterwards. In Flora 

Malesiana he largely ignored the molecular suggestions 

of relationships, I guess he found it too 

diffi cult to decide which of the results were well 

supported and which were only preliminary 

Cees in the herbarium of the Queen Sirikit Botanical Garden, 

Chiang Mai, Thailand. – Rachun Pooma 

hypotheses - surely enough we are still struggling 

to sort out the infrageneric relationships of Ficus 

and even the origin of the fi gs based on molecular 

data. Berg was a great morphologist and has 

inspired us to pursue the molecular work in even 

more detail - especially when DNA suggests obvious 

confl icts with relationships well supported by 

morphology. As Berg said, “it has to make sense, 

you can’t just say that swallows are not birds” and 

so the white crane has himself departed, but his 

love of Ficus, supportive attitude, and persistence 

and insistence on making sense of it all continues. 

― Nina Rønsted 

Honours. We knew Cees as a silent, hard-working 

man who never took part in social events like coffee 

breaks or drinks. Till his death, he tried to visit 

Leiden as much as possible, but like in Norway, he 

missed his last appointment. We also treasure him 

as a very friendly and helpful person. Although he 

had no teaching obligations in Leiden, he still 

motivated students to help revise the species of the 

Solomon Islands. We admire the way in which he 

made all arrangements for after his death: the list of 

persons who should be notifi ed, who should ‘clean’ 

his desk, etc. Cees was really a banyan tree among 

taxonomists and he will be dearly remembered. 

One Moraceae was named in honour of Cees: 

Dorstenia bergiana Hijman. Cees newly described 

or made new name combinations for 318 taxa in 

the Moraceae and Cecropiaceae and for one 

Boraginaceae, a subspecies of Myosotis, M. palustris 

(L.) Nathh. subsp. nemorosa (Besser) C.C.Berg 

& Kaastra, a result of his polyploidy interest. ― 

Peter van Welzen 

Two giants in Chiang Mai, Thailand. – Rachun Pooma

THAI FOR. BULL. (BOT.) 40: 9–13. 2012. 

Somrania, a new genus of Gesneriaceae from Thailand 

INTRODUCTION 

This contribution arose from specimen and 

fi eld based taxonomic work on the Gesneriaceae 

towards an account of the family for the Flora of 

Thailand. We present a new genus of Gesneriaceae 

with two hitherto undescribed species. The genus 

fi rst came to our attention when seeds of an 

unknown species of Gesneriaceae from Ranong 

Province in Thailand collected by a joint Royal 

Botanic Garden Edinburgh/Forest Herbarium 

Bangkok expedition were grown at the Royal 

Botanic Garden Edinburgh. When this plant fl owered 

it could not be named to genus or species. A 

second undescribed species was later collected in 

Phangnga Province in Thailand and is clearly related 

to the Ranong plant but also quite clearly not 

the same species. We describe these two species in 

the new genus Somrania D.J.Middleton, named in 

honour of Dr Somran Suddee of the Forest 

Herbarium Bangkok. 

Somrania is most similar to Damrongia Kerr 

ex Craib and, to some extent, to Loxocarpus R.Br. 

Damrongia has recently been resurrected from 

synonymy of Chirita Buch.-Ham. ex D. Don 

(Weber et al., 2011). Loxocarpus was until recently 

included in synonymy of Henckelia Spreng. 

(Weber et al., 2011; Kiew & Lim, 2011). Despite 

their straight fruits these plants belong in the twistedfruited 

group of Old World Gesneriaceae (Möller 

et al., 2009; Weber et al., 2011). The most easily 

DAVID J. MIDDLETON* & PRAMOTE TRIBOUN** 

ABSTRACT. The new genus Somrania D.J.Middleton is described along with two new species, Somrania albifl ora D.J.Middleton 

and Somrania lineata D.J.Middleton & Triboun. 

observable difference between Somrania, on the 

one hand, and Damrongia and Loxocarpus, on the 

other, is in the presence of branched hairs in 

Somrania. These hairs are completely lacking in 

the other two genera, which have their own characteristic 

indumentums of dense multicellular uniseriate 

hairs in both genera (which is generally brown 

in Damrongia and silvery in Loxocarpus). In 

Somrania albifl ora D.J.Middleton the branched 

hairs are large and cover all vegetative parts and 

the infl orescence axes; in S. lineata D.J.Middleton 

& Triboun the branched hairs are smaller and are 

not found on the infl orescence axes. The only other 

Asian species of Gesneriaceae with branched hairs 

of which we are aware are Paraboea tarutaoensis 

Z.R.Xu & B.L.Burtt and Paraboea vulpina Ridl. 

(Xu et al., 2008). In addition the corolla of 

Somrania is tubular and only slightly widened 

distally whilst that of Damrongia is infundibuliform 

and that of Loxocarpus fl at-faced or campanulate 

(Paraboea ( tarutaoensis and P. vulpina are also fl at- 

faced and differ in many other characters). Lastly 

Somrania and Loxocarpus species have plagiocarpic 

fruits whilst those of Damrongia are 

orthocarpic. 

A preliminary molecular phylogenetic study 

which includes Somrania albifl ora and S. lineata, 

and which will be published in due course, supports 

the conclusions presented here (Puglisi, pers. 

comm.). 

* Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, Scotland, U.K. Email: d.middleton@rbge.ac.uk 

** Bangkok Herbarium, Plant Variety Protection Division, Department of Agriculture, Chatuchak, Bangkok 10900, Thailand.

10 

Somrania D.J.Middleton, gen. nov. 

Similar to Damrongia and Loxocarpus but 

differs in the branched hairs on the vegetative parts 

(absent in Damrongia and Loxocarpus), and in the 

tubular corolla (infundibuliform, campanulate or 

fl at-faced in Damrongia and Loxocarpus). – Type: 

Somrania albifl ora D.J.Middleton. 

Lithophytic herbs, scapose or with a short 

fl owering stem. Leaves primarily arising at base of 

plant, more rarely opposite on short stem, weakly 

to strongly anisophyllous, petiolate; blade herbaceous, 

margin crenate or dentate, with 4–8 pairs of 

secondary veins, tertiary venation alternate percurrent, 

adaxial surface with acicular hairs and short 

glandular hairs, abaxial surface with branched 

hairs and shortly stalked or sessile glands. 

Infl orescences either scapose or on short stems, 

THAI FOREST BULLETIN (BOTANY) 40 

KEY TO THE SPECIES 

cymose. Calyx of 5 lobes almost free to the base. 

Corolla zygomorphic, tube cylindrical, limb 

2-lipped with 2 upper and 3 lower lobes, mouth 

weakly to strongly oblique. Stamens 2, inserted in 

lower half of corolla tube; fi laments straight; 

anthers adnate face to face; staminodes 3, equal or 

medial one shorter. Disk k present, margin crenate. 

Ovary 2-carpellate, unilocular; style straight or 

slightly curved; stigma with only the lower lobe 

developing, weakly and subequally laterally 

bilobed, lobes rounded. Fruit t a capsule, held at an 

angle to the pedicel, not twisted, fusiform, dehiscence 

loculicidal on upper surface only. Seeds numerous, 

unappendaged. 

Distribution.— Currently only known from 

two species on karst limestone in Peninsular 

Thailand. 

1. Corolla white throughout and with a slightly oblique mouth; branched hairs present on leaves and infl orescence axes 

S. albifl ora 

1. Corolla white with two orange-brown lines in the corolla and with a strongly oblique mouth; branched hairs on leaves but not on 

infl orescence axes S. lineata 

Somrania albifl ora D.J.Middleton, sp. nov. 

Differs from Somrania lineata in the 

branched hairs being larger and more frequently 

branched, the pure white corolla, and the less 

oblique corolla mouth. – Type: Thailand, Ranong, 

Ngao Waterfall National Park, Trail to waterfall, 

580 m alt., seed collected 20 Feb. 2006, cultivated 

as RBGE acc. no. 20060626, vouchered and 

chosen as type as Middleton 4216 6 (holotype E!; 

isotypes BKF!, E!). Fig. 1, 2A. 

Lithophytic or terrestrial herb, to 20 cm high; 

stem, when present, densely covered in frequentlybranched 

hairs. Leaves primarily arising at base of 

plant but sometimes there is a short stem with 

opposite leaves, these somewhat differing in size; 

petiole 3.5–14 cm long, densely covered by 

frequently-branched hairs; blade ovate, 1.8–16.4 x 

0.8–12 cm, 1.2–1.8 times as long as wide, apex 

acute to obtuse, base cordate, sometimes somewhat 

unequally so, margin coarsely and somewhat 

irregularly dentate, with 6–8 pairs of secondary 

veins, covered with a mixture of acicular and shorter 

glandular hairs above, densely covered with 

frequently-branched hairs all over beneath, more 

densely so on venation, and with minute shortly 

stalked to sessile glands. Infl orescences on short 

stems, 8.7–16 cm long, many-fl owered; bracts narrowly 

obovate, 2.5–5 x 0.7–1.2 mm, densely 

covered with branched hairs; peduncle 5–13.5 cm 

long, densely covered with large branched hairs; 

pedicels 5.5–7.4 mm long, glandular pubescent 

with occasional larger branched hair. Calyx tube 

very short, lobes 3.8–4 x 0.5–1 mm, narrowly 

triangular, apex acute, glandular pubescent with 

hairs of varying lengths. Corolla white throughout, 

with a tube which widens slightly and gradually 

towards lobes, weakly 2-lipped, mouth slightly 

oblique, lobes spreading, c. 13.5 mm long; tube 

9.5–10 mm long, glandular pubescent outside, 

glabrous inside; upper lobes ovate, apex rounded to 

obtuse, c. 2.5 x 4.5 mm; lateral lobes ovate, apex 

rounded to obtuse, c. 3 x 3.7 mm; lower lobe ovate, 

apex rounded, c. 3.5 x 3.5 mm. Stamens inserted 

in lower half of corolla tube at 2 mm from base 

which is 20% of tube length; fi laments very pale 

orange, straight, c. 4 mm long, glabrous; anthers c. 

2.5 x 1 mm; staminodes equal in size, c. 0.9 mm 

long. Disk k yellow, to 0.7 mm high, margin thick, 

strongly crenate. Pistil l 

c. 11.4 mm long, held in

SOMRANIA, A NEW GENUS OF GESNERIACEAE FROM THAILAND (D.J. MIDDLETON & P. TRIBOUN) 11 

upper part of corolla tube; ovary green, 3.7 mm long, 

densely covered in a mixture of short glandular 

hairs (0.1 mm long) and slightly longer (0.2 mm) 

eglandular hairs; style white, c. 7.2 mm long, 

slightly curved, pubescence as on ovary but less 

dense; stigma with only the lower lobe developing, 

weakly and subequally laterally bilobed, lobes 

rounded. Fruit t fusiform, 7–11 x 1.2–1.8 mm, densely 

and minutely pubescent. Seeds c. 0.5 x 0.2 mm. 

Distribution.— Only known from Ranong 

Province in Thailand although may also be found 

in the far south of Burma. 

Ecology.— On limestone rocks or cliffs in 

dry evergreen forest or overhanging mangrove at 

20–580 m altitude. 

Etymology.— The specifi c epithet refers to 

the pure white corollas. 

Proposed IUCN conservation assessment.— 

Data Defi cient (DD). Although this species is currently 

only known from two limestone sites in 

Ranong province both of these sites are very close 

to the Burmese border and the possible distribution 

of the species in Burma is currently unknown. 

Additional specimen studied (paratype). 

Thailand.— PENINSULAR: Ranong [Kra Buri, 

Ban Lam Liang, Tham Phra Khayang, 20 m alt., 24 

June 2006, Williams, Pooma & Poopath 2123 (A!, 

BKF!, E!)]. 

Somrania lineata a D.J.Middleton & Triboun, sp. nov. 

Differs from Somrania albifl ora in the 

branched hairs being smaller and less frequently 

branched, the two orange-brown lines in the 

corolla, and the strongly oblique corolla mouth. – 

Type: Thailand, Phangnga, Sra Nang Manohra 

Waterfall Forest Park, Trail to limestone hill, on 

moist limestone base, 100 m alt., 17 June 2010, 

Triboun 4440 (holotype BK!; isotype E!). Fig. 2B. 

Lithophytic herb, rosulate or with a stem to 

1.5 cm long supporting opposite leaves and two 

infl orescences, these leaves reduced and strongly 

anisophyllous. Leaves primarily arising at base of 

plant; petiole 3–10.2 cm long, densely covered in a 

mixture of short acicular hairs and small branched 

hairs with 2–4 branches; blade ovate, 2.5–12.5 x 

2.3–10 cm, 1.1–1.5 times as long as wide, apex 

rounded, rarely almost obtuse, base rounded 

to cordate, sometimes somewhat unequally so, 

margin somewhat irregularly crenate, with 4–8 

secondary veins on each side of midrib, densely 

covered with short acicular hairs and even shorter 

glandular hairs above, densely covered in short 

acicular and sparsely-branched hairs and sessile 

glands beneath. Infl orescences either scapose or on 

short stems, 6–12 cm long, 3–8-fl owered; bracts 

linear, c. 2 mm long, covered with short acicular and 

short glandular hairs; peduncle 4–8.2 cm long, 

densely covered in a mixture of short acicular and 

even shorter glandular hairs; pedicels 3–8 mm long, 

densely covered in a mixture of short acicular and 

even shorter glandular hairs with occasional globular 

subsessile gland. Calyx tube 1–1.5 mm, lobes 

1.8–2.5 x 0.8 mm (to 5.5 mm long in fruit), narrowly 

elliptic to triangular, apex acute with a blunt 

tip, indumentum as on pedicels. Corolla white 

throughout except for two orange-brown lines in 

line with sinuses between lower three lobes, these 

lines to about half way down inside of tube, mouth 

strongly oblique with lower lip longer than upper 

lip, c. 18 mm long; tube 10–12 mm long, c. 2 mm 

wide at base and c. 4 mm at apex, densely covered in 

a mixture of short acicular and shorter glandular 

hairs outside, glabrous inside; upper lobes squarish, 

apex rounded, c. 2.2 x 2.5 mm; lateral lobes 

orbicular to obovate, apex rounded, 2.2–3 x 2.5–3 

mm; lower lobe orbicular to obovate, apex rounded, 

3.2–3.5 x 3.2–3.5 mm. Stamens inserted at 4 

mm from corolla base which is 36% of tube length; 

fi laments straight to slightly curved near the anther, 

white, 3.5–4 mm long, slightly widening just above 

base, glabrous; backs of anthers dark purple to 

nearly black, connective white, each 1.5–1.8 x 0.5 

mm; 2 lateral staminodes c. 2.5 mm long, medial 1 

mm long. Disk k c. 0.9 mm high, weakly crenate. 

Pistil l c. 11.5 mm long, held in upper part of corolla 

tube; ovary c. 3.5 mm long, densely covered in 

short acicular hairs; style c. 7.8 mm long, densely 

covered in a mixture of short acicular hairs and 

even shorter glandular hairs; stigma with only the 

lower lobe developing, weakly and subequally lat- 

erally bilobed, lobes rounded. Fruit t fusiform, 7.5– 

8.5 x 1.3–1.5 mm, with tiny glands. Seeds c. 0.4 x 

0.2 mm. 

Distribution.— Thailand (Phangnga). 

Ecology.— On limestone rocks or cliffs in 

shade in evergreen forest at 75–100 m altitude.

12 


Figure 1. Somrania albifl ora D.J.Middleton. A. Habit; B. Stellate hairs on petiole; C. Corolla dissection; D. Stamen; E. Calyx, disk 

and gynoecium; F. Dehisced fruit and persistent style. Scale bars: A, B = 1 cm; C, E, F = 5 mm; D = 2 mm. Drawn from living 

collection which was subsequently vouchered as Middleton 4216 6 

(E).

SOMRANIA, A NEW GENUS OF GESNERIACEAE FROM THAILAND (D.J. MIDDLETON & P. TRIBOUN) 13 

Etymology.— The specifi c epithet refers to 

the two orange-brown lines in the corolla tube. 

Proposed IUCN conservation assessment.— 

Critically Endangered CR B1ab(iii). This species is 

currently only known from Tham Pha Phueng and 

Sra Nang Manohra Waterfall Forest Park, only 

about 4 km apart and with a known Extent of 

Occurrence of considerably less than 100 km 2 . 

Both are surrounded by disturbed vegetation and 

both have high levels of disturbance from tourists 

leading to a decline in the quality of the habitat. 

Additional specimen studied (paratype). 

Thailand.— PENINSULAR: Phangnga [Muang 

Phangnga, Tham Pha Phueng, 75 m alt., 15 Sept. 

2010, Middleton, Bunpha, Karaket, Lindsay, 

Phutthai, Suddee & Tetsana 5434 (BKF!, E!, K!, 

KEP!, PSU!)]. 

A B 

Figure 2. A. Somrania albifl ora D.J.Middleton (photograph D.J.Middleton); B. Somrania lineata D.J.Middleton & Triboun (photograph 

T. Phutthai). 

ACKNOWLEDGEMENTS 

We thank K. Bunpha, C. Hemrat, P. Karaket, 

S. Lindsay, T. Phutthai, S. Suddee, S. Suwanachat 

& N. Tetsana for their assistance and company in 

the fi eld; T. Phutthai for the photograph of 

Somrania lineata; and I. Güner for the illustration 

of Somrania albifl ora. 

REFERENCES 

Kiew, R. & Lim, C.L. (2011). Names and new 

combinations for Peninsular Malaysian species 

of Codonoboea Ridl. (Gesneriaceae). Garden’s 

Bulletin Singapore 62: 253–275. 

Möller, M., Pfosser, M., Jang, C.G., Mayer, V., 

Clark, A., Hollingsworth, M.L., Barfuss, 

M.H.J., Wang, Y.Z., Kiehn, M. & Weber, A. 

(2009). A preliminary phylogeny of the ‘didymocarpoid 

Gesneriaceae’ based on three molecular 

data sets: Incongruence with available 

tribal classifi cations. American Journal of 

Botany 96: 989–1010. 

Weber, A., Middleton, D.J., Forrest, A., Kiew, R., 

Lim, C.L., Rafi dah, A.R., Sontag, S., Triboun, 

P., Wei, Y.-G., Yao, T.L. & Möller, M. (2011). 

Molecular systematics and remodelling of 

Chirita and associated genera (Gesneriaceae). 

Taxon 60: 767–790. 

Xu, Z.R., Burtt, B.L., Skog, L.E. & Middleton, D.J. 

(2008). A revision of Paraboea (Gesneriaceae). 

Edinburgh Journal of Botany 65: 161–347.


Actinostachys wagneri (Schizaeaceae), a new record for Thailand 

STUART LINDSAY*, THAMARAT PHUTTHAI**, KITICHATE SRIDITH***, 

SAHUT CHANTANAORRAPINT*** & DAVID J. MIDDLETON* 

ABSTRACT. Actinostachys wagneri (Selling) C.F.Reed, a new record for Thailand, is described and illustrated. 

KEY WORDS: Actinostachys, Schizaeaceae, fern, Thailand. 

INTRODUCTION 

Actinostachys Wall. ex Hook. is a small pantropical 

genus of ferns with about 20 species. For 

most of its history the genus has been included 

within Schizaea Sm., but in recent literature the 

genera have been treated as distinct (Smith et al., 

2006, 2008; Jaman & Umi Kalsom, 2010; 

Christenhusz et al., 2011) and as the only two genera 

in Schizaeaceae (the rest having been moved to 

Lygodiaceae - see Smith et al., 2006, 2008; 

Christenhusz et al., 2011). 

The only species of Actinostachys previously 

recorded from Thailand is Actinostachys digitata 

(L.) Wall. ex C.F.Reed, which is widely distributed 

in Southeast and Peninsular Thailand (Tagawa & 

Iwatsuki, 1979; Lindsay & Middleton, 2012 onwards). 

Recently a specimen was collected in 

Phangnga that has been identifi ed as Actinostachys 

wagneri (Selling) C.F.Reed, a new record for 

Thailand. This species is described and illustrated 

here. It differs from Actinostachys digitata in having 

shorter fronds (up to 20 cm long vs. 20−35 cm 

long), narrower fronds (up to 0.7 mm wide vs. 

more than 1 mm wide), stomata widely spaced in a 

single row on each side of the midrib and close to 

it (vs. almost end-to-end in a single row on each 

side of the midrib and at some distance from it), 

sporangia mostly in 2 rows (vs. sporangia mostly 

in 4 rows), and long soral paraphyses between sporangia 

(vs. no soral paraphyses). 

Actinostachys wagneri (Selling) C.F.Reed, Bol. 

Soc. Brot. 21: 131. 1947; Jaman & Umi Kalsom, 

Fl. Pen. Malaysia, Ser. I, Ferns & Lycoph. 1: 111, 

map 1. 2010.— Schizaea wagneri Selling, Svensk 

Bot. Tidskr. 40: 278, f. 8–11. 1946; Holttum, Rev. 

Fl. Malaya ed. 1, 2: 52. 1955 [‘1954’]; Holttum, Fl. 

Males., Ser. II, Pterid. 1: 44, f. 4e−g. 1959; Holttum, 

Rev. Fl. Malaya ed. 2, 2: 52. 1968. Type: Papua 

New Guinea, Admiralty Islands, Manus Island, 

Summit of Mt Tjajiak, Grether & Wagner 4177, 

November 1945 (holotype: UC! [barcode 

UC701236]; isotypes; K! [barcode K000408661], 

S).— S. paucijuga Holttum, Gard. Bull. Singapore 

11: 267. 1947. Type: Malaysia, Kedah Peak, Haniff 

SFN 1098 (holotype: SING, n.v.). Fig. 1. 

Terrestrial or sometimes appearing to be 

epiphytic at base of trees; rhizome short-creeping, 

producing fronds close together, less than 1 mm 

diameter, with a few thick, stiff, wiry roots c. 0.2 mm 

diameter; root hairs to 3 mm long; apex of rhizome 

densely covered in coarse brown multiseptate hairs 

up to 1.5 mm long. Fronds monomorphic, erect, 

simple, linear, (3–)7.5–16.5(–20) cm long, less 

* Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, Scotland, U.K. Email: s.lindsay@rbge.ac.uk 

** Division of Biological and Natural Resources Sciences, Mahidol University (Kanchanaburi campus), Sai-Yok, 71150 

Kanchanaburi, Thailand. 

*** Herbarium, Biology Department, Prince of Songkla University, Hat Yai 90112, Songkhla, Thailand.

ACTINOSTACHYS WAGNERI (SCHIZAEACEAE), A NEW RECORD FOR THAILAND 

(S. LINDSAY, T. PHUTTHAI, K. SRIDITH, S. CHANTHANAORRAPINT & D.J. MIDDLETON) 

A B 

C D 

Figure 1. Actinostachys wagneri (Selling) C.F.Reed: A. Habit; B. Rhizome and stipe bases; C. Young fertile lobes; D. Fertile lobes 

showing sporangia in two rows. 

15

16 

than 0.7 mm wide, reddish brown and terete or 

triquete at base, gradually becoming green and 

very narrowly winged above the base (therefore, 

no distinct stipe); basal portion sometimes sinuous 

and/or horizontal (especially if growing through 

leaf litter or moss); midrib rather broad and slightly 

to prominently raised on lower surface of winged 

portion but indistinct above, other venation, if any, 

obscure; with 2-celled dark glandular hairs sparse 

throughout; stomata on the lower surface only, 

widely spaced, in 1 row on each side of the midrib 

and close to it. Sporangia produced only at the 

frond apex in a digitate cluster of (2–)3–4(–5) 

fertile lobes; fertile lobes linear, slightly twisted 

together when young but untwisting as they mature 

and separate, (3.5–)7–15(–19) mm long, less than 

0.7 mm wide, with entire margins, venation of each 

lobe a single broad medial vein slightly raised 

below but indistinct above, with 2-celled dark 

glandular hairs above. Sporangia ovoid, sessile, 

each with a large apical annulus, usually arranged 

in 2 rows (very rarely with short sections of 3 rows) 

the entire length of the lobes; long fi liform transparent 

reddish-brown paraphyses protruding between 

sporangia, 2–4 times longer than sporangia, fl at, 

twisted, ribbon-like with rounded ends and all originating 

from the vein. Spores monolete, bilateral, 

fi nely verrucose. 

Thailand.— PENINSULAR: Phangnga [Thai 

Muang, Khao Lampi-Hat Thai Muang National 

Park, 17 Feb. 2011, Phutthai, Sridith & 

Chantanaorrapint 247 7 ( PSU)]. 

Distribution.— Peninsular Malaysia, 

Singapore, Borneo, Moluccas, New Guinea, NE 

Australia. 

Ecology.— In Thailand, Actinostachys wagneri 

is only known from beach forest at sea level 

but elsewhere it has been reported from a range of 

forest types and altitudes (eg. to 440 m in Peninsular 

Malaysia and to 1600 m in New Guinea). Plants are 

usually terrestrial in leaf litter or humus, in moss or 

on rotten logs but they can sometimes appear to be 

epiphytic in moss at the base of trees. Actinostachys 


wagneri is reported to grow with Actinostachys 

digitata in New Guinea and with both Actinostachys 

digitata and Schizaea dichotoma in Peninsular 

Malaysia and Singapore. 

IUCN Conservation Status. — Least Concern 

(LC). Although this species has been collected only 

once in Thailand and is known from relatively few 

localities in Peninsular Malaysia (Kedah, Perak 

and Johor) it is very widespread and not under any 

known threat. 

Note.— Following Holttum (1955) we place 

Schizaea paucijuga Holttum in synonymy of 

Actinostachys wagneri although we have not seen 

the type material. 

REFERENCES 

Christenhusz, M.J.M., Zhang, X-C. & Schneider, 

H. (2011). A linear sequence of extant families 

and genera of lycophytes and ferns. Phytotaxa 

19: 7–54. 

Holttum (1955 [‘1954’]). A Revised Flora of 

Malaya, edition 1, vol. 2 Ferns of Malaya: 52. 

Government Printing Offi ce, Singapore. 

Lindsay, S. & Middleton, D.J. (2012 onwards). 

Ferns of Thailand, Laos and Cambodia. http:// 

rbg-web2.rbge.org.uk/thaiferns/ 

Jaman, R. & Umi Kalsom, Y. (2010). Schizaeaceae. 

In: Flora of Peninsular Malaysia Ser. 1, 1: 107– 

117. FRIM, Kepong. 

Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, 

P., Schneider, H. & Wolf, P.G. (2006). A classifi 

cation of extant ferns. Taxon 55: 705–731. 


P., Schneider, H. & Wolf, P.G. (2008). Fern 

Classifi cation. Pp. 417–467 In: Ranker, T.A. & 

Haufl er, C.H. (eds), Biology and Evolution of 

Ferns and Lycophytes, CUP, Cambridge. 

Tagawa, M. & Iwatsuki, K. (1979). In: Smitinand, 

T. & Larsen, K. (eds), Flora of Thailand, Vol. 3, 

part 1. Royal Forest Department, Bangkok.


Amomum tomrey Gagnep. (Zingiberaceae), a new record for Thailand 

INTRODUCTION 

Amomum Roxb. is a genus of 150–176 species 

of terrestrial herbs and a few epiphytes, occurring 

from Sri Lanka and India eastwards to New Guinea 

and Australia (Lamxay & Newman, 2012). The last 

complete revision of this genus throughout its 

range is over a hundred years old (Schumann, 

1904). The most recent regional revision is for 

Cambodia, Laos and Vietnam by Lamxay & 

Newman (2012). Thirty-fi ve species and two varieties 

were recognized, of which seven species were 

newly described. In Thailand, 14 Amomum species 

were listed by Larsen (1996) in the preliminary 

checklist of the family Zingiberaceae. Later, 

Sirirugsa (2001) estimated that there were about 

15−20 Amomum species in Thailand and Larsen & 

Larsen (2006), in Gingers of Thailand, listed 16 

species. In addition, Amomum micranthum Ridl. 

was recently reported from Khlong Khruea Wai 

Wildlife Sanctuaries and A. inthanonense Chaveer. 

& Tanee has been reduced to A. coriandriodorum 

S.Q.Tong & Y.M.Xia (Kaewsri et al., 2009). 

Kaewsri (2006), in an unpublished PhD thesis, reported 

31 species of Amomum for Thailand, of 

which 13 were previously recognised species and 

the rest were proposed as new species. During a 

visit to the Royal Botanic Garden Edinburgh to 

continue taxonomic work on Amomum it was discovered 

that a specimen collected in Nakhon 

Nayok province, central Thailand, matched the 

type specimen of Amomum tomrey Gagnep. from 

the Paris herbarium. This species is newly recorded 

for Thailand. Most characters of this plant match 

WITTAYA KAEWSRI* 

ABSTRACT. Amomum tomrey Gagnep. is newly recorded for Thailand. It is described and illustrated. 

KEY WORDS: Amomum, Zingiberaceae, Thailand. 

place the Thai material in Amomum tomrey var. 

tomrey except that its leaves are not hairy along the 

margin. 

DESCRIPTION 

Amomum tomrey Gagnep., Bull. Soc. Bot. France 

53: 145. 1906; Gagnep. Fl. Indo-Chine 6: 113. 

1908. Type: Cambodia, Kampong Spoe, Samrong 

Tong, 12 Apr. 1870, J.B.L. Pierre 5847 7 (lectotype 

P [barcode P00599247], designated by Lamxay & 

Newman (2012); isolectotype P [barcodes P00599248 

& P00599249]). Fig. 1A−F. 

Terrestrial herb, 1.8−2.3 m tall; rhizome c. 

0.5−1.5 cm diameter; stilt roots absent; Leafy shoot 

slender, growing in clumps, pale green to purple 

near base. Leaves 4−26; sheath green, striate, glabrous; 

ligule subleathery, glabrous, apex 2-lobed, 

round, purple, 5−8 mm long; petiole 8−10 mm long, 

glabrous; lamina oblong-lanceolate to lanceolate, 

19−45 by 6−10 cm, glabrous, base attenuate or 

rounded, apex acuminate. Infl orescence subglobose, 

c. 2.5−3.0 cm diam.; peduncle 6−13 cm long; 

peduncular bract oblong, c. 4.0 by 2.7 cm, sparsely 

pubescent, leathery, apex mucronate; bract oblong 

or obovate-oblong, c. 1.8 by 1.0 cm, outer surface 

pubescent at base, purplish green, apex obtuse, 

apiculate; bracteole tubular, c. 9 mm long, apex unequally 

bifi d, sparsely pubescent. Calyx c. 1.1 cm 

long excluding ovary, apex trifi d and shallowly 

split on one side, base white pubescent. Corolla 

creamy white, tube sparsely hairy, 1.4 cm long excluding 

ovary, dorsal lobe hooded, ovate, c. 1.2 by 

* Mahidol University Amnatcharoen Campus, 259, Moo 13, Chayangkul Rd., Muang District, Amnatcharoen Province, 37000, 

Thailand. Email: wittaya.kae@mahidol.ac.th

18 

0.8 cm, apex blunt hooded, lateral lobe narrower. 

Staminodes subulate, white, c. 1.0 mm long. 

Labellum orbiculate, hooded, c. 1.2 cm diam., base 

auriculate, apex bifi d and attenuate, margin revolute 

toward apex, white with pale yellow mid-band, 

with pinkish lateral stripes from base radiating to 

margin, base sparsely white pubescent. Stamen 

creamy white; fi lament 1−2 mm long, sparsely 

white pubescent; anther 8.0 by 2.5−3.0 mm, dehiscing 

lengthwise; anther crest 3-lobed, rounded, 

4−5 by 3 mm, glabrous. Ovary cylindric, c. 2 by 2 

mm, white pubescent; stigma cup-shaped, the aperture 

edge hairy, stylodes blunt, c. 3 mm long. Fruit 

globose to oblong, smooth with 3 longitudinal 

grooves, glabrescent, c. 2 cm diam., pinkish brown, 

fruit sessile, to 7 mm long, apex with persistent 

calyx; seed angular, 6−7 by 4−5 mm, brown, enclosed 

by white aril. 

Thailand.— CENTRAL: Nakhon Nayok 

[Muang District, Khao Phra, Hub-luek, the jungle 

route behind Ban Kluai Kluai Resort, 14°16’ 

58.1”N, 101°12’ 45.17”E, 7 June 2003, Kaewsri 29 


Table 1. The character differences between A. tomrey and A. koenigii 

(BK); ibid, in Ban Kluai Kluai Resort, 10 Feb. 

2011, Kaewsri 302 (BK)]. 

Distribution.— Laos, Cambodia, Vietnam. 

Ecology.— Dry evergreen forest, open disturbed 

places, limestone bedrock, c. 100 m. 

Flowering and fruiting during March−July. 

Conservation.— In Thailand Amomum tomrey 

has been only found in Khao Phra sub-district, 

Muang District, Nakhon Nayok Province. 

According to the FAO data (Banerjee, 1997), 

Thailand has had the most rapid deforestation in 

the last 40 years. Its forests in 1961 occupied about 

27 million hectares and by 1989 was down to 14.3 

million which is only about 28% of the country 

(Banerjee, 1997). The habitat for Amomum tomrey 

is continually being reduced but currently the 

species remains fairly widespread. IUCN red list 

category NT (IUCN, 2001). 

Note.— Amomum tomrey is relatively similar 

to A. koenigii J.F.Gmel. The main character differences 

are shown in Table 1. 

Character A. tomrey A. koenigii 

Ligule length 5−8 mm < 5 mm 

Labellum shape orbiculate rhomboid 

Flowering habit continues to fl ower during fruit 

maturation 

Fruit glabrescent glabrous 


I would like to thank the Royal Botanic 

Garden Edinburgh for supporting this work. 

Exploration work was supported by the TRF/ 

BIOTEC Special Program for Biodiversity 

Research and Training, grant BRT R153060. I am 

grateful to the staff of Ban Kluai Kluai Resort for 

helping facilitate my fi eldtrip, and to the herbaria 

which made the collections available for this work 

(BK, BKF and E). 

REFERENCES 

Banerjee, A.K. (1997). Decentralization and 

fl owers rarely found when fruit 

reaches maturity 

Devolution of Forest Management in Asia and 

the Pacifi c, FAO Working Paper No: APFSOS/ 

WP/21. Forestry Policy and Planning Division, 

Rome Regional Offi ce for Asia and the Pacifi c, 

Bangkok. 

IUCN. (2001). IUCN Red List Categories and 

Criteria : Version 3.1. IUCN Species Survival 

Commission. IUCN, Gland, Switzerland and 

Cambridge, UK. 

Kaewsri, K. (2006). Systematic studies of the genus 

Amomum Roxb. (Zingiberaceae) in 

Thailand. Ph.D. Thesis, Kasetsart University. 

Kaewsri, K., Paisooksantivatana, Y. & Veesommai,

AMOMUM TOMREY GAGNEP. (ZINGIBERACEAE), A NEW RECORD FOR THAILAND (W. KAEWSRI) 19 

U. (2009). A new record and a new synonym in 

Amomum Roxb. (Zingiberaceae) in Thailand. 

Thai Forest Bulletin (Botany) 37: 32−35. 

Lamxay, V. & Newman, M.F. (2012). A revision of 

Amomum (Zingiberaceae) in Cambodia, Laos 

and Vietnam. Edinburgh Journal of Botany 69 

(1): 99–206. 

Larsen, K. (1996). A preliminary checklist of the 

Zingiberaceae of Thailand. Thai Forest Bulletin 

(Botany) 24: 35−49. 

1 cm 

D 

Larsen, K. & Larsen, S.S. (2006). Gingers of 

Thailand. Queen Sirikit Botanic Garden. 

Sirirugsa, P. (2001). Zingiberaceae of Thailand, pp. 

63−77. In: V. Baimai and R. Kumhom. BRT 

Research Reports 2001. Biodiversity Research 

and Training Program. Jirawat Express Co., 

Ltd., Bangkok (in Thai). 

Schumann, K. (1904). Zingiberaceae. In: Engler, 

Das Pfl anzenreich 4:46. (Heft 20). Leipzig: W. 

Engelmann. 

A B 

1 cm 

Figure 1. A−F. Amomum tomerey Gagnep.: A. leafy shoot; B. leaves; C. ligule; D. fl ower E. infructescence; F. fruit (sections). 

Photographed by Wittaya Kaewsri (all from Kaewsri 302 except D from Kaewsri 29). 

1 cm 

C 

1 cm 

1 cm 

E 

F


Revision of Cyclocodon Griff. ex Hook.f. & Thomson (Campanulaceae) 

INTRODUCTION 

Cyclocodon is a genus of Campanulaceae 

close to Campanumoea Blume and Codonopsis 

Wall., and into both of which it has previously been 

synonymised. The genus was recently reinstated as 

distinct mainly based on characters of the seed coat 

or pollen (Morris & Lammers, 1997; Hong & Pan, 

1998; Lammers, 1998, 2007a, 2007b). It is distributed 

in SE Asia from Bhutan and NE India 

through S China to Japan and southeast through 

Indochina and Malesia to New Guinea. The three 

WILLEM J.J.O. DE WILDE* & BRIGITTA E.E. DUYFJES* 

ABSTRACT. This work concerns the recognition, enumeration, and discussion of 4 species of Cyclocodon Griff. ex Hook.f. & 

Thomson, a genus occurring in Southeast Asia and Malesia. A key to the species of Cyclocodon is given. Cyclocodon axillaris (Oliv.) 

W.J.de Wilde & Duyfjes, based on Campanumoea axillaris Oliv. is reinstated, as this taxon was previously included in synonymy of 

Cyclocodon lancifolius (Roxb.) Kurz. 

KEY WORDS: Campanulaceae, Cyclocodon taxonomy, Asia, Malesia. 

Table 1. Differences between Codonopsis, Cyclocodon, and Campanumoea. 

genera also differ in macro morphology, in growth 

habit, disposition of fl owers, and in fruit (capsular 

versus baccate), and for a large part also in the position 

of the sepals (calyx lobes) relative to the 

ovary (Table 1). Concerning this latter item, following 

the schematic fi gures of lengthwise fl ower 

sections as presented by Chipp (1908) and 

Moeliono (1960), the differences between the three 

genera as accepted by the present authors are depicted 

in fi gure 1. 

Codonopsis Cyclocodon Campanumoea 

root tuberous short rootstock tuberous 

growth habit twining erect twining 

phyllotaxis alternate (mostly) opposite opposite (mostly) 

position of fl owers terminal terminal axillary 

disposition of sepals apical or lateral on the either lateral on the at base of the ovary 

relative to the ovary ovary 

ovary, or at base of, or Fig. 1C 

Fig. 1A –A 1 2 

below the ovary 

Fig. 1B –B 1 3 

fi laments dilated at base (mostly) terete or dilated at base terete 

stigma-lobes and 

locules of ovary 

and fruit 

3 4 or 5 (or 6) 3 

fruits capsule berry berry 

* Netherlands Center for Biodiversity Naturalis (section NHN), Leiden University, P.O. Box 9514, 2300 RA Leiden, the Netherlands. 

Email: Duyfjes@nhn.leidenuniv.nl

REVISION OF CYCLOCODON GRIFF. EX HOOK.F. & THOMSON (CAMPANULACEAE) (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 21 

Figure 1. Insertion of the sepals relative to the ovary in Codonopsis, Cyclocodon, and Campanumoea. Note that the corolla is always 

at the apex of the ovary. Stamens are not indicated. A 1 –A 2 . Codonopsis; B 1 –B 3 . Cyclocodon; C. Campanumoea. 

For the treatment of Campanulaceae for the 

Flora of Thailand the genus has been studied again. 

After the reinstatement of Cyclocodon axillaris 

(Oliv.) W.J.de Wilde & Duyfjes, based on 

Campanumoea axillaris Oliv., we recognize four 

species in Cyclocodon. Two species occur in 

Thailand: Cyclocodon celebicus (Blume) D.Y.Hong 

and C. parvifl orus (Wall. ex A.DC.) Hook.f. & 

Thomson. Because Campanumoea axillaris was 

formerly mostly placed in synonymy of Cyclocodon 

lancifolius (Roxb.) Kurz, Codonopsis lancifolia 

(Roxb.) Moeliono or Campanumoea lancifolia 

(Roxb.) Merr., in previous taxonomic treatments 

the latter species had a much wider circumscription 

when compared to that in the present treatment. 

TAXONOMY 

CYCLOCODON 

Griff., Not. Pl. Asiat. 4: 277 (1854), nom. invalid. 

sub Art. 42.1; Griff. ex Hook.f. & Thomson, J. 

Proc. Linn. Soc., Bot. 2: 17. 1858; Miq., Fl. Ned. 

Ind. 2: 1082. 1859.— Campanumoea sect. 


Cyclocodon (Griff. ex Hook.f. & Thomson) 

C.B.Clarke in Hook.f., Fl. Brit. Ind. 3: 436 

(1881).— Lectotype (designated by Pfeiffer, 

1874): Codonopsis parvifl ora Wall. ex A.DC. = 

Cyclocodon parvifl orus (Wall. ex A.DC.) Hook.f. 

& Thomson. 

Perennial herbs, stems mostly branched, 

erect; roots fi brous from short root-stock. Leaves 

opposite. Infl orescences in loose terminal panicles 

(or solitary, terminal or axillary, not in Thailand). 

Flowers pedicellate; bracteoles present or absent; 

perianth regular, 4- or 5-merous (in Thailand), ovary 

subglobose, inferior to corolla, superior or partly 

superior to sepals, 4- or 5-locular; sepals persistent; 

corolla partially divided; fi laments not or only 

somewhat dilated at base; stigma 4- or 5-lobed. 

Fruit: a berry, 4- or 5-locular. Seeds numerous, 

somewhat fl attened, subglobose, very small. 

Note.— The lectotypifi cation of the genus 

Cyclocodon by Hong & Pan (1998) with Cyclocodon 

adnatus Griff. is rejected because this name is invalid 

and also not mentioned in Hooker & Thomson 

(1858). 

1. Flowers 4-merous. Sepals inserted at base of the ovary or below the ovary on the pedicel, also in these positions in the fruit 

4. C. parvifl orus 

1. Flowers 5- (or 6)-merous. Sepals inserted at base of the ovary or on the ovary, in fruit always inserted on the fruit 

2. Sepals pectinately long-lobed. Flowers solitary, long-pedicelled, (terminal and) axillary to normal foliage leaves. Filaments 

much widened at base 1. C. axillaris 

2. Sepals dentate-serrate, or sub-laciniate or entire. Flowers terminal, solitary or in panicles, pedicels short or long, Filaments not 

widened at base 

3. Flowers solitary on short pedicels, 2 cm long or less, terminal or axillary to normal foliage leaves, bracts small, not appearing 

as reduced leaves 3. C. lancifolius 

3. Flowers in loose terminal panicles with mostly longer pedicels, 1–6 cm long, bracts minute or larger, often appearing as 

much reduced (petiolate) leaves 2. C. celebicus

22 

1. Cyclocodon axillaris (Oliv.) W.J.de Wilde & 

Duyfjes, comb. nov.— Campanumoea axillaris 

Oliv., Hooker’s Icon. Pl., ser. 3, 8: pl. 1775. 1888.— 

Type: China, Sichuan, Mt Omei, Faber 253 (holotype 

K).— Campanumoea truncata auct. non 

(A.DC.) Endl.: Merr., Philipp. J. Sci., C 7(2): 104. 

1912.— Campanumoea celebica auct. non Blume: 

Danguy, Fl. Indo-Chine [P.H.Lecomte et al.] 3: 

693. 1930.— Codonopsis lancifolia (Roxb.) 

Moeliono subsp. lancifolia auct. non Roxb.: 

Moeliono, Fl. Mal. Ser. 1, Spermat., 6: 121. 1960.— 

Codonopsis celebica auct. non (Blume) Miq.: 

Thuan, Fl. Laos, Cambodge & Vietnam 9: 10. 

1969.— Cyclocodon lancifolius auct. non (Roxb.) 

Kurz: D.Y.Hong & Lammers, Fl. China 19: 527. 

2011 (electronic version). 

Flowers solitary, 5- or 6-merous, on axillary 

peduncles ca. 5 cm long; bracteoles minute, linear, 

ca. 5 mm long; sepals linear, with (1–)3–6 pinnately 

arranged side-lobes; fi laments much widened 

and adaxially hairy at base. Fruits green, globose, 

ca. 1 cm in diam. 

Distribution.— China (Sichuan), Taiwan, 

Japan, Laos, Vietnam, N Sumatra, Philippines (no 

material seen), Moluccas (Buru). 

Notes.— 1. In the Flora of China (Hong & 

Lammers, 2011) this species, under the name 

Cyclocodon lancifolius, is described as having purple-black 

fruits; in other Cyclocodon species the 

fruits are white (or pinkish). 

2. The name Campanumoea truncata used 

by Merrill (1912), as ascribed to Endlicher, Gen. 

Pl. 1: 515. (“1836”) 1838, does not exist. We have 

not seen Vanoverbergh 965 (N Luzon) on which 

Merrill’s record of C. truncata was based. 

2. Cyclocodon celebicus (Blume) D.Y.Hong, Acta 

Phytotax. Sin. 36, 2: 109. 1998; D.Y.Hong & 

Lammers, Fl. China 19: 527. 2011 (electronic version).— 

Campanumoea celebica Blume, Bijdr. Fl. 

Ned. Ind. 13: 727. 1826.— Campanula celebica 

(Blume) D.Dietr., Syn. Pl. 1: 758. 1839.— 

Codonopsis celebica (Blume) Miq., Fl. Ned. Ind. 

2: 566. 1857.— Codonopsis lancifolia (Roxb.) 

Moeliono subsp. celebica (Blume) Moeliono, Fl. 

Mal. Ser. 1, Spermat., 6: 121. 1960.— Cyclocodon 

lancifolius (Roxb.) Kurz subsp. celebicus (Blume) 

K.E.Morris & Lammers, Novon 9, 3: 387. 1999. 


Type: Sulawesi (Celebes), Reinwardt s.n. L0585035 

(L, lectotype, here designated), see note.— 

Canarina moluccana Roxb., Fl. Ind. (ed. 1832) 2: 

173. 1832. Type: Roxburgh, not found, specimen 

possibly seen by Roxburgh at Calcutta, originating 

from the Moluccas.— Cyclocodon adnata Griff., 

Not. Pl. Asiat. 4: 278. 1854, nom. inval.— 

Codonopsis albifl ora Griff., Not. Pl. Asiat. 4: 279. 

1854. Type: S Myanmar (Mergui ‘Mergue’), 

Griffi th 1013 (not seen).— Codonopsis leucocarpa 

Miq., Fl. Ned. Ind. 2: 565. 1857.— Cyclocodon 

leucocarpus (‘leucocarpum’) (Miq.) Miq., Fl. Ned. 

Ind. 2: 1082. 1859. Type: Sumatra (Singgalang 

Gorge) Teysmann HB 1108 8 (holotype L). Fig. 2B–E. 

Flowers commonly in few-fl owered sub-paniculate 

terminal infl orescences, 5–10 cm across, 

with pedicels 1–6 cm long; bracts often with the 

appearance of reduced leaves; bracteoles small, 

linear; sepals linear, entire or few-serrate or fewdentate; 

fi laments (sub)terete, glabrous. Fruits 

white, depressed globose, (0.5–)1 cm in diam. 

Distribution.— South China (no material 

seen), southern Myanmar, Thailand, throughout 

Malesia to New Guinea. 

Notes.— 1. In the collection Vidal et al. 6179 

(Thailand) exceptionally the fruit opens by 3 apical 

valves. 

2. In the original description Blume (1826) 

only mentions “Celebes”, and Miquel (1857), who 

obviously examined the material, mentions 

“Celebes (Reinwardt)”. This material is composed 

of two sheets in L, presumably duplicates. One 

sheet (L0585034) bears in Reinwardt’s handwriting 

“1520 Campanula caesia R. Habitat in sylvis 

ad fl umen Batudoelang ...... Celebes. Septr. 1821”. 

The other sheet (L0585035) bears in Blume’s 

handwriting: “Campanula caesia, Celebes, 

Campanumoea celebica”. We have chosen the latter 

(L0585035) as the lectotype. 

3. The collection Teysmann 5768 (L) from 

Menado (Celebes) deviates in alternating leaves 

which are glabrous on the lower surface. 

4. Some collections from Sumatra ( (Korthals 

s.n., De Voogd 322 and d 1491) 

approach Cyclocodon 

parvifl orus in that the sepals are inserted at or close 

to the base of the ovary, also in fruit. The fl owers, 

however, are 5-merous. In some fl owers the opposite 

bracteoles (reduced leaves) are inserted on the


peduncle (pedicel) close to the fl ower, resembling 

the sepals as in C. parvifl orus; however, these 

fl owers (and fruits) additionally also have true sepals 

inserted on the ovary (fruit) close to the base. 

3. Cyclocodon lancifolius (Roxb.) Kurz., Flora 

55: 303. 1872; J. Asiat. Soc. Bengal 46, 2: 210. 

1877.— Campanula lancifolia Roxb., Fl. Ind. 

(Carey & Wallich ed.) 2: 96. 1824; Fl. Ind. (ed. 

1832) 1: 505. 1832.— Campanumoea lancifolia 

(Roxb.) Merr., Enum. Phillip. Fl. Pl. 3: 587. 

1923.— Codonopsis lancifolia (Roxb.) Moeliono, 

Fl. Mal. Ser. 1, Spermat., 6: 120. 1960.— 

Cyclocodon lancifolius (Roxb.) Kurz subsp. lancifolius: 

K.E.Morris & Lammers, Novon 9, 3: 387. 

1999. Type: Chittagong, Roxburgh (holotype BM, 

BM000070748, photo seen, see note).— 

Codonopsis truncata A.DC., Monogr. Campan.: 

122. 1830; Prodr. 7: 423. 1839.— Campanula 

truncata (A.DC.) D.Dietr., Syn. Pl. 1: 757. 1839.— 

Cyclocodon truncatus (A.DC.) Hook.f. & 

Thomson, J. Proc. Linn. Soc., Bot. 2: 18. 1858.— 

Campanumoea truncata (A.DC.) Diels, Bot. Jahrb. 

Syst. 29: 606. 1901. Type: Irrawady basin, Wallich 

1301 (holotype G (photo seen); isotype K-W). 

Flowers solitary, terminal to the leafy branches; 

pedicel short, 0.5–1(–2) cm long, bracteoles narrowly 

lanceolate, ca. 5 mm long; sepals (sparsely) 

serrate-dentate; fi laments (sub)terete. Fruits whitish, 

subglobose, 5–10 mm in diam. 

Distribution.— Bangladesh (Chittagong, type) 

and NE India (Shillong). 

Notes.— There has been confusion about the 

typifi cation of Campanula lancifolia Roxb., as no 

specimen was indicated in the protologue. There 

would appear to be neither Roxburgh type material 

nor drawings at CAL, K, or BR (Sanjappa et al., 1994 

(“1991”); Sealy, 1956; Forman, 1997). However, 

BM holds a specimen annotated in Roxburgh’s 

handwriting “Campanula lancifolia” and “moist 

soil l ……” together with “ 

N 217 Massat Roxburgh 

Flow Oct 1810” in unidentifi ed handwriting but possibly 

also from Roxburgh. We are convinced that 

this specimen should be regarded as the holotype. 

A representative specimen of the species possessing 

good fruits is: India, Eastern Circle, 

Shillong, Subansiri, Palin, 15 Nov. 1964, Sastri 

BSI I ( EC) C 40628 (L). 

4. Cyclocodon parvifl orus (Wall. ex A.DC.) 

Hook.f. & Thomson, J. Proc. Linn. Soc., Bot. 2: 18. 

1858.— Codonopsis parvifl ora Wall. ex A.DC., 

Monogr. Campan.: 123. 1830; Prodr. 7: 423. 

1839.— Campanula punduana D.Dietr., Syn. Pl. 1: 

757. 1839 (non Campanula parvifl ora Lam.).— 

Campanumoea parvifl ora (Wall. ex A.DC.) 

C.B.Clarke in Hook.f., Fl. Brit. India 3: 436. 1881. 

Type: NE India, Pundua, Wallich 1300 (holotype 

G-DC, photo seen; isotype K-W, photo seen).— 

Cyclocodon distans Griff., Not. Pl. Asiat. 4: 277. 

1854, nom. inval. Fig. 2A. 

Flowers 4-merous, in poor-fl owered terminal 

panicles; bracts (bracteoles) minute; sepals 4, linear, 

few dentate-lacerate, inserted on the pedicel at 

base of ovary or up to 4 mm below ovary; fi laments 

terete, slightly widened at base, glabrous. Fruits 

whitish, globose-obovoid, 4-locular, ca. 6 mm in 

diam. 

Distribution.— NE India (Assam, Sikkim), 

Bangladesh, Bhutan, S China, Myanmar, Thailand, 

Laos. 


We thank the curators of BK, BKF, K, L and 

QBG for permitting us to study their collections. 

We thank David Middleton (E) for improving the 

manuscript, Pramote Triboun (BK) for the photo of 

the rare Cyclocodon parvifl orus, and the colleagues 

from Khon Kaen University and Queen Sirikit 

Botanic Garden, particularly Phongsak Phonsena, 

Piyakaset Suksathan, and Pitak Panyachan, with 

whom we have conducted fi eldtrips. Jan van Os 

(Leiden) made the drawing and Ben Kieft (Leiden) 

prepared the drawing for publication. 

REFERENCES 

Blume, C.L. (1826). Bijdragen tot de Flora van 

Nederlandsch Indië. Lands Drukkerij, Batavia. 

Chipp, T.F. (1908). A revision of the genus 

Codonopsis Wall. Journal Linnean Society, 

Botany 38: 374–391. 

Forman, L.L. (1997). Notes concerning the typifi - 

cation of names of William Roxburgh’s species 

of Phanerogames. Kew Bulletin 52, 3: 

513–534.

24 


A B 

C 

D E 

Figure. 2. A. Cyclocodon parvifl orus (A.DC.) Hook.f. & Thomson. B–E. Cyclocodon celebicus (Blume) D.Y.Hong. Photographed 

by Pramote Triboun (A), Brigitta Duyfjes (B–E).


Hong, D.Y. & Pan, K.Y. (1998). The restoration of 

the genus Cyclocodon (Campanulaceae) and 

its evidence from pollen and seed-coat. Acta 

Phytotaxonomica Sinica 36: 106–110. 

Hong, D.Y. & Lammers, T.G. (2011). Cyclocodon. 

Flora of China @ efl oras.org: 527–528. 

Hooker, J.D. & Thomson, T. (1858). Praecursores 

ad Floram Indicam. J. Proc. Linn. Soc., Bot. 2: 

1–29. 

Lammers, T.G. (1998). Campanulaceae. Flora of 

Taiwan, ed. 2, 4: 775–800. Taipei, Taiwan, 

ROC. 

Lammers, T.G. (2007a). World checklist and bibliography 

of Campanulaceae: 1–675. Royal 

Botanic Gardens Kew. 

Lammers, T.G. (2007b). Campanulaceae. In: K. 

Kubitzki (ed.) The families and genera of vascular 

plants 8: 26–56. Springer. 

Merrill, E.D. (1912). Sertulum Bontocense. 

Philipp. J. Sci., C, 7: 71–107. 

Miquel, F.A.W. (1857). Flora van Nederlandsch 

Indië, vol. 2, pt. 4. Van der Post, Amsterdam. 

Moeliono, B. (1960). Codonopsis. In: C.G.G.J. van 

Steenis (ed.) Flora Malesiana, Series 1, 

Spermatophyta vol. 61: 118–121. Wolters- 

Noordhoff, Groningen. 

Morris, K.E. & Lammers, T.G. (1997). 

Circumscription of Codonopsis and the allied 

genera Campanumoea and Leptocodon (Campanulaceae: 

Campanuloideae). I. Palynological 

data. Botanical Bulletin Academia Sinica 38: 

277–284. 

Pfeiffer, L. (1874). Nomenclator botanicus: 964. 

Fischer, Kassel. 

Sanjappa, M., Thothathri, K. & Das, A.R. (1994 

(“1991”)). Roxburgh’s Flora Indica drawings 

at Calcutta. Bulletin Botanical Survey India 

33, 4: 90, 193. 

Sealy, J.R. (1956). The Roxburgh Flora Indica 

drawings at Kew. Kew Bulletin [11], 2: 318.


The distribution and ecology of the purple form of Ficus montana in western Thailand 

YAOWANIT TARACHAI*, PORNWIWAN POTHASIN**, WATTANA TANMING** & STEPHEN G. COMPTON*** 

ABSTRACT. Ficus montana Burm. f. var. purpurascens (Blume) has previously only been recorded from Java, by Corner (1960). 

It is distinguished from the typical form of F. montana by the strong purple pigmentation on the underside of the leaves, which is 

retained throughout their development. During our study of Ficus montana phenology in Kanchanaburi Province, Western Thailand 

from May 2008 to April 2009 we recorded its presence for the fi rst time in Thailand and compared its relative frequency and withinsite 

distribution to that of the typical form of the plant. Plants grown from seed and cuttings, and comparisons of the pollinators of 

the two forms, provide an indication of the nature of this variation in leaf colour. Cuttings grown under similar conditions retained 

their leaf colour, all seeds from typical plants produced typical offspring, but purple-leaved plants produced a mixture. The pollinator 

of the two forms is the same species of fi g wasp, Kradibia (= Liporrhopalum) tentacularis (Grandi). These results suggest the variation 

is due to an inherited colour polymorphism. 

KEY WORDS: Agaonidae, fi g tree, leaf colour, polymorphism, Western Thailand. 

INTRODUCTION 

Ficus montana Burm.f. and its pollinator fi g 

wasp (Liporrhopalum ( tentacularis (Grandi), syn. 

Kradibia tentacularis (Grandi), Agaonidae) have 

been intensively studied under laboratory conditions, 

but this small fi g tree is relatively poorly 

known in the wild (Raja et al., 2008a,b; Tarachai et 

al., 2008; Zavodna et al., 2005). F. montana is a 

widely-distributed functionally dioecious species 

(Subgenus Sycidium, Section Sycidium) recorded 

from Myanmar, Thailand, Peninsular Malaysia, 

Java, Sumatra and Borneo (Berg & Corner 2005). 

Despite its wide distribution, F. montana is recorded 

infrequently and Berg (in Berg and Corner, 2005) 

commented that it was “remarkably… poorly 

represented in herbarium collections”. This lack of 

records refl ects its cryptic growth form: it is a small 

shrub, often less than 50 cm tall, with stems that 

are often thinly-spaced and emerge above-ground 

within other vegetation, and small inconspicuous 

fi gs (Fig. 1). Leaf shape is also variable: entire, 

lobed or toothed, and sometimes varies within the 

same plant. It also frequently lacks any fi gs to help 

distinguish it from young plants of other Ficus 

species. Population sizes are also typically small. 

In his review of the fi g trees of Thailand, 

Corner (1965) recorded F. montana from 

Chanthaburi, Nakhon Sawan and Ratchaburi provinces, 

mainly from riparian situations, but he did 

not mention the purple-leaved colour form of the 

plant. Even in the recently published; Ficus, in 

Flora of Thailand vol. 10 part 4, Berg (2011) did 

not describe the purple leaf form of this species. 

Originally described as a separate species, F. montana 

Burm.f. var. purpurascens (Blume) Corner 

(1960) has previously only been recorded from 

Java. It is distinguished from the typical form of F. 

montana by the strong purple pigmentation on the 

underside of the leaves, which is retained throughout 

their development (Fig. 2). In the fi eld, the upper 

side of the leaves also sometimes appear to be a 

darker shade of green. Within plants, leaf colouration 

is consistent, either purple or green below, but 

young leaves of the typical form are sometimes 

purple beneath, although this quickly disappears 

once they are fully open. 

Here we record for the fi rst time the presence 

of the purple form of F. montana in Thailand, record 

its relative frequency and compare its withinsite 

distribution to that of the typical form of the 

* Faculty of Architecture and Environmental Design, Maejo University, Chiang Mai, Thailand. Email: yaowanit@mju.ac.th 

** Faculty of Science, Chiang Mai University, Chiang Mai, Thailand. 

*** Faculty of Biological Sciences, University of Leeds, Leeds LS2 9JT, UK.

THE DISTRIBUTION AND ECOLOGY OF THE PURPLE FORM OF FICUS MONTANA IN WESTERN THAILAND 

(Y. TARACHAI, P. POTHASIN, W. TANMING & S.G. COMPTON) 

plant. Plants grown from seed and cuttings, and 

comparisons of the pollinators of the two forms, 

provide an indication of the nature of this variation 

in leaf colour. 

METHODS 

Ficus montana herbarium specimens (recently 

determined by C.C. Berg) in the Chiang Mai 

Herbarium (CMU) and in the Bangkok Forest 

Herbarium (BKF) were examined in January 2007 

and February 2009 respectively to look for examples 

of specimens with purple leaves. We also carried 

out surveys for F. montana plants growing in nine 

stream-side locations in Western Thailand (Fig. 3) 

in 2008 and 2009. Their location, leaf colour and 

leaf shape were recorded. Cuttings were taken and 

a small number of mature fi gs that were present 

were removed to determine plant sex, to compare 

pollinators and to grow progeny under controlled 

conditions. 

Pollinator females were reared from male 

fi gs obtained from a purple-leafed individual in 

Phu Toei, Suphan Buri in February 2009. They 

were compared with fi g wasps from green-leafed 

plants collected from Erawan National Park, 

Kanchanaburi and the CIFOR estate, Bogor Java, 

Indonesia. 

RESULTS 

No specimens of F. montana var. purpurascens 

were found in the herbarium collections. F. 

montana was present along most of rocky stream 

beds in western Thailand that were examined, 

though it was never abundant (Table 1: no plants 

were detected at Sai Yok Noi waterfall). The plants 

were all growing in moderately shaded situations, 

either on rocks or stream-side sediments, with the 

tree canopy immediately above them or close by. F. 

montana var. purpurascens was present at several 

of the sites, but at others all the plants had the typical, 

green leaves. Where present, purple-leaved plants 

were usually in a minority, with the exception of 

the stream below the Ta Pern Kee Noi waterfall, 

where most of the plants had purple-coloured 

leaves. Overall, 19% of the 89 plants were of var. 

purpurascens. 

Purple- and green-leaved plants were often 

found growing in close proximity, sometimes with 

stems inter-mixed, and there were no obvious ecological 

differences in the substrates where they 

were established, nor the amount of cover/shade 

from the trees above them. The spatial inter-mixing 

of the two colour forms can be seen in their distribution 

along the stream at Erawan (Table 2). 

Purple-leaved forms of both male and female 

plants were present, and leaf colour was also independent 

of leaf shape. 

When cuttings of the purple form obtained 

from Erawan were grown in the same soil as plants 

with typical colouration they maintained their original 

colouration, indicating that the colour variation 

is not environmental in origin. Only greenleaved 

progeny were reared from three fi gs on 

three green-leaved female plants. 31 seedlings 

were grown from one fi g on a single purple-leaved 

female parent (Fig. 4); nine were green-leaved, the 

rest purple-leaved. There were no intermediates. 

Some of the fi g wasp pollinators reared from 

typical and purple-leaved plants wereLiporrhopalum 

tentacularis (Grandi). This is the normal pollinator 

across Indonesia and Thailand (Wiebes, 1994). 

However, individuals of a second, yellow, 

Liporrhopalum species were also reared from the 

single var. purpurascens fi g collected at Phu Toei, 

together with individuals of L. tentacularis. Further 

collecting will be needed to establish the nature of 

the relationship between this second agaonid species 

and F. montana var. purpurascens. 

DISCUSSION 

In Western Thailand, F. montana is typically 

a species of riparian habitats, whereas further 

south, in Sumatra and Java, it occurs more generally 

in open forest undergrowth, degraded forest and 

open spaces including parks (Zavodna et al., 2005, 

SGC Pers. Obs.). Its more restricted distribution in 

Thailand presumably refl ects the more seasonal 

rainfall experienced in the western and northern 

parts of its range. 

F. montana var. purpurascens has apparently 

not been reported previously other than from Java, 

yet purple-leaved plants were present at most of the 

western Thailand sites we surveyed. The pollinator 

fi g wasp L. tentacularis was reared from male 

27

28 

plants with both green and purple leaves, supporting 

the conclusion of Corner (1960) that they represent 

colour forms of the same species, though the 

relationship with a second species of pollinator remains 

unexplained. Furthermore, the consistency 

in leaf colour of individuals grown from cuttings, 


and that green-leaved progeny were obtained from 

a purple-leaved parent, indicate that the difference 

is likely to refl ect a genetic dimorphism, rather 

than an environmentally-induced response. Leaf 

colour is not restricted to one or other plant sex 

(both male and female purple-leaved plants were 

Table 1. Ficus montana colour morph frequencies in western Thailand. Note that it is often diffi cult to 

distinguish between adjacent individuals because of the spreading growth form of the plants, and the frequencies 

are therefore only approximate. 

Province Location Year Typical Purple 

Kanchanaburi Pha Tad 2008 3 0 

Sai Yok Yai 2008 1 0 

Dai Chong Tong waterfall 2008 8 6 

Erawan National Park 2009 18 6 

Huai Mae Khamin 2009 31 0 

Chaloem Rattanakosin National Park 2009 2 0 

Kra Teng Jeng Waterfall 2010 >40 0 

Suphan Buri Ta Pern Kee Noi waterfall, Phu Toei 

National Park 

2009 1 5 

Phetchaburi Kaeng Krachan National Park 2009 8 0 

Total >112 17 

Table 2. The sequence of colour forms of Ficus montana along the western bank of the Erawan waterfalls 

stream, arranged from north-south. The distance between plants 1 and 24 was approximately 1,500 metres. 

Plant Leaf colour Plant Leaf colour 

1 Green 13 Green 

2 Green 14 Purple 




6 Purple 18 Green 






12 Purple 24 Green

THE DISTRIBUTION AND ECOLOGY OF THE PURPLE FORM OF FICUS MONTANA IN WESTERN THAILAND 

(Y. TARACHAI, P. POTHASIN, W. TANMING & S.G. COMPTON) 

Figure 1. Ficus montana Burm.f.: A. habit; B. entire leaves; C. lobed leaves and D. fi gs. 

A 

B C 

A D 

B C 

Figure 2. Ficus montana Burm.f. var. purpurascens (Blume) Corner: A. habit; B. and C. lobed/toothed leaves; D. entire leaves and fi gs. 

D 

29

30 

recorded) and appears to be independent of leaf 

shape (as with typical plants, some purple-leaved 

individuals had simple leaves, others had leaves 

that were lobed or toothed). In addition, our observations 

suggest that green leaves may display simple 

genetic dominance over purple leaves, but formal 

breeding experiments would be needed to 

confi rm this. 

Red or purple pigmentation on the underside 

of mature leaves is common amongst understory 

plants growing in shady conditions in tropical forests 

(Lee et al., 1979). In most plants, including 

Ficus species, this colouration results from the accumulation 

of anthocyanins (Lee and Collins, 

2001). Proposed benefi ts for having an anthocyanin 

layer have included increased heat retention and 

‘back-scattering’ of light to increase rates of photosynthesis. 

Both these mechanisms have recently 

been discounted, and it has been suggested that its 

key benefi t may be to provide protection for plants 

that are shaded for much of the day, but are also 

subject to occasional damaging irradiance from 

brief periods of exposure to bright sunlight (Hughes 

et al., 2008). The F. montana growing along stream 

and rivers in Thailand will be exposed to just such 

a mixture of light and shade conditions, but purpleleaved 

individuals are nonetheless in a minority at 

most sites, suggesting that the costs associated with 

anthocyanin accumulation in the leaves often outweigh 

its potential benefi ts. 


We thank of the head of Erawan National 

Park and the head of Khuean Srinakarindra 

National Park for their supporting, Mr. Prasop 

Thongkeaw, at Erawan National Park and Mr. 

Somboon Daoruang, at Khuean Srinakarindra 

National Park, for fi eld work help. 

REFERENCES 

Berg, C.C. & Corner, E.J.H. (2005). Moraceae - 

Ficus. Flora Malesiana Series I (Seed Plants) 

Volume 17/Part 2. National Herbarium of the 

Netherlands, Leiden. 


Berg, C.C., Pattharahirantricin, N. & Chantarasuwan, 

B. (2011). Moraceae. In: Flora of Thailand, 

Vol. 10, pp. 475−675. 

Corner E.J.H. (1960). Taxonomic Notes on Ficus 

Linn., Asia and Australasia. IV. Subgen. Ficus 

Sect. Sycidium Miq. The Gardens’ Bulletin 

Singapore 17: 442−485. 

Corner, E.J.H. (1965). Check-list of Ficus in Asia 

and Australasia with a key to identifi cation. 

The Gardens’ Bulletin Singapore 21: 1−186. 

Hughes, N.M., Vogelmann, T.C. & Smith, W.K. 

(2008). Optical effects of abaxial anthocyanin 

on absorption of red wavelengths by understorey 

species: revisiting the back-scatter hypothesis. 

Journal of Experimental Botany 59: 

3435−3442. 

Lee, D.W, Lowry, J.B. & Stone, B.C. (1979). 

Abaxial anthocyanin layer in leaves of tropical 

rain forest plants: enhancer of light capture in 

deep shade. Biotropica 11: 70−77. 

Raja, S., Suleman N. & Compton, S. G. (2008a). 

Why do fi g wasps pollinate female fi gs? 

Symbiosis 45: 25−28. 

Raja, S., Suleman, N., Compton, S. G. & Moore, J. 

C. (2008b). The mechanism of sex ratio adjustment 

in a pollinating fi g wasp. Proceedings of 

the Royal Society of London B 275: 

1603−1610. 

Tarachai Y., Compton, S. G. & Trisonthi, C. (2008). 

The benefi ts of pollination for a fi g wasp. 

Symbiosis 45: 29−32. 

Wiebes, J.T. (1994). The Indo-Australian Agaoninae 

(Pollinators of Figs). Amsterdam, North-Holland. 

Zavodna, M., Arens, P.,Van Dijk, P.J., Partomihardjo, 

T., Vosman, B. & van Damme, J.M.M. (2005). 

Pollinating fi g wasps: genetic consequences of 

island recolonization. Journal of Evolutionary 

Biology 18: 1234−1243.


Three new species of Piper r (Piperaceae) from Thailand 

CHALERMPOL SUWANPHAKDEE*, DAVID A. SIMPSON** & PRANOM CHANTARANOTHAI* 

ABSTRACT. Three new species of Piper r (Piperaceae) from Thailand, P. doiphukhaense, P. khaoyaiense and P. smitinandianum are 

described and illustrated. 

KEY WORDS: Piper, new species, Thailand. 

INTRODUCTION 

Piperr L. is the largest genus of the family 

Piperaceae, with c. 1,050 species (Mabberley,1008) 

distributed mainly in tropical and subtropical regions. 

In Thailand, 46 taxa (45 species and two 

varieties) are enumerated (Suwanphakdee, 2012). 

Morphologically the genus is rather uniform, characterized 

by simple, alternate leaves and jointed 

stems with enlarged nodes. Branches are brittle. 

The stem anatomy is unusual for dicotyledons 

because it retains scattered vascular bundles in 

mature tissues (Metcalfe & Chalk, 1957). In preparation 

for publication of the Piperaceae for the 

Flora of Thailand, it has become necessary to publish 

a number of novelties. 

The unidentifi ed specimens deposited in 

BKF and QBG possess morphological characters 

which are not matched with known species. The 

collections consist of immature infl orescence and 

mature fruits that are clearly distinguishable from 

other known taxa. Moreover, some materials were 

recollected in the fi eld for examination by the fi rst 

author. Additional specimens from AAU, BK, BKF, 

BO, C, G, G-DC, K, K-W, KEP, KKU, L, PSU, 

QBG & SING (Thiers, 2012) were also consulted 

and studied. 

NEW SPECIES 

1. Piper doiphukhaense Suwanph. & P. Chantar. 

sp. nov., differs from P. macropiper r Pennant in its 

glabrous, pinnately veined leaves and smaller 

fruits. Type: Thailand, Nan, Doi Phu Kha National 

Park, 24 Aug. 2001, P. Srisanga & C. Maknoi 2095 

(holotype QBG). Fig. 1. 

Woody climber, dioecious, glabrous; nodes 

swollen with climbing roots. Leaves with petioles 

1.0–1.5 cm long, glabrous; stipules hood-like, lanceolate, 

glabrous, caducous; lamina subcoriaceous 

or coriaceous, ovate or elliptic, asymmetric, 7–10 

by 2–4 cm, base acute or oblique, apex acuminate 

or aristate, margin undulate, upper surface dark 

green, lower surface pale green; venation pinnate 

with 2–3 veins per side. Infl orescence e a terminal or 

leaf opposed catkin, pendulous, cylindrical, greenishwhite; 

rachis hairy, with dense fl owers; fl oral bract 

peltate ca. 1 mm diam., with stalk ca. 1.5 mm long 

and hairy at base. Infructescence 5–12 by 0.3–0.5 

cm, pendulous; peduncles 2.0–2.5 cm long, gla- 

brous. Fruit t free, sessile, ellipsoid-subglobose, ca. 

1.5 by 1 mm, dense on rachis, base rounded, apex 

acute with persistent stigma and fl oral bract. 

Thailand.— NORTHERN: Nan [Doi Phu Kha 

National Park, 24 Aug. 2001, P. Srisanga & C. 

Maknoi 2095 (holotype QBG)]. 

Distribution.— Endemic to Thailand. 

Ecology.— In shaded areas by streams and 

waterfalls in dry evergreen forest. Flowering and 

fruiting; August. 

Etymology.— The specifi c epithet of this 

* Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen, 40002, 

Thailand. Email: PP_SCKU56@yahoo.com 

** Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK.

32 

species refers to the locality, Doi Phu Kha National 

Park, Nan Province where the plant was collected. 

Note.— A comparison of morphological 

2. Piper khaoyaiense Suwanph. & D.A. Simpson. 

sp. nov., in general appearances similar to P. attenuatum 

Buch-Ham. ex Miq., but differs most obviously 

in its rounded or peltate fl oral bract with 

ciliate margin, stamens in a dense cluster of 5–7, 

larger and oblong-globose fruits. Type: Thailand, 

Nakhon Ratchasima, Khao Yai National Park, 9 

April 2004, C. Suwanphakdee 153 (holotype BKF; 

isotypes BK, KKU, QBG). Figs. 2 & 4. 

Woody climbers, dioecious, glabrous or puberulous; 

nodes swollen with climbing roots. 

Leaves with petioles 1–1.5 cm long; stipules hoodlike, 

lanceolate, glabrous or puberulous, caducous; 

lamina chartaceous, young leaves cordate to broadly 

cordate, rarely aristate, mature leaves ovate, 

rarely cordate, symmetric or asymmetric, 8–11 by 

6–9 cm, base oblique, cuneate, rarely cordate, apex 

acuminate or aristulate, margin undulate, upper 

surface green, lower surface pale green or yellowish 

green, glabrous, puberulous or glabrescent; venation 

pinnate, with 2 (–3) vein per side, glabrous 

or glabrescent. Infl orescence a terminal or leaf-opposed 

catkin, pendulous, cylindrical, yellow or 


features indicates that this species appears most 

closely related to Piper macropiper r Pennant rather 

than to Piper pedicellatum C.DC. see below. 

Character P. doiphukhaense P. macropiper P. pedicellatum 

Leaves 

: texture semicoriaceous coriaceous chartaceous 

: indumentum glabrous glabrous pilose 

: vein pinnate palmate pinnate 

Petiole glabrous puberulous, pilose, puberulous, pilose, 

hirsute 

velutinous 

Peduncle 

Floral bract 

glabrous glabrous velutinous 

: shape peltate peltate rounded 

: margin glabrous glabrous ciliate 

: stalk 

Fruit 

hairs at base hairs at base sessile 

:shape ellipsoid-subglobose ellipsoid globose 

:size c. 1.5 by 1 mm 1–1.5 by 0.5–0.8 mm 1–1.5 mm diam. 

:character unfl eshy, nut-like unfl eshy, nut-like fl eshy, drupe 

yellowish-green; rachis hairy, with dense fl owers; 

fl oral bract rounded ± short stalk, ca. 1 mm diam., 

margin ciliate. Male infl orescence 4–7 by 0.1–0.2 

cm; peduncles 1–2 cm long. Male fl ower: stamens 

in a dense cluster of 5–7, uniformly distributed 

throughout infl orescence, fi lament 0.6–0.8 mm long, 

anther 2-valved, slightly exserted at anthesis, 0.5– 

0.7 mm long, laterally dehiscent. Female infl orescence 

4–6 by 0.1–0.2 cm; peduncles 1–2 cm long. 

Female fl ower: ovary ±globose, stigma star-shaped 

3–5-lobed, 2–3 mm diam., hairy. Infructescence 

8–16 by 1–2 cm, pendulous, cylindrical, green; pe- 

duncles 0.5–3 cm long. Fruit t free, sessile, ± globose, 

4–5 by 3–4 mm, dense or sparse on rachis, base 

rounded, apex mucronulate with persistent stigma 

and fl oral bract, ripening fruit yellow, dark orange 

or red. 

Thailand.— EASTERN: Nakhon Ratchasima 

[Khao Yai National Park, 16 July 1962, T. T 

Smitinand 7443 (BKF), 9 April 2003, C. 

Suwanphakdee 52 (BK, BKF, KKU), 9 April 2004, 

C. Suwanphakdee 153 (holotype BKF; isotypes 

BK, KKU, QBG]; SOUTH-EASTERN: Prachin Buri

THREE NEW SPECIES OF PIPER (PIPERACEAE) FROM THAILAND (C. SUWANPHAKDEE, D.A. SIMPSON & P. CHANTARANOTHAI) 33 

Figure 1. Piper doiphukhaense Suwanph. & P. Chantar.: A. Branch with infructescences; B. & C. Side and top views of fl oral bract; 

D. A portion of infructescence; E. Fruit.

34 

[Khao Yai National Park, 8 March 2009, C. 

Suwanphakdee 264 (BK, BKF, KKU)]. 


Ecology.— Shaded or slightly open area 

along streams or waterfalls in evergreen forest. 

Flowering and fruiting; March to July. 


Etymology.— The specifi c epithet is referred 

to Khao Yai National Park, eastern Thailand where 

the type specimen was collected. 

Note. — Morphological differences between 

P. khaoyaiense and the similar P. attenuatum are 

shown below. 

character P. khaoyaiense P. attenuatum 

Stem glabrous or puberulous glabrous 

Petiole 

Floral bract 

puberulous hispidulous 

: shape rounded or peltate oblong (fl owering), spathulate 

(fruiting) 

: margin ciliate glabrous 

Stamen 

Fruit 

in a dense cluster of 5–7 2–4 

: shape oblong-globose ovoid-globose 

: size 4–5 by 3–4 mm 2–3.5 by 2–3 mm 

3. Piper smitinandianum Suwanph. & P. Chantar. 

sp. nov., similar to P. sclerophloeum C.DC. but 

differs by its small shrub, coriaceous leaves, with 

6–8 pairs of pinnate veins and narrowly ovid fruit 

shape with persistent style and spine-like stigma. 

Type: Narathiwat, Waeng, 12 June 1970, T. T 

Smitinand 10907 7 (holotype BKF). Fig. 3. 

Small shrubs, 0.5–1 m high, dioecious, glabrous; 

nodes swollen without climbing root. Leaves 

with petioles 1–2 cm long; stipules hood-like, oblonglanceolate, 

glabrous, caducous; lamina coriaceous, 

elliptic or elliptic-ovate, asymmetric, 16–27 by 

7–16 cm, base cuneate or oblique, apex acute to 

acuminate rarely aristate, margin undulate; venation 

pinnate with 6–8 veins per side. Infl orescence 

a terminal or leaf-opposed catkin, erect, oblongsubglobose; 

rachis hairy, with dense fl owers; fl oral 

bract rounded, ca. 1 mm diam. Infructescence a terminal, 

erect, leaf-opposed, cylindrical, ca. 4 by 

1–1.5 cm; peduncles ca. 1 cm long. Fruit t free, sessile, 

narrowly ovoid, 0.6–0.8 by 0.2–0.3 cm diam., 

dense on rachis, with persistent style and stigma 

spine-like and fl oral bract. 

Thailand.— PENINSULAR: Narathiwat [Waeng, 

20 Sept. 1965, C. Phengklai & T. T Smitinand 1188 

(BKF), same locality, 12 June 1970, T. T Smitinand 

10907 7 (holotype BKF), same locality, 22 Nov. 

1971, C.S.S. 268 (BKF)]. 


Ecology.— Open area in evergreen forest. 

Flowering and fruiting; September to November. 

Etymology.— The new species is named in 

honour of Prof. Tem Smitinand (1920–1995), outstanding 

Thai botanist and forester, who also initiated 

the Flora of Thailand Project. 


We gratefully thank the directors, curators 

and staff of herbaria cited above for permission to 

study the specimens and references. We also would 

like to thank Teerawut Srisuk, La-Ongdao 

Leukhachon & Orathai Kerdkaew for the line 

drawings. This work was supported by the Applied 

Taxonomic Research Center, Khon Kaen


Figure 2. Piper khaoyaiense Suwanph. & D.A. Simpson: A. Branch with infl orescences; B. Infructescence; C–D. A portion of male 

and female infl orescences; E–F. Side and top views of fl oral bract; G. Ovary; H. Stamen.

36 


1 cm 

Figure 3. Piper smitinandianum Suwanph. & P. Chantar.: Branch with an infructescence.


University grant ATRC_R5303, Khon Kaen 

University, Thailand. 

REFERENCES 

Mabberley, D.J. (2008). Mabberley’s plant-book, 

A portable dictionary of plants, their classifi - 

cation and uses. 3rd edition, Cambridge 

University Press. 

Metcalfe, C.R. & Chalk, L. (1957). Anatomy of the 

Dicotyledons- leaves, stem and wood in relation 

to taxonomy with notes on economic uses. 

Claredon Press, Oxford. 

Suwanphakdee, C. (2012). Systematics and utilization 

of family Piperaceae in Thailand. Ph. D. 

Thesis, Khon Kaen University. 

Thiers, B. (2012). Index Herbariorum: A global directory 

of public herbaria and associated staff. 

New York Botanical Garden’s Virtual Herbarium. 

http://sweetgum.nybg.org/ih/available on: 9 Mar. 

2012. 

Figure 4. Piper khaoyaiense Suwanph. & D.A. Simpson: A. Male infl orescences; B. Female infl orescences; C. Infructescence; D. 

Fruits (Photos by C. Suwanphakdee).


INTRODUCTION 

The delimitation of Lobelia species 

(Campanulaceae, subfam. Lobelioideae in Asia has 

long been confusing due to sometimes great superfi 

cial resemblance of the species and unclear typifi - 

cation of the names. For instance: the treatment of 

Lobelia trigona Roxb. by Arnott (1841) for India is 

a mixture of two species, and the circumscription 

of L. alsinoides by Moeliono in Flora Malesiana 

(1960) and by Thuan in Flora of Indochina (1969) 

is much too wide. Especially the group of closely 

allied species of small stature around the older 

name L. alsinoides posed problems when writing 

up Lobelia for Flora of Thailand (De Wilde & 

Duyfjes, expected in 2012). 

Although uncertainties remain, the enumeration 

of species here presented covers the whole 

Asian-Malesian area. In that area 21 species of 

small Lobelias can be recognized. As lesser-sized 

Lobelias we consider those plants usually not 

growing over 20(–40) cm tall, and having stems 

not thicker than 3 mm in diam. Lammers (2011), 

when defi ning 18 sections for 415 species in the 

whole world-wide genus Lobelia uses the term 

‘gracile’ for plants with stems of 5 mm diam. or 

less. In New Guinea four of the species are 

The lesser-sized Lobelias of Asia and Malesia 

WILLEM J.J.O. DE WILDE & BRIGITTA E.E. DUYFJES* 

ABSTRACT. Recognition, enumeration, and discussion of 21 species of lesser-sized Lobelias in South and South East Asia and 

Malesia are presented. A key to the lesser Lobelias, primarily based on seed shape and growth habit, is given. In Indochina and in 

the Malesian area the name Lobelia alsinoides Lam. is replaced by L. dopatrioides Kurz var. cantonensis (Danguy) W.J.de Wilde & 

Duyfjes. In the remaining area of L. alsinoides, mainly India, that species is divided into two varieties, viz. var. alsinoides and var. 

trigona (Roxb.) W.J.de Wilde & Duyfjes. Lobelia dioica R.Br. is a new record for Malesia. Lobelia reinwardtiana (C.Presl) A.DC. 

and Lobelia thorelii E.Wimm. are reinstated. Lobelia zeylanica L. var. walkeri C.B.Clarke is raised to species rank, L. walkeri 

(C.B.Clarke) W.J.de Wilde and Duyfjes. Lobelia reinwardtiana (C.Presl) A.DC. var. megalantha W.J.de Wilde & Duyfjes (India), L. 

dopatrioides Kurz var. kradungensis W.J.de Wilde & Duyfjes (Thailand), L. reptans W.J.de Wilde & Duyfjes (New Ireland), and L. 

serratifolia WJ.de Wilde & Duyfjes (Bougainville Isl.) are described as new. 

KEY WORDS: Campanulaceae, Lobelioideae, Lobelia taxonomy, nomenclature, Asia, Malesia. 

endemics from high altitude, not of direct interest 

for Thailand, and these have not been further studied. 

Furthermore L. dioica, a lowland species from 

Australia, is a new record for New Guinea. These 

fi ve species are included in the key presented in 

this study, which was framed as a result of the 

study of species to be accepted or considered for 

Thailand. Of the remaining 16 species of lesser 

Lobelias in Asia and Malesia, 10 species, including 

the reinstated L. thorelii, are accepted to occur in 

Thailand. The Indochinese and Malesian populations 

formerly identifi ed as L. alsinoides are now 

assigned to L. dopatrioides and segregated at the 

rank of variety. Lobelia alsinoides proper is largely 

confi ned to India and divided into two varieties, 

var. alsinoides and var. trigona. Lobelia reinwardtiana, 

with a newly described variety, var. megalantha, 

is reinstated for a widespread species occurring 

in S India and Malesia, with a closely 

related species, L. walkeri in Sri Lanka. Another 

species, L. reptans, possibly related to L. reinwardtiana 

and endemic to New Ireland, and the species 

L. serratifolia, endemic to Bougainville Isl., are 

newly described. Lobelia dopatrioides var. kradungensis 

is newly described for Thailand. 

The fl ower (corolla) colour of L. nummularia 

Lam. is largely pinkish; that of L. chinensis Lour. 

* Netherlands Center for Biodiversity Naturalis (section NHN), Leiden University, P.O. Box 9514, 2300 RA Leiden, the Netherlands. 

Email: Duyfjes@nhn.leidenuniv.nl

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 39 

pale pinkish or pale bluish, and in highland New 

Guinea species the corolla is some shade of red or 

chocolate coloured. All other Asian lesser Lobelias 

have blue fl owers. 

SE Asian lesser Lobelias all have alternate 

leaves. In the fi eld they may be confounded with 

especially Lindernia of Scrophulariaceae, a genus 

with opposite leaves and superior ovary. 

All species, with the principal synonyms, are 

enumerated with literature references, description, 

distribution, and notes. For complete synonymy 

and references one is referred to Lammers (2007), 

and for infrageneric classifi cation to Lammers (2011). 

The name L. loochooensis Koidz. (1929) 

from Kume and Okinawa (Ryukyu Islands) is listed 

by Lammers (2007) as a standing species. It is 

similar to the Australian L. quadrangularis R.Br. 

Its status is not evaluated in the present treatment. 

In order to assist in the fi nding of a name of a 

collected specimen, an enumeration of the species 

per geographic region is presented at the end of this 

article. 

CHARACTERS 

For the recognition of lesser-sized Lobelia 

specimens the following characters are of major 

importance and they preferably need to be employed 

in the given sequence: 

1. Shape of the seed, either r ellipsoid or rtrigo- nous. This is not diffi cult to determine, because the 

plants easily and quickly produce fruits with seeds 

and hence seeds are generally present in a collection. 

A discussion of the characters of seed anatomy 

and seed shape in Lobelia in general is presented 

by Lammers (2011). 

2. Growth habit: plant either r strictly erect 

with only one main stem and with roots only at its 

very base (not always easy to decide when collections 

contain only apical branches of the plant or 

suberect branches thereof), or r plant with (few or) 

several ramifi cations, growing directly from near 

the place of the main roots, with the branches sagging, 

ascending, or decumbent, or creeping with adventitious 

roots at least in the (very) lowest part.— 

Fig. 1. 

Figure 1. Growth forms of Lesser Lobelias. A–D. erect forms; E–G. decumbent forms. A–A'. L. heyneana; B. L. griffi thii; C. L. 

terminalis; D. L. dopatrioides var. dopatrioides; E. L. dopatrioides var. cantonensis; F. L. dopatrioides var. kradungensis; G. L. 

zeylanica.

40 

3. Shape and size of leaves. Distinction 

should be made between the lower r leaves, i.e. those 

in the lower part of the plant and not bearing fl owers 

in their axils, and the upper r leaves bearing sin- 

gle fl owers in their axils (sometimes these latter 

leaves are small so that they may be called bracts). 

4. Whether the stem is (sub)terete, angular, or 

distinctly winged. Useful as an additional character, 

but not always easy to ascertain in dried material. 

Lobeliia. alsinoides, L. heyneana, and L. reinwardtiana 

have distinctly angular or winged stems. 

(Sub)terete stems may be somewhat fl eshy in the 

living state and become winged when dry. 

5. Whether only two anthers are bearded at apex, or 

all fi ve anthers are bearded at apex. 

6. Whether the two anterior fi laments are 

wider, or all fi ve fi laments are equal in width. This 

character needs further investigation. It was used 

by Thuan in his key to the species in Flora of Indo- 

China (1969), discriminating between his widely 

conceived L. alsinoides and few other smaller 

Lobelias as accepted by him. As for the present 

treatment, this character appeared not easily to be 

ascertained in (boiled) herbarium specimens and 

possibly it is not practical in living plants either. 

We have refrained from recording this character. 


KEY TO THE LESSER-SIZED LOBELIASS OF ASIA AND MALESIA 

SECTIONS 

Of the 18 sections recognized by Lammers 

(2011) in Lobelia, fi ve sections occur in SE Asia, 

of which three sections comprise the lesser 

Lobelias. The 21 species at present recognized for 

Asia are divided over the sections as follows, 

largely corroborated by the species as they appear 

in the presented key: 

1. Sect. Delostemon (E.Wimmer) J.Murata: L. alsinoides, 

L. chevalieri, L. chinensis, L. dopatrioides, 

L. griffi thii, L. microcarpa, L. nummularia, L. 

terminalis, L. thorelii, and L. zeylanica (with 

bibracteolate pedicels, bearded anthers with tufts 

of fi liform hairs, and striate seed coat). 

2. Sect. Hypsala (C.Presl) Lammers: L. archboldiana, 

L. brachyantha, L. conferta, L. dioica, L. 

donanensis,and L. serratifolia (with ebracteolate, 

or less often bibracteolate pedicels, a bristle rather 

than a tuft of hairs on the anthers, and reticulate 

(not striate) seed coat). 

3. Sect. Stenotium (C.Presl) Lammers: L. hainanensis, 

L. heyneana, L. reptans, L. reinwardtiana 

and L. walkeri (with ebracteolate or bibracteolate 

pedicels, tufts of fi liform hairs on the anthers, and 

striate seed coat. 

Note.— Four narrowly endemic, high altitude species of E New Guinea (2000 m or more), are keyed-out 

and have been marked with an asterisk. They are mentioned in the enumeration of species below, but not 

further described; one is referred to Moeliono (1960). A fi fth high altitude species in New Guinea is the 

wide spread L. nummularia Lam., a species also occurring at lower altitudes. 

1. Plant dioecious, fl owers unisexual 7. L. dioica 

1. Flowers hermaphroditic 

2. Plant stemless. Plants from high altitude in E New Guinea 

3. Leaves membranous, margin (sub)entire, not glandular 2. L. archboldiana* 

3. Leaves coriaceous, margin glandular 8. L. donanensis* 

2. Plant with creeping, ascending, or erect stems 

4. Corolla less than 5 mm long. Plants from high altitude in E New Guinea 

5. Leaves 1–3 mm petiolate 3. L. brachyantha* 

5. Leaves sessile, blade semi-amplexicaul 6. L. conferta* 

4. Corolla more than 5 mm long 

6. Corolla (sub)unilabiate, lobes all more or less similar and placed in one row, dorsal slit wide, ca 90º or more 

7. Fruit berry-like (capsular fruits in certain specimens of L. nummularia from Philippines, Sulawesi or New Guinea excepted). 

Leaves (sub)circular, coarsely dentate 14. L. nummularia 

7. Fruit a capsule. Leaves not circular, dentate or not 

8. Leaves elliptic, to 5 mm wide, margin (sub)entire 5. L. chinensis 

8. Leaves (broadly) ovate-elliptic, 10–20 mm wide, margin serrate-dentate [Bougainville Isl.] 17. L. serratifolia 

6. Corolla bilabiate, upper lip 2-lobed, lower lip 3-lobed, dorsal slit, at least towards base, narrow, ca. 10º wide or less 

9. Seed ellipsoid. Only 2 anthers bearded at apex. Stem winged or angular 

10. Plant (main stem) erect. Flowers small, corolla 3–5 mm long 12. L. heyneana


10. Plant with lower part of main stem or (basal) branches decumbent or creeping. Flowers larger, corolla 6 mm long or more 

11. Plant stout, with long, rooting runners (1–)2–2.5 mm in diam. Pedicel 40–60 mm long, 2 or 3 times longer than sustaining 

leaf or bract. [New Ireland.] 16. L. reptans 

11. Plant less stout, branches decumbent, 1–2 mm in diam. Pedicel 10–20(–35) mm long, longer or shorter than sustaining 

leaf or bract 

12. All leaves broad, ± circular or rhombic or ovate, at base shortly cuneate into petiole 2–5 mm long. Plant delicate, 

sub-annual; stems 1–1.5 mm in diam. [Sri Lanka.] 20. L. walkeri 

12. Only lower leaves subcircular; other leaves and bracts narrower, sessile, base rounded or at base tapering forming 

a 1–6(–10) mm long petiole. Plant stouter, perennial, stems (1–)1.5–2 mm in diam. 

13. Stem 2- or 3-winged. [S India, Sumatra, Java.] 15. L. reinwardtiana (with 2 varieties) 

13. Stem 4-angular [Hainan.] 11. L. hainanensis 

9. Seed trigonous. All anthers bearded at apex. Stem angular or terete 

14. Plant with one (or more) main stem(s), all erect or ± zig-zag, not decumbent or sagged in lower portion, not rooting from 

the lower nodes 

15. Plant, especially the ovary, usually wholly long-hairy. Pedicel long, 15–40 mm long 18. L. terminalis 

15. Plant (ovary) (almost) glabrous. Pedicel 10–20(–25) mm long 

16. Leaves all small and narrow, ca 8 mm long or less. Flowers small, corolla ca 5 mm long 10. L. griffi thii 

16. Leaves larger and broader, ca 10 mm long or more 

17. Lower leaves (narrowly) elliptic. Flowers larger, corolla (6–)8–10 mm long 9a. L. dopatrioides (var. dopatrioides) 

17. Lower leaves subcircular (upper leaves narrower). Flowers small, corolla 4–5 mm long 13. L. microcarpa 

14. Plant with one or more main stem(s), commonly at base ± sagged, decumbent or creeping, commonly rooting from (the 

lower) nodes 

18. Leaves all with distinct petiole (2–)4–10 mm long. Ovary (usually) hairy 21. L. zeylanica 

18. Only the lower leaves petiolate, or all leaves (sub)sessile. Ovary glabrous or hairy 

19. Lower and upper leaves all broad, subcircular or ovate 

20. Stem (when fresh) angular, winged when dry. Ovary glabrous 1. L. alsinoides (with 2 varieties) 

20. Stem (when fresh) subterete, neither angular nor winged when dry. Ovary usually hairy 

21. Corolla ca 10 mm long, 3-coloured: upper lobes pink-lilac, lower lobes dark blue, with two white markings 

near the throat 19. L. thorelii 

21. Corolla 4–7(–8) mm long, all lobes uniformly blue (white markings excepted) 18. L. terminalis 

19. Leaves narrower, the lower ones (broadly) elliptic, the upper ones elliptic or narrowly elliptic. Ovary glabrous 

22. Fruit (ovary) for more than halfway superior 4. L. chevalieri 

22. Fruit largely (wholly) inferior 9. L. dopatrioides (with 3 varieties) 

ENUMERATION OF SPECIES 

1. Lobelia alsinoides Lam., Encycl. [J. Lamarck et 

al.] 3 (2): 588. 1792; E.Wimm., Ann. Naturhist. 

Mus. Wien 41: 348. 1948; Pfl anzenr. [Engler] 

IV.267b (107. Heft) II. Teil: 571. 1953.— Type: 

Herb. Lamarck. Lobelia. a cap. b. spei. Sonnerat 

s.n. (P-LAM 00356087, image available from 

http://www.lamarck.cnrs.fr/herbier.php), see note. 

For synonyms see the varieties. 

Subperennial herb, glabrous; branches few or several, 

ascending; stems angular or winged. Leaves: 

petiole to 4 mm long or leaves subsessile; lower 

and upper blades similar, subcircular or broadly 

ovate. Bracteoles present. Flowers: ovary glabrous; 

corolla bilabiate, 5–7 mm long; anthers all 

bearded at apex. Fruit t a capsule, glabrous. Seeds 

trigonous. 

Distribution.— See under the varieties. 

Notes.— 1. Lobelia alsinoides was for a long 

time regarded as a species with a much wider distribution 

than here accepted. Lammers (1992) 

roughly regarded the north-eastern part of its then 

distributional area to represent a separate subspecies, 

subsp. hancei (H.Hara) Lammers, but we 

found that also the Indo-Chinese and Malesian material 

belongs to this taxon. We accept it on species 

level, under the older name L. dopatrioides, with 

three varieties, although some specimens from the 

area where L. alsinoides and L. dopatrioides overlap 

are sometimes not easy to name. 

2. We recognise two sympatric varieties in L. 

alsinoides, different from the varieties as accepted 

by Wimmer (1953). 

3. The type of L. alsinoides (Sonnerat s.n.; 

P00356087) was said to originate from the Cape 

(“Cap de Bonne-Espérance”) but it must have been 

collected rather in Asia, presumably in southern 

India or in Sri Lanka, because plants similar to L. 

alsinoides are not known from Africa.

42 


KEY TO THE VARIETIES 

1. Lower leaves (sub)circular, upper leaves subcircular or ovate-elliptic a. var. alsinoides 

1. All leaves somewhat narrower, ovate or broadly elliptic b. var. trigona 

a. var. alsinoides.— Lobelia stipularis Roth ex 

Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 

5: 67. 1819; Roth, Nov. Pl. Sp.: 144. 1821.— Type: 

S India, near Mangalore, Hohenacker 1617 7 (neotype 

L, here designated). 

Lower leaves (sub)circular, upper leaves subcircular 

or ovate-elliptic. 

Distribution.— Northern and southern India; 

Sri Lanka; Myanmar, NW Thailand, east (perhaps) 

to approximately the lower Salween River basin; 

one odd collection Veldkamp 7157 7 (BO, L, K), 

from Sumatra (Toba Lake: Samosir Isl.). Possibly 

also China, but no collections seen. 

Ecology.— In paddy fi elds and forest edges, 

300—1000 m altitude. 

Note.— Neotypifi cation of L. stipularis. The 

description of L. stipularis (Roth in ms) was based 

on Heyne material that was sent to Roth (in 

Germany) under the name L. zeylanica, but this 

original material is no longer extant. Clarke (1881) 

placed L. stipularis in the synonymy of L. trigona, 

arguing in a note under the latter species that 

Heyne’s material at K and BM (and named L. zeylanica 

by Heyne) belongs to L. trigona, but that 

Roth’s description (published in 1821) rather 

points to a similar though different species. Roth’s 

manuscript was published, somewhat shortened, 

by Schultes (1819), which was followed by Roth’s 

own elaborate description (1821), without mentioning 

Schultes. We agree with Clarke’s view, i.e. 

that Roth’s description also contains elements of a 

second species (in fact our present L. walkeri), and 

that the material at the time at the hand of Roth was 

a mixture. In order to stabilize the current concept 

of L. stipularis Roth ex Schult. as being a synonym 

of L. alsinoides var. alsinoides, we designate a neotype. 

Lobelia alsinoides var. alsinoides is a taxon 

with distinct bracteoles (‘stipules’) and (sub)circular 

lower leaves. In L. walkeri the bracteoles at the 

base of the pedicel are absent or very small, ca 0.5 

mm long only. 

b. var. trigona (Roxb.) W.J.de Wilde & Duyfjes, 

stat. nov.— Basionym: Lobelia trigona Roxb., Fl. 

Ind. (Carey & Wallich ed.) 2: 111. 1824; Wight, 

Icon. Pl. Ind. Orient. t. 1170. 1848.— Type: not 

indicated; the drawing in Roxburgh Icones, ined. 

(K, http://apps.kew.org/fl oraindica/home) gives a 

good picture, but the specimen in Herbarium 

Hookerianum (K), indicated by Hooker as from 

Roxburgh, is herewith designated as the lectotype. 

All leaves ovate or broadly elliptic. 

Distribution.— N & S India, Sri Lanka (?), 

north-eastern boundary in India not certain. 

2. Lobelia archboldiana (Merr. & L.M.Perry) 

Moeliono, Fl. Males., Ser. 1, Spermat. 6: 131. 

1960.— Pratia archboldiana Merr. & L.M.Perry, 

J. Arnold Arb. 30: 59. 1949.— Type: Papua New 

Guinea, Murray Pass, Brass 4943 (holo GH, not 

seen). 

This is a low-growing “stem-less” herb, endemic 

of eastern New Guinea, from high altitude, 

2840–3600 m. 

3. Lobelia brachyantha Merr. & L.M.Perry, J. 

Arnold Arbor. 22: 385. 1941.— Type: Indonesia: 

West Papua (NE of Lake Habbema), Brass 11570 

(holo GH, not seen; iso L). 

This is a creeping herb, only known from one 

collection, of high mountain habitat (2350 m altitude) 

in western New Guinea. 

4. Lobelia chevalieri Danguy, Bull. Mus. Natl. 

Hist. Nat., Ser. 2. 1: 263. 1929; Fl. Indo-Chine 

[P.H.Lecomte et al.] 3: 683, f. 76: 9. 1930.— Type: 

Vietnam, Annam, Chevalier 30607, (lecto 

P00243509, designated by Danguy, 1930; photo 

seen). 

As taken from the protologue (Danguy, 1929) 

the characters relevant for our present species description 

are as follows: 

Annual herb, glabrous, branches several, decumbent; 

stem angular. Leaves (sub)sessile; all 

blades elliptic to lanceolate. Bracteoles present.


Flowers: ovary glabrous, for the greater part superior; 

corolla bilabiate, 8–10 mm long; anthers all 

bearded at apex. Fruit t a capsule, glabrous, largely 

superior, sepals refl exed. Seeds trigonous. 

Distribution.— Vietnam (Annam, between 

Dabang and Dran). 

Ecology.— Altitude between 200 and 1200 

m, exact altitude not known. 

Notes.— 1. Pencil sketches by the author of 

the superior ovary (and other details) are on the 

lectotype sheet (P00243509). The superior ovary is 

also depicted in Danguy (1930, l.c.). 

2. Lammers (2007) placed L. hancei in the 

synonymy of L. chevalieri, maybe disregarding the 

fact that the latter species was described as having 

a largely superior ovary. We consider L. hancei as 

synonym of L. dopatrioides var. cantonensis. 

Lobelia chevalieri is known only from two collections. 

If the superior position of the ovary is due to 

an aberrant condition, which could be ascertained 

when more similar material becomes available 

from the type locality, then this name would go under 

L. dopatrioides var. cantonensis. 

5. Lobelia chinensis Lour., Fl. Cochinch. 2: 514. 

1790; Merr., Trans. Amer. Philos. Soc. 24: 382. 

1935.— Type: Hong Kong, Lamont 403A, barcode 

L0585148 (neotype L, here designated, see note 1). 

— L. radicans Thunb., Trans. Linn. Soc. London 2: 

330. 1794.— Pratia thunbergii G.Don, Gen. Hist. 

3: 700. 1834.— Isolobus radicans (Thunb.) A.DC., 

Prodr. (DC.) 7, 2: 353. 1839.— Type: not indicated, 

but no. 21068 in herb. Thunberg (UPS) agrees, 

(lectotype no. 21068 in herb. Thunb UPS, here 

designated).— Lobelia caespitosa Blume, Bijdr. 

Fl. Ned. Ind. 13: 729. 1826.— Type: Java, Mt 

Gede, Blume s.n., barcode L0820633 (holo L).— 

Pratia radicans G.Don, Gen. Hist. 3: 700. 1834.— 

Isolobus roxburghianus A.DC., Prodr. (DC.) 7, 2: 

353. 1839.— Lobelia roxburghiana (A.DC.) 

Heynh., Nom. Bot. Hort. 1: 471. 1840.— Type: 

Wallich 1308 (holo K, 2 sheets). 

Perennial herb, glabrous, caespitose, branches 

ascending, rooting at lower nodes; stem subterete; 

roots fi brous. Leaves distichous; petiole absent; 

lower and upper blades similar, (broadly) elliptic. 

Bracteoles absent. Flowers: ovary glabrous; corolla 

unilabiate, 10–15 mm long; only 2 anthers 

bearded at apex. Fruit t a capsule, glabrous. Seeds 

compressed, ellipsoid. 

Distribution.— China; widely naturalized 

a.o. in Sri Lanka, Thailand, and Indonesia (Java). 

Ecology.— Wet places; at low altitudes. 

Notes.— 1. Loureiro (1790) gave ‘Canton’ as 

provenance for L. chinensis, but did not indicate 

the type. As no material of Loureiro could be 

traced, a neotype is chosen here. 

2. The species is said to be often not fruitsetting 

outside its area of origin. 

6. Lobelia conferta Merr. & L.M.Perry, J. Arnold 

Arbor. 30: 59. 1949.— Type: Papua New Guinea, 

Mt Albert Edward, Brass 4417 7 (holo GH, not 

seen). 

This is a high-altitude (3680 m) prostrate 

herb from Papua New Guinea. 

7. Lobelia dioica R.Br., Prodr. Fl. Nov. Holland.: 

565. 1810.— Type: Australia, Gulf of Carpentaria, 

Brown s.n. (BM, not seen). 

This annual herb was so far known only from 

Australia. However, L. dioica also occurs in New 

Guinea. It is known there from two collections: one 

from West Papua (Van Royen 4906) 6 and one from 

Papua New Guinea ( (Henty & Foreman NGF 

49367). 7 Moeliono (1960) did not recognize these 

collections, the reason that L. dioica was omitted in 

Flora Malesiana. 

8. Lobelia donanensis P.Royen, Kew Bull. 20: 

305. 1966; Steenis, Fl. Males., Ser. 1, Spermat. 6, 

addenda: 928. 1972.— Type: Papua New Guinea, 

Milne Bay Distr., Mt Donana, Cruttwell 1250 (holo 

K; iso L). 

A middle altitude herb from eastern Papua 

New Guinea, once found at 2250 m altitude. 

9. Lobelia dopatrioides Kurz, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 39 (2): 77. 1870; Kerr, Fl. 

Siam. 2: 303. 1936. 

Annual or (sub)perennial herb, glabrous, 

with one or more main stems; stems erect or suberect 

and commonly sagged in lower portion, or

44 

decumbent, not rooting at the nodes; stem angular 

or subterete, not winged. Leaves: lower ones sessile 

or short-petiolate, and blades (broadly) elliptic; 

upper ones (sub)sessile, and blades elliptic or narrowly 

elliptic, apex acute. Bracteoles present or 



not obvious. Flowers: pedicel 10–20(–25)mm 

long; ovary glabrous; corolla bilabiate, 6–12 mm 

long; all anthers bearded at apex. Fruit t a capsule. 

Seeds trigonous. 

1. Plant erect a. var. dopatrioides 

1. Plant suberect or decumbent 

2. Flowers small, corolla 6–9 mm long b. var.cantonensis 

2. Flowers large, corolla 10–12 mm long c. var. kradungensis 

a. var. dopatrioides.— Lobelia griffi thii Hook.f. 

& Thomson var. dopatrioides (Kurz) Kurz, J. Asiat. 

Soc. Bengal, Pt. 2, Nat. Hist. 46 (2): 211. 1877.— 

Type: Myanmar, Arracan, Kurz s.n. (holo CAL, 

not seen; iso K). 

Erect annual herb, with one or few main 

branches; stem angular. Leaves sessile, lower 

blades (5–)10–20 mm long, upper blades (5–)10 

mm long. Flowers: corolla (6–)8–10 mm long. 

Distribution.— S Myanmar; Thailand; 

Indochina. 

Ecology.— In wet places, open marshy and 

grassy grounds, open areas on sandy soil; from sea 

level to 520 m altitude. Flowering and fruiting all 

year round. 

Notes.— Lobelia dopatrioides var. dopatrioides 

and L. griffi thii both have an erect growth 

habit but the latter differs in having much smaller 

leaves. Lobelia dopatrioides var. dopatrioides may 

be confused with both L. alsinoides and L. dopatrioides 

var. cantonensis. However, both are distinct 

in having basically sagged or ascending main 

branches, although the difference with var. cantonensis 

is sometimes gradual and may be infl uenced 

by habitat. 

The fl owers are sometimes recorded as white. 

b. var. cantonensis (Danguy) W.J.de Wilde & 

Duyfjes, comb. nov.— Basionym: Lobelia chinensis 

Lour. var. cantonensis Danguy forma cantonensis, 

Fl. Indo-Chine [P.H.Lecomte et al.] 3: 681. 

1930.— L. alsinoides Lam. var. cantonensis 

(Danguy) E.Wimm., Ann. Naturhist. Mus. Wien 

56: 360. 1948.— Type: China, Canton, Hance 634 

(lecto W, here designated, not seen; iso K).— L. 

chinensis Lour. var. cantonensis Danguy forma 

elongata Dangui, Fl. Indo-Chine [P.H.Lecomte et 

al.] 3: 681. 1930.— Syntypes: Tonkin, Quang-yen, 

Balansa; Cambodia, Kampot, Harmand, Lecomte 

& Finet, Geoffray; Annam Thua-thien Eberhardt 

(lectotype not yet designated.— L. hosseusii 

E.Wimm., Repert. Spec. Nov. Regni Veg. 26: 2. 

1929.— Type: Thailand, Doi Chiang Dao, Hosseus 

462 (holo W, photo seen; iso K).— L. hosseusii 

E.Wimm. var. villosa Kerr, Fl. Siam. 2: 304. 

1936.— Type: Thailand, Put 1950 (holo K).— L. 

hancei H.Hara, J. Jap. Bot. 17: 23. 1941.— L. alsinoides 

Lam. subsp. hancei (H.Hara) Lammers, 

Bot. Bull. Acad. Sin. (Taipei) 33(3): 287. 1992.— 

Type: China, Canton, Sampson in Hance 634 (holo 

GH, photo seen; iso K). 

Annual or subperennial herb with one or several 

branches, ascending; stem subterete or angular. 

Lower leaves: petiole up to 3 mm long and 

blades (broadly) elliptic; upper leaves sessile and 

blades narrowly elliptic or linear. Flowers: corolla 

6–9 mm long. 

Distribution.— NE India(?), east through 

China (Hong Kong, Guangzhou: Canton, Hainan, 

Kung Tung) and Indochina (including Thailand 

and Vietnam: Tonkin, Annam to Japan, and southeast 

to New Guinea. 

Ecology.— Common in wet areas like moist 

sandy and marshy soils, on open grassy ground, 

and edges of fl ooded rice fi elds; from sea level to 

200 m altitude. Flowering and fruiting all year 

round. 

Notes.— Most material of the present L. 

dopatrioides var. cantonensis was formerly included 

in a widely accepted L. alsinoides, but the distinctness 

from proper, now more narrowly conceived 

L. alsinoides, was blurred by the inclusion


of material of quite distinct species, e.g. L. reinwardtiana, 

L. terminalis, and L. zeylanica. In general, 

the corolla of Lobelia dopatrioides var. cantonensis 

is larger (6–9 mm long) than that of L. 

alsinoides (5–7 mm long). 

c. var. kradungensis W.J.de Wilde & Duyfjes, 

var. nov. A varietate cantonensis habitu validiore 

(sub)perennanti, fl oribus lazulineis, corolla 10–12 

mm longa differt.— Typus: Thailand, Loei, Phu 

Kradueng, 16 Jan. 1966, Hennipman 3703 (holo 

BKF, iso L). 

Perennial herb, often purplish tinged, stems 

ascending, one or several growing from primary 

root-tuft, not or but occasionally rooting from lower 

nodes; stem subterete, not winged. Leaves: blade 

10–20 by 5–10 mm, upper blades smallest. 

Bracteoles 0.5–1 mm long. Flowers: sepals 3–3.5 

mm long; corolla 10–12 mm long, corolla lobes 

uniformly bright dark blue, except for white markings 

in the throat, lower 3 lobes broadly obovate, 

each ca 3 mm wide. Fruit t subglobose, rounded at 

base, ca 3 by 3 mm. Seeds trigonous. 

Distribution.— Endemic to Phu Kradueng 

and its surroundings, Loei, NE Thailand. 

Ecology.— Common in open grassy places 

on slightly peaty sandy soil (on the plateau of Phu 

Kradueng,); 800–1300 m altitude. 

Notes.— Variety kradungensis obviously is 

close to the widely conceived and very widespread 

L. dopatrioides var. cantonensis, but on the plateau 

of Phu (= Mt) Kradung the latter is replaced by distinct 

plants of a stouter and perennial habit and 

with conspicuously large dark blue fl owers. 

10. Lobelia griffi thii Hook.f. & Thomson, J. Proc. 

Linn. Soc., Bot. 2: 28. 1858; Kerr, Fl. Siam. 2: 303. 

1936.— Type: Myanmar, Mergui, Griffi th 660 

(holo K, two sheets). 

Annual herb, glabrous, erect, few- or manybranched; 

stem (sub)terete, not winged. Leaves: 

petiole absent; lower and upper blades similar, elliptic 

(or linear), to 5(–8) mm long, apex acute. 

Bracteoles present. Flowers: pedicel 10–20 mm 

long; ovary glabrous; corolla bilabiate, 5–6 mm 

long; anthers all bearded at apex. Fruit t a capsule, 

glabrous. Seeds trigonous. 

Distribution.— Myanmar, Thailand, Laos, 

Cambodia, Vietnam (Annam, Tonkin), northern 

Peninsular Malaysia. 

Ecology.— Wet and dry fallow rice fi elds, 

open grassy fi elds, lower montane pine-oak forest; 

sea level to 250–1300 m altitude. 

Notes.— Various authors, e.g. Clarke (1881) 

and Lammers (2007), united L. dopatrioides with 

L. griffthii. Apart from having different leaf size 

and shape, the former has (mostly) a different 

growth habit, stems more or less sagged at base, or 

ascending, and larger fl owers, with corolla 6–12 

mm long. 

11. Lobelia hainanensis E.Wimm., Ann. Naturhist. 

Mus. Wien 41: 348. 1948; Pfl anzenr. (Engler) 

IV.267b (107. Heft) II. Teil: 506, f. 76: l. 1953; 

Lammers, World Checkl. & Bibliogr. 

Campanulaceae: 424. 2007; D.Y.Hong & Lammers 

in D.Y.Hong, Ge Song, Lammers & L.L.Klein, Fl. 

China, 19: 557. 2011 (electronic version).— Type: 

Hainan, Liang 64115 (NY, not seen (drawing made 

from the type published in Wimmer, l.c.: fi g. 76: l. 

1953); iso S). 

Subperennial ascending or erect herb, glabrous; 

stem 4-angular. Leaves: petiole absent; lower 

and upper blades similar, spathulate-oblong or 

elliptic, ca 15 mm long. Bracteoles not seen. 

Flowers: pedicels shorter than the leaves; ovary 

glabrous; corolla bilabiate, ca 10 mm long; all anthers 

bearded at apex. Fruit t and seeds unknown. 

Distribution.— Hainan, only known from the 

type-collection. 

Ecology.— Possibly lowland; altitude not 

recorded. 

Notes.— We have seen an isotype of L. hainanensis 

at S, which bears fl owers but no fruits. 

According to its protologue and the corresponding 

drawing (Wimmer, l.c.: fi g. 76: l. 1953) this plant 

looks most similar to L. reinwardtiana from S India 

and Indonesia (Sumatra and Java). We found 

Wimmer’s description correct, except that the 

S-specimen shows all anthers bearded at apex. For 

the key to the species it is assumed that it has ellipsoid 

seeds, but this can only be ascertained when 

fruiting material becomes available. 

12. Lobelia heyneana Schult., Syst. Veg., ed. 15

46 

bis [Roemer & Schultes] 5: 50. 1819.— Type: 

Heyne s.n. in herb. Roth (B100154255, photo 

seen).— L. decurrens Roth, Nov. Pl. Sp.: 145. 

1821, nom. inval. (non Cav., Icon. 6: 13, tab. 521. 

1801).— Type: Heyne s.n. in herb. Roth 

(B100154255, photo seen).— L. trialata D.Don, 

Prodr. Fl. Nepal.: 157. 1825.— Type: Nepal, 

Hamilton s.n. (BM, not seen).— L. subincisa 

A.DC., Prodr. (DC.) 7: 367. 1836.— Type: Wallich 

1310 (holo K).— L. subracemosa Miq., Fl. Ned. 

Ind. 2: 576. 1857.— Type: Java, Horsfi eld 1339 

(holo K).— L. subracemosa var. β rigidior r Miq., 

Fl. Ned. Ind. 2: 576. 1857.— Type: Java, Horsfi eld 

728 (holo K).— L. trialata D.Don var. lamiifolia 

C.B.Clarke, Fl. Brit. India [J. D. Hooker] 3: 425. 

1881.— Type: S India, Wight 1278 (lecto K, here 

designated).— L. trialata D.Don var. asiatica 

Chiov., Res. Sci. Somalia Ital. 1: 109. 1916.— 

Locality and type not indicated.— L. bialata Merr. 

in Philipp. J. Sci., C 7(2): 105. 1912.— Type: 

Philippines, Luzon, Bontoc, Vanoverbergh 902 

(PNH, not seen).— L. micrantha Hook., Exot. Fl. 

1: t. 44. 1823, nom. inval. (non Kunth, Nov. Gen. 

Sp. [H. B. K.] 3: 316. 1820). 

Annual herb, glabrous, with one single erect 

main stem, sometimes with long lateral branches 

from above the base; stem 3-winged. Leaves: petiole 

up to 4 mm long; lower blades broadly ovate or 

rhombic, with narrowed base, decurrent on the 

petiole, upper blades similar, but somewhat narrower 

and smaller. Bracteoles present. Flowers: 

ovary glabrous; corolla bilabiate, 3–4(–5) mm 

long; only 2 anthers bearded at apex. Fruit t a cap- 

sule, glabrous. Seeds compressed, ellipsoid. 

Distribution.— N & S India; Sri Lanka; east 

to China; south-east through Indochina to Indonesia 

(the Lesser Sunda Islands); sea level to 2000 m altitude; 

not in wet localities. Also known from 

Africa (e.g. Ethiopia). 

Ecology.— Common; 750–1700 m altitude. 

Notes.— Lobelia heyneana has in the past 

been mixed up with L. stipularis (here treated as a 

synonym of L. alsinoides var. alsinoides), but the 

latter differs in having a procumbent growth form 

and a larger corolla, 5–7 mm long. Both species are 

also distinct in the shape of the seeds, which are 

ellipsoid in L. heyneana, whereas they are trigonous 

in L. alsinoides. 


13. Lobelia microcarpa C.B.Clarke, Fl. Brit. India 

[J. D. Hooker] 3: 424. 1881.— Type: Myanmar, 

Moulmein, Lobb 410 (lecto K, here designated). 

Annual herb, erect, glabrous, with one stem 

or with several main branches from the base, not 

rooting from lower nodes; stem (sub)terete, somewhat 

fl eshy at base; roots ‘spongy’, whitish. 

Leaves: petiole absent; lower blades subcircular, 

upper blades (narrowly) elliptic. Bracteoles minute. 

Flowers: pedicel 10–20(–25) mm long; ovary 

glabrous; corolla bilabiate, 4–5(–6) mm long; an- 

thers all bearded at apex. Fruit t a capsule, glabrous, 

(hemi)globose, small, 2–2.5 mm in diam. Seeds 

trigonous. 

Distribution.— India (Calcutta?, Andaman 

Islands), Myanmar (Tenasserim, Moulmein), S 

China (Canton), Thailand, Laos, Cambodia. 

Examined collections.— India: S Andaman, 

23 Nov. 1973, Balakrishnan 632 (L), deviating, 

see note; near Calcutta, anno 1937, Helfer 134 (L). 

Myanmar: Moulmein, Lobb 410 (K, lectotype). 

Thailand: Chanthaburi, 17 Oct. 1956, 200 m alt., 

Smitinand 3579 (BKF). Cambodia: Mekong River, 

12º 40 N’; 106º 05’ E, 04 August 2007, Maxwell 

07-508 (L). 

Ecology.— Wet places on poor soil at low 

altitudes. 

Notes.— 1. Lobelia microcarpa is characterized 

by its annual, erect growth habit, by circular 

lower leaves, and short corolla, 4–5(–6) mm long. 

Possibly the ‘spongy’ whitish roots are also 

diagnostic. 

2. Unusual large plants, 40 cm tall, from 

South Andaman ( (Balakrishnan 632) may belong 

here. 

14. Lobelia nummularia Lam., Encycl. [J. Lamarck 

et al.] 3(2): 589. 1792.— Type: Commerson s.n. in 

Herb. Lamarck. “ex Java” [also annotated “2, 

Piddingtonia Alph. DC.”], (P-LAM photo 

seen).— Lobelia begoniifolia Wall., Asiat. Res. 13: 

377. 1820.— Pratia begoniifolia (Wall.) Lindl., 

Edwards’s Bot. Reg. 16: t. 1373. 1830.— Type: 

Nepal, Wallich 1306 6 (holo K, 2 sheets).— Lobelia 

horsfi eldiana Miq., Fl. Ned. Ind. 2: 577. 1857.— 

Type: Java, Sello, Horsfi eld, not seen.— L. angulata 

auct. non G.Forst., Fl. Ins. Austr.: 58. 1786: 

Moeliono, Fl. Mal. Ser. 1, Spermat. 6: 133. 1960.


Low creeping herb, glabrous or hairy, rooting 

at the nodes; stem terete; roots fi brous. Leaves distichous; 

petiole distinct; lower and upper blades 

similar, subcircular, dentate on margins. Bracteoles 

absent. Flowers: ovary glabrous or hairy; corolla 

sub-bilabiate, 5–12 mm long; only 2 anthers beard- 

ed at apex. Fruit t berry-like (also capsular in 

Philippines, Sulawesi and New Guinea), ellipsoid 

or globose. Seeds compressed, ellipsoid, with reticulate 

surface. 

Distribution.— SE Asia to Australia; also 

naturalized in S America. 

Notes.— Lobelia nummularia is very variable 

in growing habit and it is recorded from sea 

level to high up in the mountains (New Guinea). 

Plants from eastern Malesia are peculiar as they 

sometimes have capsular fruits; see Moeliono 

(1960: 133, as L. angulata Forst.). 

15. Lobelia reinwardtiana (C.Presl) A.DC., Prodr. 

(DC.) 7(2): 367. 1839.— Rapuntium reinwardtianum 

C.Presl, Prodr. Monogr. Lobel.: 14. 1836.— 

Type: Java Reinwardt s.n. (holo L0846159).— 

Lobelia dichotoma Miq., Fl. Ned. Ind. 2: 576. 

1857.— Type: Indonesia, Java, Horsfi eld 327 7 (holo 

K).— L. aligera Haines, J. Proc. Asiat. Soc. Bengal 


15: 316. 1920.— L. zeylanica L. var. aligera 

(Haines) Haines, Bot. Bihar Orissa 4: 501. 1922.— 

L. dichotoma Miq. var. aligera (Haines) E.Wimm., 

Ann. Naturhist. Mus. Wien 56: 345. 1948.— Type: 

India, mountains of Chota Nagpur, Haines s.n., not 

seen.— L. dichotoma Miq. var. pilosella E.Wimm, 

Repert. Spec. Nov. Regni Veg. 38: 78. 1935.— 

Type: Java, Docters van Leeuwen 2364 (L).— L. 

zeylanica auct. non L., Sp. Pl.: 932. 1753: 

C.B.Clarke, Fl. Brit. India [J. D. Hooker] 3: 425, 

p.p. 1881. Fig. 2. 

(Sub)perennial herb, glabrous or sparsely 

hairy, branches few, ascending; stem 2- or 3-winged 

(winged when dry). Leaves subsessile, or with a 

petiole up to 5 mm long; lower blades broadly 

ovate, rhombic, or elliptic, at base long and narrowly 

decurrent on the petiole, upper blades only 

slightly narrower and somewhat smaller. Bracteoles 

absent or minute, 0.5–1 mm long, linear. Flowers: 

ovary (sub)glabrous; corolla bilabiate, 7–12 mm 

long; only two anthers bearded at apex. Fruit t a cap- 

sule, glabrous. Seeds compressed, ellipsoid. 

Distribution.— S India; Indonesia (Sumatra, 

Java). 

Ecology.— At high altitudes, (1200–)1700– 

2200 m. 

1. Corolla 7–8 mm long a. var. reinwardtiana 

1. Corolla 10–12 mm long b. var. megalantha 

a. var. reinwardtiana 

Corolla 7–8 mm long. 

Distribution.— South and Central India, 

Indonesia. 

Ecology.— Possibly confi ned to areas between 

1000 and 2000 m altitude. 

Notes.— In various fl ora treatments (e.g. 

Moeliono, 1960) material of L. reinwardtiana was 

mistaken with Lobelia heyneana, an annual species 

with one single, usually strictly erect main stem, 

and smaller fl owers. 

b. var. megalantha W.J.de Wilde & Duyfjes, 

var. nov.— A varietate typica fl oribus maioribus, 

corolla 10–12 mm longa differt.— Typus: India, 

Kerala, Umaiya Malai-Devicolam, Shetty, Bot. 

Survey India 27332 (holo K). 

Corolla 10–12 mm long. 

Distribution.— India. 

Ecology.— Flowers blue; 2150 m altitude. 

Examined collections.— Fischer 3336 6 ( K); 

Shetty, Bot. Survey India 27332 (K). 

16. Lobelia reptans W.J.de Wilde & Duyfjes, sp. 

nov.— Lobeliae reinwardtianae similis stolonibus 

longioribus validioribus, pedicellis 40–60 mm longis, 

fl oribus maioribus, fructibus ca 10 mm longis 

differt.— Typus: Papua New Guinea, New Ireland, 

East-coast, 21 Oct. 1975, Sands,Pattison, Wood & 

Croft 2455 (holo K; iso K). Fig. 3.

48 


Figure 2. Lobelia reinwardtiana (C.Presl) A.DC. var. reinwardtiana: A. Habit; B. detail of stem; C. mature fl ower bud, corolla laid 

open; D. ellipsoid seed, frontal view and cross section (all: Junghuhn s.n.; L0846173).


Figure 3. Lobelia reptans W.J.de Wilde & Duyfjes: A. Habit; B. capsule; C. seed (all: Wood & Croft 2455).

50 

Perennial herb, glabrous, with epigeous 

creeping shoots up to 80 cm long, rooting at the 

nodes, with erect shoots from nodes to 15 cm long; 

stem (1–)2–2.5 mm in diam., somewhat winged 

when dry. Leaves: petiole (1–)2–4 mm long; all 

blades ovate-elliptic. Bracteoles absent. Flowers: 

pedicel 2 or 3 times longer than sustaining leaf, 

40–60 mm long; ovary glabrous; corolla bilabiate, 

seen in bud only. Fruit t a capsule, glabrous, ob- 

ovoid, ca 10 by 4.5 mm. Seeds somewhat compressed, 

ellipsoid, with fi nely reticulately sculptured 

surface. 

Distribution.— Endemic to Papua New 

Guinea (New Ireland); only known from the 

type-collection. 

Ecology.— Middle-altitude forest, open 

mossy places; in damp disturbed vegetation on 

very steep slope at the side of stony landslip; creeping 

in partial shade; 1575 m altitude. Flowering & 

fruiting in October. 

Notes.— The seed coat of L. reptans is similar 

to type D as designated by Murata (1992). 

17. Lobelia serratifolia W.J.de Wilde & Duyfjes, 

sp. nov.— A congeneribus asiaticis minoribus foliis 

(anguste) ovato-ellipticis 8–20 mm latis margine 

serrato-dentato differt.— Typus: Papua New 

Guinea, Bougainville Isl., Port-Mine Road, 19 

June 1974, Womersley NGF 46500 (holo K, iso A, 

BRI, CANB, L). Fig. 4. 

Perennial herb, glabrous, with thin creeping 

leafl ess stem to ca 50 cm long, ascendent at apex 

and producing (10–)20–30 cm long vertical shoots, 

possibly also from the rooting nodes; stem terete, 

unwinged, 1–1.5 mm in diam. Leaves distichous, 

8–12(–20) per shoot, the lower ones withered; petiole 

(1–)2 mm long; blade narrowly ovate-elliptic, 

20–35 by 8–20 mm, base rounded, margin at each 

side with 10–20 teeth, 1–1.5 mm long, apex acuteacuminate. 

Bracteoles absent. Flowers axillary, 

solitary, glabrous; pedicel 30–60(–70) mm long; 

ovary turbinate, ca 3 mm long; sepals narrowly triangular, 

ca 3 mm long, at base 0.7(–1) mm wide; 

corolla blue, sub-unilabiate,( 8–)10–13 mm long, 

united at base for 2–3 mm (except dorsal slit), 

lobes very minutely hairy at throat, oblong-linear, 

acute; fi laments ca 2.5 mm long, connate at apex, 

only 2 fi laments fi nely hairy about halfway; 


synandrium glabrous, ca 1.5 mm long; style ca 4.5 

mm long, broadened at base, glabrous, stigma 

(stigma lobes) 2, each ca 0.3 mm long. Fruit t a cap- 

sule, long-obconical, 6–7 by 3.5 mm, at apex with 

short-conical extension above somewhat refl exed 

persistent sepals. Seeds numerous, fl attened, broadly 

ellipsoid (to nearly circular) in outline, ca 0.5 

mm long, with minutely sculptured surface. 

Distribution.— Only known from the typecollection: 

Papua New Guinea, Bougainville Isl., 

Port-Mine Road, Crown Prince Range, Kieta 

District, 6º 20’ S; 155º 30’ E. 

Ecology.— On open ground after road building; 

900 m altitude. Flowering & fruiting in June. 

Note.— The creeping growth-habit, distichous 

leaves, and sub-unilabiate corolla suggest a 

relationship with L. chinensis. 

18. Lobelia terminalis C.B.Clarke, Fl. Brit. India 

[J. D. Hooker] 3: 424 (incl. var. minuta C.B.Clarke). 

1881.— Type: Sikkim, Clarke s.n. (lectotype K, 

here designated). 

Annual herb, soft-hairy, sometimes glabrescent 

(see note), erect or suberect and branching 

from (somewhat) above the base; stem subterete; 

roots fi brous, not ‘spongy’. Leaves: petiole ± 

winged, to 2 mm long, or subsessile; lower blades 

subcircular or broadly ovate, upper blades smaller, 

the highest also narrower. Bracteoles absent or 

present (see note). Flowers: pedicel comparatively 

long, 15–40 mm long; ovary soft hairy (or glabrous, 

see note); corolla bilabiate, all lobes uniformly 

blue, white marking excepted, 4–7(–8) mm 


hairy, glabrescent or glabrous (see note). Seeds 

trigonous. 

Distribution.— India (Bengal), China 

(Yunnan), Thailand, Laos. 

Ecology.— Between 300–800 m altitude. 

Notes.— The collection Van Beusekom et al. 

3621, Kanchanaburi, from limestone, is somewhat 

deviant, since it consists of a mixture of hairy and 

subglabrous specimens. The latter have conspicuous 

bracteoles (2–)3–6 mm long, sometimes foliaceous, 

the ovaries and capsules are glabrous, and 

these plants are reminiscent of L. microcarpa. 

Possibly they are of hybrid origin; they also come


Figure 4. Lobelia serratifolia W.J.de Wilde & Duyfjes: A, B. Habit; C. leaf; D. fl ower, in dorsal view; E. capsule; F. seed (all: 

Womersley NGF 46500).

52 

close to L. thorelii, a species with a more prostrate 

growth habit and 3-coloured fl owers. 

19. Lobelia thorelii E.Wimm., Repert. Spec. Nov. 

Regni Veg. 26: 3, t. 71, f. 3. 1929.— Type: Laos, 

Expédition du Mékong, rivière d’Ubon, anno 1866, 

Thorel s.n.(holotype P P0243511, photo seen). 

Annual herb, sparingly soft-hairy, especially 

on ovary (partly glabrescent), few-branched at 

base, branches ascending but not or rarely rooting 

at the lower nodes; stem subterete (low-winged 

when dry). Leaves: petiole up to 2 mm long; lower 

blades subcircular, narrowly cuneate at base, upper 

blades smaller, only somewhat narrower, (narrowly)ovate. 

Bracteoles linear, 2 mm long. Flowers: 

pedicel 15–60 mm long; ovary soft-hairy; corolla 

bilabiate, bright blue with two white markings near 

the throat, but upper lobes lilac (pinkish), 7–10 mm 


hairy (or glabrescent). Seeds trigonous. 

Distribution.— Northeastern and eastern 

Thailand, Laos (type), can be expected in 

Cambodia; in harvested paddy fi elds. 

Ecology.— In open areas in rice fi elds, on 

small dikes and in harvested rice fi elds, also in 

open wet areas; 130–300 m altitude. 

Examined collections: Thailand: North- 

Eastern: Udon Thani, Kumphawapi, 27 Nov. 2010, 

Phonsena et al. 6614 (BKF, KKU, L); Maha 

Sarakham, Borabue, Bo Yai, Ban Bo Yai, 14 Dec. 

2010, Phonsena et al. 6640 (BKF, KKU, L). 

Eastern: Nakhon Ratchasima, Pak Thong Chai, 24 

Dec. 1924, Kerr 8099 (L); Nakhon Ratchasima, 

Korat, 22 Jan. 1931, Kerr 19927 7 ( L); Buri Ram, 27 

Nov. 1976, Phengklai et al. 3468 (BKF); ibid., 23 

Dec. 2005, Pooma et al. 6100 (BKF, L); ibid., 20 

Dec. 2005, Pooma et al. 5959 (BKF, L); Surin, 27 

Nov. 2007 Norsaengsri et al. 3126 6 ( QBG). 


Notes.— 1. Edible; the whole plant, with 

stems, leaves and fl owers, raw served with rice. 

2. Occurs occasionally gregarious, brightly 

blue colouring large patches in the fi elds. 

3. Lobelia thorelii is similar to L. terminalis, 

the latter either with a single rather erect main 

stem, or branching higher up or towards the base, 

and with smaller fl owers, uniformly blue (except 


for white markings on lower lip). 

20. Lobelia walkeri (C.B.Clarke) W.J.de Wilde & 

Duyfjes, stat. nov.— Basionym: Lobelia zeylanica 

L. var. walkeri C.B.Clarke, Fl. Brit. India [J. D. 

Hooker] 3: 425. 1881.— Type: Ceylon, Walker 

123, the lower specimen (lectotype K, here 

designated). 

Rather delicate annual or subperennial herb, 

glabrous, few-branched, ascending; stem ca 1 mm 

in diam., 3-winged (when dry). Leaves: petiole 

1–4(–7) mm long; all blades subcircular, at the 

very base shortly attenuate into petiole. Bracteoles 

ca 0.5 mm long. Flowers: ovary glabrous; corolla 

bilabiate, 6–7 mm long; only 2 anthers bearded at 

apex. Fruit t a capsule, glabrous. Seeds ± compressed, 

ellipsoid. 

Distribution.— Sri Lanka. 

Examined collections.— Cramer 4447 7 ( L); 

Hepper 4450 (K); Kostermans 23020 (L), 24230 

(L); Van Beusekom 1483 (L); Walker 123 (K, 

lectotype). 


Notes.— Clarke (l.c.) cites 2 collections: 

Walker, and Wight. The collection by Wight t ( K) is 

however L. alsinoides. 

21. Lobelia zeylanica L., Sp. Pl. 2: 932. 1753; 

Moeliono, Fl. Mal. Ser. 1, Spermat. 6: 128. 1960; 

C.E.Jarvis, Order out of chaos: 639. 2007.— Type: 

Herb. Linnaeus, no. 1051.41 (LINN), left hand 

specimen: Osbeck? “18 zeylanica” (lecto LINN, 

here designated).— L. succulenta Blume, Bijdr. Fl. 

Ned. Ind. 13: 728. 1826.— Type: Java, Buitenzorg, 

Blume 760 (holo L; iso L).— L. affi nis G.Don, 

Gen. Hist. 3: 709. 1834.— Type: India, Sylhet, 

Wallich 1311 (holo K, two sheets).— L. subcuneata 

Miq., Fl. Ned. Ind. 2: 574. 1857.— Type: Java, 

Banjoemas, Horsfi eld s.n. (U0000945 at L, photo 

seen).— L. lobbiana Hook.f. & Thomson, J. Proc. 

Linn. Soc., Bot. 2: 28. 1858.— L. affi nis G.Don 

var. lobbiana (Hook.f. & Thomson) C.B.Clarke, 

Fl. Brit. India [J. D. Hooker] 3: 424. 1881.— L. 

succulenta Blume var lobbiana (Hook.f. & 

Thomson) E.Wimm., Ann. Naturhist. Mus. Wien 

56: 361. 1948.— L. zeylanica L. var. lobbiana 

(Hook.f. & Thomson) Y.S.Lian, Fl. Reipubl.


Popularis Sin. 73(2): 149. 1938.— Type: India, 

Khasia Mountains, Lobb s.n. (holo K).— 

Dortmanna trigona (Roxb.) Kuntze var. nummulariifolia 

Kuntze, Revis. Gen. Pl. 2: 380. 1891 

(‘nummulariaefolia’).— Type from Bengal, not 

further indicated, not traced.— Lobelia macraeana 

E.Wimm, Ann. Naturhist. Mus. Wien 56: 361. 

1948; Pfl anzenr. (Engler) IV.267b (107. Heft) II. 

Teil: 575, f. 93: f. 1953.— Type: Sri Lanka, Macrae 

1227 7 (holo CGE, not seen). 

(Sub)perennial herb, usually hairy to some 

degree, ascendant, rooting at lower nodes; stem 

subterete; roots fi brous. Leaves: petiole (2–)4–10 

mm long; lower and upper blades similar, broadly 

ovate. Bracteoles absent (always?). Flowers: ovary 

usually (soft) hairy; corolla bilabiate, 5–15 mm 


usually hairy. Seeds trigonous. 

Distribution.— Widespread, from N and S 

India, Sri Lanka, east to China (including Taiwan), 

south-east to whole of Malesia (not yet found in 

Indonesia: Lesser Sunda Islands and Moluccas); 

also Solomon Islands, naturalized in Fiji. 

Ecology.— Damp, (half)shaded places, from 

sea level to 1200 m altitude. 

Notes.— 1. Lobelia zeylanica is readily distinct 

by a decumbent growth habit, some hairiness 

especially on ovary (and fruit), and distinctly petiolate 

leaves. 

2. Subglabrous forms are known from Sri 

Lanka. 

3. We regard L. lobbiana as a synonym, representing 

a large-fl owered local form, with corollas 

to 15 mm long, from Khasia Mts. Similar forms 

with large corollas are also known from S China 

and Sumatra. The larger fl owered form gradually 

merges into smaller fl owered specimens, and the 

variation apparently is found all over the wide area 

of L. zeylanica. 

4. We regard L. macraeana as a synonym, although 

in the protologue bracteoles at the base of 

the fruiting pedicel are mentioned. 

DOUBTFUL SPECIES 

Lobelia luzoniensis (Pers.) Merr., Enum. Philipp. 

Fl. Pl. 3: 588. 1923; Moeliono, Fl. Males., Ser. 1, 

Spermat. 6: 128. 1960 (in note under L. alsinoides); 

Lammers, World Checkl. & Bibliogr. 

Campanulaceae: 433. 2007.— L. fi liformis Lam. 

var. β luzoniensis Pers., Syn. Pl. 2: 214. 1806; 

Schult., Syst. Veg., ed. 15 [Roemer & Schultes] 5: 

61. 1819; A.DC., Prodr. (DC.) 7: 368. 1839.— L. 

fi liformis auct. non Lam., Encycl. [J. Lamarck & 

al.] 3(2): 588. 1792: Cav., Icon. 6: 7, t. 511, f. 1. 

1801; Miq., Fl. Ned. Ind. 2: 577. 1857; Fern.-Vill., 

Nov. App.: 121. 1880. 

Notes.— Typifi cation: Lobelia fi liformis var. 

luzoniensis was based on the description and drawing 

by Cavanilles (l.c.), who mentioned that these 

were made after a collection by Née (at MA?; not 

seen) from near Santa Cruz, at Laguna, Luzon, 

Philippines. 

The description and fi gure, which describes 

and depicts a strictly erect growing specimen with 

small roundish leaves at base and larger narrow 

leaves higher up, cannot be judged as identical 

with any later Lobelia collection from that region, 

despite the two spceimens added by Merrill (l.c.), 

and discussed by Moeliono (l.c.). 

Until truly similar plants are found in the area 

(Luzon), we regard the name as possibly based on 

a plant from a remote locality, and as doubtful for 

Asia. 

LIST OF NAMES AND SYNONYMS OF LESSER-SIZED LOBELIAS OF ASIA AND MALESIA 

Lobelia 

affi nis G.Don (1834) = L. zeylanica 

affi nis G.Don var. lobbiana (Hook.f. & Thomson) 

C.B.Clarke (1881) = L. zeylanica 

aligera Haines (1920) = L. reinwardtiana 

alsinoides Lam. (1792) 

alsinoides Lam. var. cantonensis (Danguy) 

E.Wimm. (incl. f. elongata Danguy) (1948) = L. 

dopatrioides var. cantonensis 

alsinoides Lam. var. hirta E.Wimm. (1948) [see L. 

chinensis var hirta] = possibly L. thorelii 

alsinoides subsp. hancei (H.Hara) Lammers 

(1992) = L. dopatrioides var. cantonensis

54 

alsinoides var. trigona g (Roxb.) W.J.de Wilde & 

Duyfjes (2012) 

archboldiana (Merr. & L.M.Perry) Moeliono 

(1960) 

arenarioides (C.Presl) A.DC. (1839) = L. 

heyneana 

arfakensis Gibbs (1917) = L. nummularia 

begoniifolia Wall. (1820) = L. nummularia 

bialata Merr. (1912) = L. heyneana 

brachyantha y Merr. & L.M.Perry (1941) 

caespitosa Blume (1826) = L. chinensis 

chevalieri Danguy (1929) 

chinensis Lour. (1790) 

chinensis Lour. var. cantonensis Danguy f. 

cantonensis (1930) = L. dopatrioides var. 

cantonensis 

chinensis Lour. var. cantonensis Danguy f. 

elongata Danguy (1930) = L. dopatrioides var. 

cantonensis 

chinensis Lour. var. hirta Danguy (1930) = 

possibly L. thorelii 

conferta Merr. & L.M.Perry (1949) 

decurrens Roth (1821) = L. heyneana 

dichotoma Miq. (1857) = L. reinwardtiana 

dichotoma Miq. var. aligera (Haines) E.Wimm. 

(1948) = L. reinwardtiana 

dichotoma Miq. var. pilosella E.Wimm. (1935) = 

L. reinwardtiana 

dioica R.Br. (1810) 

donanensis P.Royen (1966) 

dopatrioides p Kurz (1870) 

dopatrioides p Kurz var. cantonensis (Danguy) 

W.J.de Wilde & Duyfjes 

dopatrioides p Kurz var. kradungensis g W.J.de Wilde 

& Duyfjes (2012) 

fi liformis Lam. var. β luzoniensis Pers. (1806) = 

doubtful species 

griffi g thii Hook.f. & Thomson (1858) 

griffi thii Hook.f. & Thomson var. dopatrioides 

(Kurz) Kurz (1877) = L. dopatrioides var. 

dopatrioides 

hainanensis E.Wimm. (1948) 

hancei H.Hara (1941) = L. dopatrioides var. 


cantonensis 

heyneana y Schult. in Roem. & Schult. (incl. var. 

lamiifolia (C.B.Clarke) E.Wimm) (1819) 

horsfi eldiana Miq. (1857) = L. nummularia 

hosseusii E.Wimm. (1929) = L. dopatrioides var. 

dopatrioides 

hosseusii E.Wimm. var. villosa Kerr (1936) = L. 

dopatrioides var. dopatrioides 

javanica Thunb. (1825) = L. nummularia 

lobbiana Hook.f. & Thomson (1858) = L. 

zeylanica 

luzoniensis (Pers.) Merr. (1923) = doubtful 

species 

macraeana E.Wimm. (1948) = L. zeylanica 

micrantha Hook., non Kunth, nom. inval. = L. 

heyneana 

microcarpa p C.B.Clarke (1881) 

nummularia Lam. (1792) 

obliqua D.Don (1825) = L. nummularia 

paradoxa E.Wimm. (1929) = L. nummularia 

radicans Thunb. (1794) = L. chinensis 

reinwardtiana (C.Presl) A.DC. (1839) 

reinwardtiana (C.Presl) A.DC. var. megalantha g 

W.J.de Wilde & Duyfjes (2012) 

reptans p W.J.de Wilde & Duyfjes (2012) 

roxburghiana (A.DC.) Heynh. (1840) = L. 

chinensis 

serratifolia W.J.de Wilde & Duyfjes (2012) 

stipularis Roth ex Schult. (1819) = L. alsinoides 

var. alsinoides 

subcuneata Miq. (1857) = L. zeylanica 

subincisa A.DC. (1839) = L. heyneana 

subracemosa Miq. (incl. var. β rigidor) (1857) = 

L. heyneana 

succulenta Blume (1826) = L. zeylanica 

succulenta Blume var lobbiana (Hook.f. & 

Thomson) E.Wimm. (1948) = L. zeylanica 

terminalis C.B.Clarke (1881) 

terminalis C.B.Clarke var. minuta C.B.Clarke = L. 

terminalis 

thorelii E.Wimm. (1929) 

trialata D.Don (1825) = L. heyneana 

trialata D.Don var. asiatica Chiov. (1916) = L.

heyneana 


trialata D.Don var. lamiifolia C.B.Clarke (1881) = 

L. heyneana 

triangulata Roxb., nom. nud. (Hort. Beng.: 16. 

1814) 

trigona Roxb. (1824) = L. alsinoides var. trigona 

walkeri (C.B.Clarke) W.J.de Wilde & Duyfjes 

(2012 

zeylanica y L. (1753) 

zeylanica L. var. aligera (Haines) Haines (1921) = 

L. reinwardtiana 

zeylanica L. var. lobbiana (Hook.f. & Thomson) 

Y.S.Lian (1938) = L. zeylanica 

zeylanica L. var. parvifl ora Danguy (1930) = L. 

heyneana 

zeylanica L. var. walkeri C.B.Clarke (1881) = L. 

walkeri 

Dortmanna 

Dortmanna trigona (Roxb.) Kuntze var. nummulariifolia 

Kuntze (1891) = L. zeylanica 

Isolobus 

keri A.DC. (1839) = L chinensis 

roxburghianus A.DC. (1839) = L. chinensis 

Piddingtonia 

nummularia (Lam.) A.DC. (1839) = L. 

nummularia 

Pratia 

archboldiana Merr. & L.M.Perry (1949) = L. 

archboldiana 

begoniifolia (Wall.) Lindl. (1830) = L. 

nummularia 

ovata Elmer (1909) = L. zeylanica 

papuana S.Moore (1917) = L. nummularia 

podenzanae S.Moore (1917) = L. nummularia 

radicans G.Don (1834), (not Lobelia radicans 

Thunb.) = L. chinensis 

thunbergii G.Don (1834) = L. chinensis 

torricellensis Lauterb. (1905) = L. zeylanica 

wollastonii S.Moore (1917) = L. nummularia 

Rapuntium 

arenarioides C.Presl (1836) = L. heyneana 

reinwardtianum C. Presl (1836) = L. 

reinwardtiana 

LISTS OF LESSER LOBELIA SPECIES PER REGION 

S India, south of the Ganges River, and Sri Lanka 

alsinoides (2 varieties) 

heyneana 

reinwardtiana (2 varieties) 

walkeri 

zeylanica 

N India east to Myanmar, west of the 

Irrawaddy River 

alsinoides var. alsinoides 

chinensis 

heyneana 

nummularia 

terminalis 

zeylanica 

China (Japan) 

alsinoides var. alsinoides (no specimen seen) 

chinensis 

hainanensis (Hainan) 


heyneana 

microcarpa (Hong Kong) 

nummularia 

terminalis 

zeylanica 

Indochina, including E Myanmar east of the 

Irrawaddy River 

alsinoides var. alsinoides 

chevalieri 

chinensis 

dopatrioides (3 varieties) 

griffi thii 

heyneana 

microcarpa 

nummularia

56 

terminalis 

thorelii 

zeylanica 

Malesia, excluding New Guinea 

alsinoides var. alsinoides (Sumatra: Samosir Isl. 

only) 

chinensis 


griffi thii 

heyneana 

nummularia 

reinwardtiana var. reinwardtiana 


We thank the curators of BK, BKF, K, L, P, 

QBG, S and TCD for permitting us to study their 

collections. Mr. Brian Franzona (GH) is thanked 

for providing a photo of the type of Lobelia hancei. 

We thank the colleagues from Khon Kaen 

University and Queen Sirikit Botanic Garden, particularly 

Phongsak Phonsena and Piyakaset 

Suksathan, with whom we have conducted 

fi eldtrips. We thank Thomas Lammers (OSH) for 

valuable comments on our manuscript. Jan Frits 

Veldkamp translated the descriptions of the new 

species into Latin, the Leiden artist Jan van Os 

made the drawings and Ben Kieft (Leiden) prepared 

the drawings for publication. 

REFERENCES 

Arnott, G.A.W. (1841). In: W.J. Hooker (ed.) 

Icones Plantarum 3, pl. 358. Longman, Rees, 

Orme, Brown, Green & Longman, London. 

Clarke, C.B. (1881). Campanulaceae. In: J.D. 

Hooker (ed.) Flora of British India 3 (3): 421– 

442. Secretary of State for India in Council. 

Danguy, P. (1929). Contribution à la fl ore de l’Indo- 

Chine. Bulletin du muséum national d’histoire 

naturelle, Serie 2, 1: 263. 

Koidzumi, G. (1929). Botanical Magazine. XXX 

[Shokubutsu-gaku zasshi]. [Tokyo Botanical 

Society]. Tokyo 43: 406. 

De Loureiro, J. (1790). Flora Cochinchinensis 2: 

514. Lisboa. 

Lammers, T.G. (1992). New combinations for 


zeylanica 

New Guinea 

archboldiana 

brachyantha 

conferta 

dioica 

donanensis 


nummularia 

reptans (New Ireland) 

serratifolia (Bougainville Isl.) 

zeylanica 

Asian Campanulaceae. Botanical Bulletin of 

Academia Sinica 33: 285–287. 

Lammers, T.G. (2007). World checklist and bibliography 

of Campanulaceae: 1–675. Royal 

Botanic Gardens Kew. 

Lammers, T.G. (2011). Revision of the infrageneric 

classifi cation of Lobelia L. (campanulaceae: 

Lobelioideae). Ann. Missouri Bot. Gard. 98, 1: 

37–62. 

Loureiro see: De Loureiro. 

Moeliono, B. (1960). Lobelia. In: C.G.G.J. Van 

Steenis (ed.) Flora Malesiana, Series 1, 

Spermatophyta vol. 6: 121–136. Wolters- 

Noordhoff, Groningen. 

Murata, J. (1992). Systematic implication of seed 

coat morphology in Lobelia (Campanulaceae- 

Lobelioideae). J. Fac. Sci. Univ. Tokyo. Sect. 

3, Bot. 15: 155–172. 

Roth, A.W. (1921). Novae plantarum species: 144– 

145. Vogler, Halberstadt. 

Schultes, J.S. (1819). Lobelia. In: J.J. Roemer & 

J.A. Schultes, Systema vegetabilum ed. 15, 5: 

35–73. Stuttgart. 

Thuan, N.V. (1969). Campanulaceae. In: A. 

Aubréville & M.L. Tardieu-Blot (eds) Flore du 

Cambodge, du Laos et du Vietnam 9: (3–53). 

Muséum national d’histoire naturelle, Paris. 

Wimmer, F.E. (1953). Campanulaceae-Lobelioideae. 

In: H.G.A. Engler & F.L.E. Diels (eds) Das 

Pfl anzenreich 4, 276b: 261–813. Akademie- 

Verlag, Berlin.


The Dipterocarpaceae of Hala-Bala Forest Complex, 

Narathiwat and Yala Provinces, Peninsular Thailand 

MANOP POOPATH*, DUANGCHAI SOOKCHALOEM** & THAWATCHAI SANTISUK* 

ABSTRACT. Seven genera, 43 species and one subspecies of the family Dipterocarpaceae were recorded from the Hala-Bala forest 

complex, Narathiwat and Yala provinces, Peninsular Thailand. Eleven species and one subspecies of dipterocarps are new records 

for Thailand. Identifi cation keys to the genera and species of Dipterocarpaceae encountered in the Hala-Bala forest complex are also 

provided. 

KEY WORDS: Dipterocarpaceae, dipterocarp, Hala-Bala forest, Thailand. 

INTRODUCTION 

Botanical surveys on species diversity of 

Dipterocarpaceae in the Hala-Bala forest complex 

in Narathiwat and Yala provinces, Peninsular 

Thailand were conducted extensively during 2004– 

2006 by the fi rst author. The Hala-Bala forest complex 

which adjoins the northern border of 

Peninsular Malaysia to the south, comprises two 

major protected areas (Bang Lang National Park 

and Hala-Bala Wildlife Sanctuary) covering noncontiguous 

forest areas of about 618,750 rais or 

990 square kilometers (Fig. 1). The Dipterocarps 

of Hala-Bala Forest are predominantly Malesian in 

their affi nities as the forest is the northernmost part 

of the tropical evergreen rain forest of the Malesian 

region. Before the turn of the 21st Century, information 

and knowledge on the taxonomy, ecology 

and geography of Thai dipterocarps were rather 

sketchy. The Dipterocarpaceae of Thailand was 

fi rst treated in Florae Siamensis Enumeratio by 

Craib (1925) who recorded 10 genera, 46 species 

and 7 varieties. Fewer than half of the treated dipterocarp 

species were recorded from the Hala-Bala 

forest complex. The most recent published checklist 

of Dipterocarpaceae in Thailand provided by 

Pooma & Newman (2001) recorded 8 genera, 63 

species and 2 subspecies, of which 33 species were 

located in the Hala-Bala forest complex. The surveys 

conducted for the present study have so far 

resulted in 7 genera, 43 species and 1 subspecies 

for the Hala-Bala forest complex. It is noteworthy 

that two-thirds of the total number of dipterocarp 

species found in Thailand occur exclusively in this 

area. These dipterocarps are well established in 

moist evergreen rain forests mainly at low altitudes, 

of which only a few reach to a maximum of 

1,200 m. Ecologically the evergreen dipterocarps 

of the Hala-Bala forest complex can be divided 

into three types, namely lowland evergreen dipterocarp 

forest (below 300 m), hill evergreen dipterocarp 

forest (300–800 m), and upper evergreen 

dipterocarp forest (800–1,200 m). Most species 

thrive in the lowland and hill evergreen dipterocarp 

forest types, mostly below 500 m altitude. Eleven 

species sporadically reach upper evergreen dipterocarp 

forest, for example: Anisoptera curtisii, 

Dipterocarpus acutangulus, Parashorea stellata, 

Hopea montana, Shorea gratissima, and Vatica 

cuspidata (Fig. 2). 

DIPTEROCARPACEAE Blume 

The family Dipterocarpaceae comprises 17 

genera and 535 or more species worldwide. In 

Thailand, 63 species and 2 subspecies in 8 genera 

were previously recorded (Pooma & Newman 

2001). Seven genera, 43 species and 1 subspecies 

have now been recorded in the Hala-Bala forest 

* The Forest Herbarium, National Parks, Wildlife and Plant Conservation Department, Bangkok, Thailand 10900. Email: 

poomanop@hotmail.com 

** Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok, Thailand.

58 


Figure 1. Satellite imagery (2001) of the Hala-Bala forest complex in Narathiwat-Yala provinces.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND 

(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK) 

Figure 2. Ecological distribution of the evergreen dipterocarps in the Hala-Bala forest complex, Narathiwat and Yala provinces. 

59

60 

complex resulting in a total of 8 genera, 78 species 

and 1 subspecies of Dipterocarpaceae for Thailand. 

The only genus found in Thailand but not in Hala- 

Bala is Cotylelobium, represented by C. lanceolatum 

Craib. It is found in drier habitats of lowland, 


KEY TO THE GENERA 

semi-evergreen or seasonal rain forest on old sand 

dunes along the eastern coast of Peninsular 

Thailand, a habitat type not found in the Hala-Bala 

forest complex. 

1. Fruiting calyx lobes valvate, developed into unequal wings, or not developed. Ovary half to partly inferior 

2. Fruiting calyx lobes developed into 2 long and 3 short wings, partially united at base into a tube enclosing less than half of nut 

body, or not developed into wings; 2 long wings with 5–7 longitudinal veins. Infl orescence mostly thyrse. Stamens 15 3. Vatica 

2. Fruiting calyx lobes developed into 2 long wings and 3 much shorter wings or lobes, united at base forming a tube enclosing at 

least basal half of nut body; 2 long wings with 3 longitudinal veins. Infl orescence raceme or raceme-panicle. Stamens 15–30 

3. Fruiting calyx tube entirely adnate with nut body. Corolla not exceeding 1 cm in length, white or yellowish white. Leaf lateral 

veins joining, forming intramarginal veins; venation scalariform- reticulate 1. Anisoptera 

3. Fruiting calyx tube partially adnate with nut body. Corolla longer than 2.5 cm in length, white, tinged with pink to orange 

stripes. Leaf lateral veins not forming intramarginal veins; venation scalariform 2. Dipterocarpus 

1. Fruiting calyx lobes imbricate, developed into subequal or very unequal wings, rarely not developed. Ovary superior 

4. Fruiting calyx lobes developed into 5 subequal wings much exceeding nut body, or into 5 suborbicular lobes not exceeding nut 

body 

5. Fruiting calyx lobes developed into 5 subequally long wings, contracted at base to slender stalks not enclosing nut body; nut 

ovoid, lenticellate. Anther and ovary puberulous 4. Parashorea 

5. Fruiting calyx lobes developed into 5 coriaceous, suborbicular lobes enclosing less than half of nut body; nut ellipsoid to conical, 

smooth. Anther and ovary glabrous 5. Neobalanocarpus 

4. Fruiting calyx lobes unequally developed, at base embracing nut body tightly or loosely 

6. Fruiting calyx lobes developed into 2 long wings and 3 short lobes, or not developed. Leaf lateral veins pinnate or dryobalanoid 

to subdryobalanoid-pinnate (Fig. 3); venation reticulate or scalariform 6. Hopea 

6. Fruiting calyx lobes developed into 3 long and 2 short wings, rarely subequal, or not developed. Leaf lateral veins pinnate; 

venation obviously scalariform 7. Shorea 

Figure 3. Leaf vanation types of Hopea.



1. ANISOPTERA 

Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot.: 65. 

1841.— Mocanera Blanco, Fl. Filip., ed. 1: 446. 

1837.— Antherotriche Turcz., Bull. Soc. Imp. 

Naturalistes Moscou 2: 505. 1846.- Scaphula 

R.Parker, Repert Spec. Nov. Regni Veg. 30: 326. 


1932.— Hopeoides Cretz., J. Jap. Bot. 17(7): 408. 

1941. 

Eleven species are distributed from Bangladesh 

to Burma, Thailand, Indochina and Malesia. Four 

species are recorded for Thailand of which three 

species are in the Hala-Bala forest complex. 

1. Leaves bright golden lepidote and tomentose beneath. Flower bud long-obconical. Stamens 25; apical appendages fi liform, much 

exceeding anthers. Stylopodium short-cylindric 1. A. curtisii 

1. Leaves rusty lepidote or glabrescent beneath. Flower bud ovoid. Stamens 15; apical appendages mucronate, much shorter than 

anthers. Stylopodium discoid 

2. Leaves rusty lepidote beneath; apex long-acuminate, 5–10 mm long; lateral veins 10–12(–14) per side; venation reticulate 

2. A. laevis 

2. Leaves glabrescent beneath; apex short-acuminate, 3–5 mm long; lateral veins (12–)15–18 per side; venation scalariform-reticulate 

3. A. scaphula 

1. Anisoptera curtisii Dyer ex King, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 62: 100. 1893; Symington, 

Malayan Forest Rec. 16: 204, t. 98, f. 1. 1943; 

Smitinand, FAO-RAPA, Dipterocarps of South 

Asia: 39. 1985; Ashton, Fl. Mal. Ser. I, 

Spermatophyta 9: 329. 1982; Pooma & Newman, 

Thai For. Bull. (Bot.) 29: 115. 2001. Fig. 16. 

Thailand.— PENINSULAR: Satun, Yala (Bannang 

Sata, Than To, Betong districts), Narathiwat 

(Sukhirin & Waeng districts). 

Distribution.— Peninsular Malaysia, Sumatra. 

(Fig. 4). 

Ecology.— Common, confi ned to ridges or 

hilltops in lowland and hill evergreen dipterocarp 

forest; alt. 150–850 m. Flowering: April. Fruiting: 

July. 

Additional specimens.— Poopath-A45, 6 

May 2004, Than To, Yala (BKF); Poopath-A67, 10 

April 2005, Than To, Yala (BKF); Poopath-A71, 

10 April 2005, Than To, Yala (BKF);Poopath-A101, 

15 July 2005, Than To, Yala (BKF); Poopath-B31, 

9 Sept. 2004, Sukhirin, Narathiwat (BKF); Poopath 

-B123, 12 July 2005, Sukhirin, Narathiwat (BKF). 

2. Anisoptera laevis Ridl., Fl. Mal. Pen. 1: 219. 

1922; Symington, Malayan Forest Rec. 16: 205, t. 

96, 97, 98, f. 2. 1943; Ashton, Fl. Mal. Ser. I, 

Spermatophyta 9: 335. 1982.— A. glabra auct. non 

Kurz: Ridl., Agric. Bull. Straits Fed. Malay States. 

1: 60. 1901.— A. thurifera auct. non Blume: 

Foxw., Philipp. J. Sci., C. 6: 257, p.p. 1911.— A. 

mindanensis auct. non Foxw.: Wyatt-Sm., Malayan 

Forest Rec. 18: 77, p.p. 1955. Figs. 16, 21A. 

Thailand.— PENINSULAR: Narathiwat (Sukhirin 

& Waeng districts). 

Distribution.— Peninsular Malaysia, Sumatra, 

Borneo (Brunei, Sarawak, Sabah). (Fig. 4). 

Ecology.— Rare, sporadically found on welldrained, 

undulating to ridged terrains in lowland 

and hill dipterocarp forest; alt. 150–550 m. 

Flowering: May–June. Fruiting: October. 

Additional specimens.— Poopath-B126, 28 

Oct. 2005, Waeng, Narathiwat (BKF); Poopath-B50, 

24 July 2004, Waeng, Narathiwat (BKF); 

Poopath-B122, 12 July 2005, Sukhirin, Narathiwat 

(BKF); Poopath-B125, 10 Aug. 2005, Sukhirin, 

Narathiwat (BKF). 

3. Anisoptera scaphula (Roxb.) Kurz., Bull. Jard. 

Bot. Buitenzorg III, 9: 102, f. 6. 1927; Symington, 


Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 352. 

1982; Pooma & Newman, Thai For. Bull. (Bot.) 

29: 115, f. 1. 2001. Fig. 16. 

Thailand.— NORTHERN: Tak (Taksin National 

Park: Kra bak yai); SOUTH-WESTERN: Kanchanaburi; 

PENINSULAR: Chumphon (Lang Suan), Ranong 

(Kapoe, Kam Phuan), Surat Thani (Na San), 

Phangnga (Takua Pa), Krabi (Ao Luek, Khao 

Phanom), Nakhon Si Thammarat (Chawang), Yala 

61

62 


Figure 4. Distribution maps of Anisoptera curtisii, A. laevis, A. scaphula and Dipterocarpus acutangulus in Peninsular Thailand- 

Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors)



(Betong & Than To districts), Narathiwat (Sukhirin 


Distribution.— Bangladesh (Chittagong), 

Lower Burma, Peninsular Malaysia. (Fig. 4). 

Ecology.— Rare, sporadically found along 

stream courses on undulating to ridged terrains in 

lowland and hill evergreen dipterocarp forest; alt. 

100–650 m. Flowering: March–April. Fruiting: May. 


May. 2005, Betong, Yala (BKF); Poopath-A90, 25 

May. 2005, Betong, Yala (BKF); Poopath-B137, 

26 Aug. 2006, Waeng, Narathiwat (BKF). 

Note.— Although already previously known 

from Thailand the locality records for Narathiwat 


and Yala provinces are new. 

2. DIPTEROCARPUS 

C.F.Gaertn., Suppl. Carp.: 500. 1805.— Oleoxylon 

Roxb., Trans. Soc. London Encour. Arts 23: 413. 

1805.— Mocanera Blanco, Fl. Filip., ed. 1: 446. 

1837.— Pterygium Corrêa, Gen.: Pl. 1013. 1840.— 

Duvaliella F.Heim, Bull. Mens. Soc. Linn. Paris 

2(127): 1011. 1892. 

The genus Dipterocarpus consists of about 

70 species, distributed from Sri Lanka, India, S. 

China, Burma, Indochina and Malesia. Sixteen 

species are recorded for Thailand of which eight 

species are in the Hala-Bala forest complex. 

1. Fruiting calyx tube longitudinally and narrowly winged or angled 

2. Fruiting calyx broadly ellipsoid, 4–8 cm long, with 5 straight to slightly undulate and narrow wings exceeding 5 mm in width; 

2 large apical wings 15–20 cm long, 3–4 cm broad. Leaves usually 15–24 cm long; petioles 5–8 cm long 5. D. grandifl orus 

2. Fruiting calyx tube ovoid or subglobose, less than 3 cm long, with 5 straight and narrow wings or angles less than 5 mm broad; 

2 large apical wings less than 15 cm long. Leaves usually 6–14 cm long; petioles less than 3.5 cm long 

3. Fruiting calyx tube ovoid, 2.7–3.2 cm long, glabrous; 2 large apical wings 10–14 cm long, 2.5–3.5 cm broad. Leaves sparsely 

puberulous to glabrous beneath 1. D. acutangulus 

3. Fruiting calyx tube subglobose, 1.5–2 cm long, pubescent; 2 large apical wings 6–9 cm long, 1.8–2 cm broad. Leaves densely 

tufted hairy beneath 2. D. costatus 

1. Fruiting calyx tube smooth, never angled 

4. Stipules densely villous inside, glabrous outside. Young twigs glabrous. Leaves glabrous on both surfaces. Fruiting calyx tube 

glabrous; 2 large apical wings slightly revolute at bases; 3 short lobes prominently revolute 7. D. kerrii 

4. Stipules glabrous inside, tomentose to pubescent outside. Young twigs more or less hairy, or glabrous. Leaves more or less hairy 

or glabrous on both surfaces, or glabrous above, sparsely hairy beneath. Fruiting calyx tube pubescent to glabrous; 2 large and 

3 short apical wings revolute or not at bases 

5. Young twigs with tufted long hairs, hairs 3-4 mm long. Leaves densely tufted long hairy on both surfaces. Fruiting calyx tube 

ellipsoid; 2 long apical wings 7–9 cm long 3. D. crinitus 

5. Young twigs sparsely hairy or glabrous, hairs less than 2 mm long. Leaves subglabrous to glabrous on both sides, or tomentose 

hairy beneath. Fruiting calyx tube ovoid or subglobose; 2 long apical wings more than 11 cm long 

6. Young twigs tomentose. Leaves tomentose beneath. Fruiting calyx tube ovoid; 2 long apical wings 11–17 cm long; 3 short 

lobes slightly exceeding half the length of calyx tube 4. D. gracilis 

6. Young twigs glabrous to subglabrous. Leaves glabrous or sparsely hairy beneath. Fruiting calyx tube subglobose; 3 short 

lobes less than half the length of calyx tube 

7. Leaves glabrous on both surfaces, or pubescent along midribs and main veins beneath; lateral veins 15–21 per side; apex 

bluntly acuminate; petioles tomentose to pubescent. Fruiting calyx wings 16–23 cm long, 2.5–4.2 cm broad 

8. D. retusus 

7. Leaves glabrous, shiny above, puberulous except glabrous midribs and main veins beneath; lateral veins 9–14 per side; apex 

acute-mucronate; petioles glabrous. Fruiting calyx wings 13–18 cm long, 2–2.8 cm broad 6. D. hasseltii 

1. Dipterocarpus acutangulus Vesque, Compt. 

Rend. Hebd. Séances Acad. Sci. 78: 626. 1874; J. 

Bot. 12: 150. 1874; Symington, Malayan Forest 

Rec.16: 166, t. 76, fi g. 2. 1943; Ashton, Fl. Mal. 

Ser. I, Spermatophyta 9: 322. 1982.— D. appendiculatus 

Dyer, J. Bot. 12: 152. 1874, non Scheff. 

1870.— D. tawaensis Slooten, Bull. Jard. Bot. 

Buitenzorg. III (8): 313, f. 6. 1927.— D. helicopteryx 

Slooten, Bull. Jard. Bot. Buitenzorg III (16): 

441, f. 4. 1940. Figs. 16, 21B. 

Thailand.— PENINSULAR: Yala (Bannang 

Sata & Betong districts). 

Distribution.— Peninsular Malaysia, Borneo 

(Brunei, Kalimantan, Sabah, Sarawak). (Fig. 4). 

63

64 

Ecology.— Very rare, sporadic on high ridges 

and hilltops in hill and upper evergreen dipterocarp 

forests; alt. 600–1,050 m. Flowering: April. 

Fruiting: July–August. 


July 2004, Bannang sata, Yala (BKF); Poopath- 

A64, 20 Oct. 2004, Bannang Sata, Yala (BKF); 

Poopath-A65, 9 April 2005, Bannang sata, Yala 

(BKF); Poopath-A83, 22 May 2005, Betong, Yala 

(BKF). 

2. Dipterocarpus costatus C.F.Gaertn., Suppl. 

Carp.: 50, t. 187. 1805; Symington, Malayan Forest 

Rec. 16: 173, t. 81, f. 1. 1943; Smitinand, Thai For. 

Bull. (Bot.) 12: 33. 1980; Ashton, Fl Mal. I, 9: 321. 


29: 124. 2001.— D. insularis Hance, J. Bot. 14: 

241. 1876.— D. artocarpifolius Pierre, Fl. Forest. 

Cochinch., t. 213. 1889.— D. parvifolius F.Heim, 

Bot. Tidsskr. 25: 43. 1903. Fig. 16. 

Thailand.— NORTHERN: Chiang Mai, 

Lampang, Phrae, Tak, Sukhothai; NORTH-EASTERN: 

Phetchabun, Nong Khai, Nakhon Phanom; 

EASTERN: Nakhon Ratchasima, Ubon Ratchathani; 

SOUTH-WESTERN: Kanchanaburi; CENTRAL: 

Nakhon Nayok; SOUTH-EASTERN: Sa Kaeo, Trat; 

PENINSULAR: Chumphon, Ranong, Surat Thani, 

Phangnga, Phuket, Krabi, Nakhon Si Thammarat, 

Trang, Satun, Yala (Bannang Sata, Than To, Betong 

districts), Narathiwat (Ba Choh, Waeng, Sukhirin 

districts). 

Distribution.— Bangladesh, Burma, Andaman 

& Nicobar Islands, Laos, S. Vietnam, Cambodia, 

Peninsular Malaysia. (Fig. 5). 

Ecology.— Common often gregarious on 

ridges and hilltops in lowland and hill evergreen 

dipterocarp forests; alt. 250–850 m. Flowering: 

Febuary–March. Fruiting: May–June. 


July 2004, Than To, Yala (BKF); Poopath-A1, 20 

March 2005, Than To, Yala (BKF); Poopath-A68,10 


17 May 2005, Than To, Yala (BKF). 

3. Dipterocarpus crinitus Dyer in Hook.f., Fl. 

Brit. Ind. 1(2): 296. 1874; Symington, Malayan 

Forest Rec. 16: 175, t. 82, f. 1. 1943; Smitinand, 


Thai For. Bull. (Bot.) 12: 34. 1980; Ashton, Fl. 

Mal. Ser. I, Spermatophyta 9: 299. 1982; Pooma & 

Newman, Thai For. Bull. (Bot.) 29: 126. 2001.— 

D. hirtus Vesque, Compt. Rend. Hebd. Séancea 

Acad. Sci. 78: 627. 1874.— D. tampurau auct. non 

Korth: Burck, Ann. Jard. Bot. Buitenzorg 6: 198. 

1887. Figs. 16, 21C–D. 

Thailand.— PENINSULAR: Narathiwat (restricted 

to the border area in Waeng district of 

Narathiwat and Kelantan, Malaysia). 

Distribution.— Peninsular Malaysia, Sumatra 

and Borneo. (Fig. 5). 

Ecology.— Very rare, gregarious on undulating 

areas and low hills in lowland evergreen 

dipterocarp forest; alt. ca. 100 m. Fruiting: 

July–August. 

Additional specimen.— Poopath-B134, 23 

Aug. 2006, Waeng, Narathiwat (BKF). 

Note.— A newly recorded species for 

Thailand, Dipterocarpus crinitus can be distinguished 

by its thick undulating leaves with very 

long and dense fuscicle hairs and its ellipsoidal 

fruit. The species was previously recorded for 

Thailand from Pattani by Foxworthy (1932) and 

further cited as such by Symington (1943), 

Smithinand et al. (1980) and Ashton (1982). 

Pooma & Newman (2001) reported that there were 

no specimens of this species to be found in AAU, 

BK, BKF, C, CMU, E, K and PSU. Later on, Pooma 

(2003) deleted D. crinitus as a species for Thailand 

because of this ambiguity. This study is the fi rst confi 

rmed record of D. crinitus for Thailand with a 

distribution limited to Hala-Bala forest. 

4. Dipterocarpus gracilis Blume, Bijdr. Fl. Ned. 

Ind.: 224. 1825; Symington, Malayan Forest Rec. 

16: 177, t. 83, f. 2. 1943; Smitinand, Thai For. Bull. 

(Bot.) 12: 35.1980; Ashton, Fl. Mal. Ser. I, 

Spermatophytes 9: 301. 1982; Pooma & Newman, 

Thai For. Bull. (Bot.) 29: 127. 2001.— D. pilosus 

Roxb., Fl. Ind. ed. 1832: 615. 1832.— D. skinneri 

King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62(2): 

91. 1893; Craib, Fl. Siam. 1:137. 1925.— D. turbinatus 

var. andamanicus King, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 62(2): 92. 1893.— D. schmidtii 

F.Heim, Bot. Tidsskr. 25: 42. 1903; Craib, Fl. Siam. 

1: 137. 1925.— D. angustialatus F.Heim, Bot. 

Tidsskr. 25: 43. 1903; Craib, Fl. Siam. 1: 133.



Figure 5. Distribution maps of Dipterocarpus costatus, D. crinitus, D. gracilis and D. grandifl orus in Peninsular Thailand-Malaysia 

(♦ Pooma (2003), ▲ Symington (1943) and * Authors) 

65

66 

1925.— D. andamanicus (King) Tewary & 

A.K.Sarkar, Proc. 3rd Round-table Conf. 

Dipterocarpaceae: 542. 1987. Fig. 16. 

Thailand.— EASTERN: Nakhon Ratchasima; 

SOUTH-WESTERN: Kanchanaburi, Prachuap Khiri 

Khan; CENTRAL: Nakhon Nayok; SOUTH-EASTERN: 

Rayong, Trat; PENINSULAR: Chumphon, Ranong, 

Surat Thani, Phangnga, Nakhon Si Thammarat, 

Trang, Satun, Yala (Bannang Sata, Than To, Betong 

districts), Narathiwat (Waeng & Sukhirin districts). 

Distribution.— India (Assam), Bangladesh, 

Burma, Andaman & Nicobar Islands, Peninsular 

Malaysia, Singapore, Borneo (Kalimantan, 

Sabah), Java, Philippines. (Fig. 5). 

Ecology.— Common, scattered on undulating 

to ridged terrains or along stream courses in 

lowland and hill evergreen dipterocarp forests; altitude 

up to ca. 600 m. Flowering: January–April. 


Additional specimens.— Poopath-B1, 30 April 

2004, Waeng, Narathiwat (BKF); Poopath-B38, 22 

July 2004, Waeng, Narathiwat (BKF); Poopath 

-B42, 22 July 2004, Waeng, Narathiwat (BKF). 

5. Dipterocarpus grandifl orus (Blanco) Blanco, 

Fl. Filip., ed. 2: 314. 1845; Ridl., Fl. Mal. Pen. 1: 

216. 1922; Symington, Malayan Forest Rec. 16: 

178, t. 84, f. 1.1943; Smitinand, Thai For. Bull. 

(Bot.) 12: 36. 1980; Ashton, Fl. Mal. Ser. I, 


Thai For. Bull. (Bot.) 29: 128. 2001.— Mocanera 

grandifl ora Blanco, Fl. Filip. ed. 1: 451. 1837.— 

Dipterocarpus griffi thii Miq., Ann. Mus. Bot. 

Lugd.-Bat. 1: 213. 1864.— D. pterygocalyx Scheff., 

Natuurk. Tijdschr. Ned. Ind. 31: 347. 1870. Fig. 16. 

Thailand.— NORTH-EASTERN: Nong Khai 

(Bueng Kan), Nakhon Phanom (Phu Langka); 

PENINSULAR: Chumphon (Ban Map Ammarit), 

Ranong (Lam Liang), Surat Thani (Ban Bang Bao, 

Ratchaprapha Dam), Phangnga (Ko Surin), Phuket 

(Ko Pu), Nakhon Si Thammarat (Ron Phibun, 

Chawang), Trang (Khao Chong, Kantang, Talibong, 

Ko Libong), Satun (Ko Adang), Yala (Bannang 

Sata), Narathiwat (Waeng & Muang districts). 

Distribution.— Lower Burma, Andaman & 

Nicobar Islands, Southern Vietnam, Peninsular 

Malaysia, Singapore, Borneo (Kalimantan), 


Sumatra, Philippines. (Fig. 5). 

Ecology.— Very common, scattered or gregariously 

found on hill slopes and ridges in lowland 

and hill evergreen dipterocarp forests; alt. 

50–500 m. Flowering: April. Fruiting: May-June. 


2005, Waeng, Narathiwat (BKF); Poopath-B100, 

14 May 2005, Waeng, Narathiwat (BKF). 

6. Dipterocarpus hasseltii Blume., Fl. Javae 2: 22. 

1829; Craib, Fl. Siam. 1: 135. 1925; Symington, 


Smitinand, Thai For. Bull. (Bot.) 12: 37. 1980; 



29: 129. 2001.— D. kerrii auct. non King: Tardieu- 

Blot, Fl. Indo-Chine, Suppl. 1(3): 336, f. 33. 1943. 

Fig. 16. 

Thailand.— SOUTH-WESTERN: Ratchaburi, 

Kanchanaburi; SOUTH-EASTERN: Trat; PENINSULAR: 

Ranong, Phangnga, Nakhon Si Thammarat, Yala 

(Bannang Sata, Than To districts), Narathiwat 

(Sukhirin). 

Distribution.— Andaman & Nicobar Islands, 

Southern Vietnam, Peninsular Malaysia, Borneo, 

Sumatra, Java, Philippines. (Fig. 6). 

Ecology.— Rare, sporadically found near 

streams in hilly areas in lowland evergreen dipterocarp 

forest; alt. 150–250 m. Flowering: April–May. 


Additional specimens.— Poopath-A7, 5 May 

2004, Than To, Yala (BKF); Poopath-A69, 10 

April 2005, Than To, Yala (BKF); Poopath-A60, 6 

Aug. 2004, Than To, Yala (BKF). 

7. Dipterocarpus kerrii King, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 62(2): 93. 1893; Ridl., 

Fl.Mal. Pen. 1: 215. 1922; Craib, Fl. Siam. 1: 

136.1925; Symington, Malayan Forest Rec.16: 

181, t. 85, f. 1. 1943; Smitinand, Thai For. Bull. 



Thai For. Bull. (Bot.) 29: 131, f. 3. 2001.— D. obconicus 

Foxw., Leafl . Philipp. Bot. 6: 1951. 

1913.— D. cuneatus Foxw., Philipp. J. Sci., C 13 

(3): 178. 1918.— D. perturbinatus Foxw., Philipp. 

J. Sci., C 13(3): 177. 1918. Fig. 17.



Thailand.— SOUTH-WESTERN: Ratchaburi; 

PENINSULAR: Ranong, Surat Thani, Phangnga, 

Phuket, Krabi, Nakhon Si Thammarat, Phatthalung, 

Trang, Satun, Songkhla, Yala (Bannang Sata, Than 

To, Betong districts). 

Distribution.— Burma, Andaman & Nicobar 

Islands, Peninsular Malaysia, Singapore, Borneo 

(Brunei, Sabah, Sarawak), Philippines. (Fig. 6). 

Ecology.— Common, often gregarious along 

stream courses, on hilly terrains in lowland evergreen 

dipterocarp forest; altitude up to ca. 500 m. 

altitude. Flowering: December-February. Fruiting: 

July. 

Additional specimen.— Poopath-A26, 13 

June 2004, Banglang Dam, Mae Wad, Than To, 

Yala (BKF). 

8. Dipterocarpus retusus Blume, Catalog.: 77. 


89, f. 1. 1943; Smitinand, Thai For. Bull. (Bot.) 12: 

40. 1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 

308. 1982; Pooma & Newman, Thai For. Bull. 

(Bot.) 29: 136. 2001.— D. macrocarpus Vesque, 

Compt. Rend. Hebd. Seances Acad. Sci. 78: 627. 

1874.— D. tonkinensis A.Chev., Bull. Econ. 

Indoch. 4: 43. 1927. Fig. 17. 

Thailand.— NORTHERN: Chiang Rai, Tak; 

EASTERN: Nakhon Ratchasima; SOUTH-WESTERN: 

Kanchanaburi; SOUTH-EASTERN: Chanthaburi; 

PENINSULAR: Phangnga, Krabi, Narathiwat 

(Sukhirin). 


Distribution.— India (Assam), Burma, Laos, 

Cambodia, Vietnam, Southern China, Peninsular 

Malaysia, Sumatra, Java, Lesser Sunda Islands. 

(Fig. 6). 

Ecology.— Rare, gregarious on high ridges 

and hilltops in hill evergreen dipterocarp forest; alt. 

700–900 m. Flowering: May–July. Fruiting: 

October–November. 


May 2004, Sukhirin, Narathiwat (BKF); Poopath- 

B20, 24 Oct. 2003, Sukhirin, Narathiwat (BKF); 

Poopath-B74, 15 Oct. 2004, Sukhirin, Narathiwat 

(BKF). 

Note.— A new locality recorded from 

Sukhirin district of Narathiwat. 

3. VATICA 

L., Mant. Pl. 2: 152. 1771.— Retinodendron Korth, 

Verh. Nat. Gesch. Ned. Bezitt., Bot. 55. 1841.— 

Pteranthera Blume, Mus. Bot. 2: 30. 1852.— 

Retinodendropsis F.Heim, C.R. Assoc. Fr. Pau. 

1892: 470. 1893.— Pachynocarpus Hook.f., Trans. 

Linn. Soc. 23: 159, pl. 22. 1860.— Elaeogene 

Miq., Fl. Ned. Ind., Eerste bijv. 460. 1861.— 

Synaptea Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist. 39(2): 65. 1870. 

The genus Vatica consists of 71 species, distributed 

from Sri Lanka, India to Burma, Hainan, 

Indochina and Malesia. Thirteen species are recorded 

for Thailand of which eight species are in 

the Hala-Bala forest complex. 

1. Fruiting calyx developed into 5 subequal, long wings, or 5 very thick lobes forming a woody cup tightly enclosing nut body 

2. Fruiting calyx developed into 5 refl exed, broadly ovate-elliptic wings, slightly exceeding the length of broadly ovoid-subglobose 

nut 1. V. bella 

2. Fruiting calyx developed into a woody cup with 5 short, very thick woody lobes, enclosing more than half the length of woody nut 

3. Leaf lateral veins 11–14 per side; venation scalariform; apex obtuse, or broadly and bluntly acute; base obtuse; petioles 1.5–2 

cm long. Woody nuts 3–4 cm long, almost entirely (except the apical part) surrounded by adnate woody fruiting calyx cup 

7. V. stapfi ana 

3. Leaf lateral veins 7–9(–11) per side; venation reticulate or scalariform-reticulate; apex acute to acuminate; base acute; petiole 

0.8–1.5 cm long. Woody nut not exceeding 3 cm in length, 1/3–2/3 of nut body enclosed by adnate woody fruiting calyx cup 

with 5 short, broadly obtuse lobes 8. V. umbonata 

1. Fruiting calyx developed into 2 long and 3 short wings, all exceeding nut body 

4. Fruiting calyx wings forming a short basal tube tightly enclosing 1/2–3/4 of nut body. Corolla white 6. V. odorata 

4. Fruiting calyx wings free down to a broad base subtending nut body. Corolla white, or pinkish-red 

5. Leaves oblong or oblanceolate-oblong, 15–26 cm long, rusty pubescent beneath; lateral veins 19–25 pairs; venation obviously 

scalariform. Corolla white 5. V. nitens 

5. Leaves oblong, 4–14 cm long, puberulous or glabrous beneath; lateral veins 9–14 pairs; venation reticulate or scalariformreticulate. 

Corolla white, or pinkish-red 

67

68 


Figure 6. Distribution maps of Dipterocarpus hasseltii, D.kerrii, D. retusus and Vatica bella in Peninsular Thailand-Malaysia (♦ 

Pooma (2003), ▲ Symington (1943) and * Authors)



6. Fruiting calyx wings 3.3–4.6 cm long. Corolla white 3. V. lowii 

6. Fruiting calyx wings exceeding 5 cm in length. Corolla white or pinkish red 

7. Corolla white. Fruiting calyx wings 7–9 cm long; nut broadly ovoid, 1–1.5 cm long. Young leaves yellowish green. Leaves 

9–14 cm long, glabrous above, puberulous to glabrescent beneath 2. V. cuspidata 

7. Corolla pinkish red. Fruiting calyx wings 5–7.5 cm long; nut subglobose, 0.8–1 cm long. Young leaves reddish pink. Leaves 

6–12 cm long, glabrous on both surfaces 4. V. maingayi 

1. Vatica bella Slooten, Bull. Jard. Bot. Buitenzorg, 

III, 9: 102, f. 6. 1927; Symington, Malayan Forest 

Rec. 16: 218, t. 104. F. 2. 1943; Ashton, Fl. Mal. 

Ser. I, Spermatophyta 9: 352. 1982; Pooma, Thai 

For. Bull. (Bot.) 30: 13. 2002. Figs. 17, 22A. 

Thailand.— PENINSULAR: Narathiwat 


Distribution.— Peninsular Malaysia. (Fig. 6). 

Ecology.— Rare, sporadically found on undulating 

areas in lowland evergreen dipterocarp 

forest; altitude up to ca. 300 m. Flowering: March– 

April. Fruiting: Janualy–Febuary. 


Oct. 2004, Sukhirin, Narathiwat (BKF); Poopath- 

B133, 1 April 2006, Waeng, Narathiwat (BKF). 

2. Vatica cuspidata (Ridl.) Symington, Malayan 

Forester. 3: 200. 1934; Malayan Forest Rec.16: 

219, t. 106, f. 1. 1943; Ashton, Fl. Mal. Ser. I, 

Spermatophyta 9: 546. 1982.— Synaptea cuspidata 

Ridl., J. Straits Branch Roy. Asiat. Soc. 82: 172. 

1920; Fl. Mal. Pen. 1: 242. 1922.— S. maingayi 

Ridl., Fl. Mal. Pen. 1: 240. 1922.— Vatica maingayi 

auct. non Dyer: Slooten, Bull. Jard. Bot. Buitenzorg 

III, 9: 85, 1927. Figs. 17, 23D. 

Thailand.— PENINSULAR: Yala (Betong), 

Narathiwat (Sukhirin). 


Ecology.— Rare, frequently gregarious on 

high ridges or hilltops in hill evergreen dipterocarp 

forest; alt. 800–900 m. Flowering: April–July. 

Fruiting: September–October. 


Sept. 2005, Betong, Yala (BKF); Poopath-B86, 22 

May 2005, Betong, Yala (BKF); Poopath-B108, 24 

Oct. 2003, Sukhirin, Narathiwat (BKF); 

Poopath-B132, 31 March 2006, Sukhirin, 



Thailand. V. V cuspidata is one of the group of 

species with 2 long wings. It is very similar to V. V 

lowii and V. V maingayi. V. V cuspidata has brown dry 

leaves, a 1.5–2.5 cm long petiole, a white fl ower, 

and 6.5–9 cm long fruit wings with a sightly dilated 

base. V. V lowii has brown dry leaves, a 0.6–1.4 

cm long petiole, the fl ower colour is unknown, and 

3.3–4.6 cm long fruit wing with a slightly dilated 

base. V. V maingayi has reddish-brown dry leaves, a 

1.5–2 cm long petiole, a red fl ower, and 5–7.5 cm 

long winged fruit with a valvate non-dilated base. 

3. Vatica lowii King, J. Asiat. Soc. Bengal, Pt. 2, 

Nat. Hist. 62(2): 103, p.p. 1893; emend. Symington, 



1982.— V. perakensis King, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 62(2): 103. 1893; Symington, 

Malayan Forest Rec. 16: 226, t.111, f.1. 1943.— 

Synaptea lowii (King) Ridl., Fl. Mal. Pen.1: 241. 

1922. Figs. 17, 23C. 


(Sukhirin). 


Ecology.— Rare, scattered on hill slopes or 

ridges in lowland evergreen dipterocarp forest; alt. 

300–400 m. Flowering: April. Fruiting: April 

–September. 

Additional specimen.— Poopath-B136, 25 

Aug. 2006, Sukhirin, Narathiwat (BKF). 

Note.— The distinguishing characteristics of 

Vatica lowii are described under V. V cuspidata. V. V 

lowii was reported from Ranong province by 

Smitinand et al. (1980). However no specimens 

from Ranong have been found (Pooma & Newman 

2001). Symington (1943) and Ashton (1982) reported 

that V. V lowii was distributed in Malaysia in 

a small area of Perak, along the border with 

Kelantan. That locality is close to Narathiwat 

Province, Thailand. It is unlikely that V. V lowii is 

also to be found in Ranong in the upper part of 

Peninsular Thailand, where it would be disjunct 

69

70 

from the main population by approximately 500 

km. This study is, therefore, the fi rst confi rmed record 

of V. V lowii in Thailand where it is known only 

from Hala-Bala forest. 

4. Vatica maingayi Dyer, Fl. Brit. India. 1: 302. 

1874; Symimgton, J. Malayan Branch Roy. Asiat. 

Soc. 19(2): 151. 1943; Malayan Forest Records 16: 


Spermatophyta 9: 369, f. 47. 1982.— Synaptea 

maingayi Ridl., Fl. Mal. Pen. 1: 240. 1922.— S. 

lowii Ridl., Fl. Mal. Pen.1: 241. 1922.— Vatica 

macroptera Slooten ex Thorenaar, Meded. 

Proefstat. Boschw. 16: 120, t. 21. 1926.— V. aperta 

Slooten, Bull. Jard. Bot. Buitenzorg III(17): 250. 

1942. Figs. 17, 23A–B. 

Thailand.— PENINSULAR: Songkhla (Boripat 

waterfall), Narathiwat (Sukhirin & Waeng 

districts). 

Distribution.— Peninsular Malaysia, Singapore, 

Sumatra, Borneo (Sarawak, Sabah). (Fig. 7). 

Ecology.— Very rare, sporadic on undulating 

areas to hillsides or along stream in lowland and 

hill evergreen dipterocarp forests; alt. 100–500 m. 

Flowering: April. Fruiting: June–July. 

Additional specimens.— Poopath-B56, 8 Oct. 

2004, Sukhirin, Narathiwat (BKF); Poopath-B95, 

5 April 2005, Sukhirin, Narathiwat (BKF); Poopath- 

B106, 15 May 2005, Sukhirin, Narathiwat (BKF); 


(BKF). 


Thailand. The distinguishing characteristics of 

Vatica maingayi are given under V. V cuspidata. 

5. Vatica nitens King, J. Asiat. Soc. Bengal, Pt.2, 

Nat. Hist. 62(2): 104. 1893; Symington, Malayan 

Forest Rec. 16: 223, t. 109, f. 2. 1943; Ashton, 

Man. Dipterocarp. Brunei: 78, f. 10. 1964; Fl. Mal. 

Ser. I, Spermatophyta 9: 367. 1982.— Synaptea nitens 

(King) Ridl., Fl. Mal. Pen. 1: 241. 1922.— 

Vatica cuspidata auct. non (Ridl.) Symington: 

Browne, Forest Trees Sarawak & Brunei: 100. 

1955. Figs. 17, 22B. 





(Sarawak, Sabah). (Fig. 7). 

Ecology.— Rare, sporadic on undulating terrains 

or hill slopes in lowland evergreen dipterocap 

forest; altitude up to ca. 300 m. Flowering: April. 

Fruiting: July. 


April 2005, Waeng, Narathiwat (BKF); Poopath- 

B101, 14 May 2005, Waeng, Narathiwat (BKF); 

Poopath-B109, 9 July 2005, Waeng, Narathiwat 

(BKF). 


Thailand. The distibguishing characteristics for 

Vatica nitens are in the large oblong leaves (14–25 

x 4–7.5 cm) and the numerous pairs of secondary 

veins (19–25 pairs) in a genus where other species 

generally do not have more than 15 pairs. In addition, 

it has scalariform venation rather than the reticulate 

venation or scalariform-reticulate venation 

in other speces. It has the largest fruit wings (7.5– 

10.5 x 2–3.3 cm) of the genus in Thailand. These 

wings have a distinctly dilated and recurved base. 

6. Vatica odorata (Griff.) Symington, J. Malay. 

Br. Roy. Asiat. Soc. 19: 156. 1941; Malayan Forest 

Rec. 16: 224, t. 105, f. 2. 1943; Smitinand, Thai 

For. Bull. (Bot.) 12: 84. 1980; Ashton, Fl. Mal. Ser. 

I, Spermatophyta 9: 360. 1982; Pooma & Newman, 

Thai For. Bull. (Bot.) 29: 181. 2001; Pooma, Thai 

For. Bull. (Bot.) 30: 14. 2002.— Synaptea odorata 

Griff., Not. Pl. Asiat. 4: 516. 1854.— Vatica faginea 

Dyer in Hook.f., Fl. Brit. Ind. 1(2): 301. 1874; 

Craib, Fl. Siam. 1: 141. 1925.— V. grandifl ora 

Dyer in Hook.f., Fl. Brit. Ind. 1(2): 301. 1874.— V. 

astrotricha Hance, J. Bot. 14: 241. 1876.— 

Synaptea faginea (Dyer) Pierre, Fl. Forest. 

Cochinch. 3(15): t. 243. 1891.— S. dyeri Pierre, Fl. 

Forest. Cochinch. 3(15): t. 241. 1891.— Vatica 

curtisii King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 

62(2): 106. 1893.— Perissandra laotica Gagnep., 

Bull. Soc. Bot. France 95: 27. 1948. Fig. 17. 

Thailand.— NORTHERN: Lamphun, Phrae, 

Lampang, Phitsanulok; EASTERN: Nakhon 

Ratchasima; SOUTH-EASTERN: Chanthaburi, 

Prachinburi; SOUTH-WESTERN: Uthai Thani, 

Prachuap Khiri Khan; PENINSULAR: Chumphon, 

Ranong, Surat Thani, Krabi, Satun, Songkhla, Yala 

(Betong).



Figure 7. Distribution maps of Vatica cuspidata, V. lowii, V. maingayi, and V. nitens in Peninsular Thailand-Malaysia (♦ Pooma 

(2003), ▲ Symington (1943) and * Authors) 

71

72 

Distribution.— Burma (Tenasserim), Laos, 

Cambodia, Vietnam, S. China, Peninsular 

Malaysia, Singapore, Borneo (Brunei, Kalimantan, 

Sabah), Philippines. (Fig. 8). 

Ecology.— Common, gregarious on dry ridges 

in lowland and hill evergreen dipterocarp forests; 

alt. 450–700 m. Fruiting: September. 

Additional specimens.— Poopath-A105-a, 2 

Sept. 2005, Betong, Yala (BKF); Poopath-A105-b, 

2 Sept. 2005, Betong, Yala (BKF). 

7. Vatica stapfi ana (King) Slooten, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 62(2): 136. 1893; 

Symington, Malayan Forest Rec. 16: 228, t. 112, f. 

2. 1943; Smitinand, Thai For. Bull. (Bot.) 12: 86. 

1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 


(Bot.) 29: 184. 2001.— Pachynocarpus stapfi anus 

King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62(2): 

135. 1893; Craib, Fl. Siam. 1: 140. 1925.— P. wallichii 

(Dyer) King, J. Asiat. Soc. Bengal, Pt. 2, Nat 

Hist. 62(2): 135. 1893.— P. grandifl orus Ridl., J. 

Fed. Malay States Mus. 10: 127. 1920; Craib, Fl. 

Siam. 1:140. 1925. Figs. 17, 22C. 

Thailand.— PENINSULAR: Surat Thani, Satun, 

Yala (Betong & Than To districts), Narathiwat 

(Waeng). 

Distribution.— Cambodia, Peninsular Malaysia, 

Sumatra. (Fig. 8). 


to ridged terrains in lowland evergreen dipterocarp 

forest; alt. to ca. 300 m. Flowering: March. 

Fruiting: March–October. 


Sept. 2004, Waeng, Narathiwat (BKF); Poopath- 

B87, 8 March 2005, Waeng, Narathiwat (BKF). 

8. Vatica umbonata (Hook. f.) Burck, Ann. Jard. 

Bot. Buitenzorg 6: 232. 1887; Ashton, Fl. Mal. Ser. 

I, Spermatophyta 9: 349. 1982; Pooma, Thai For. 

Bull. (Bot.) 30: 15. 2002.— Pachynocarpus umbonatus 

Hook.f., Trans. Linn. Soc. London, Bot. 23: 

159, t. 22. 1860; Ridl., Fl. Mal. Pen. 1: 249,.1922, 

p.p..— Vatica verrucosa Burck, Ann. Jard. Bot. 

Buitenzorg 6: 232, t. 29, f. 5. 1887.— Pachynocarpus 

verrucosus (Burck) F.Heim, Rech. Dipt.: 107. 

1892; Ridl., Fl. Mal. Pen.1: 249. 1922, p.p.. 


— Vatica cupularis Slooten, Bull. Jard. Bot. 

Buitenzorg III(9): 132, t. 13. 1927.— V. stapfi ana 

Browne, Forest Trees Sarawak & Brunei: 102. 

1955. Figs. 17, 22D. 


Sata, Betong, Than To districts). 

Distribution.— Peninsular Malaysia, Borneo, 

Philippines. (Fig. 8). 

Ecology.— Rare, scattered on ridges or hilltops 

in hill evergreen dipterocarp forest; alt. 300– 

800 m. Flowering: April. Fruiting: December 

–January. 

Additional specimens.— (typical V. V umbonata): 

Poopath-A19, 19 Oct. 2004, Than To, Yala (BKF); 

Poopath-A31, 19 Oct. 2004, Bannang Sata, Yala 

(BKF); Poopath-A38, 19 Oct. 2004, Than To, Yala 

(BKF). (variant, V. V umbonata) Poopath-B28, 22 

June 2004, Waeng, Narathiwat (BKF); Poopath- 

B72, 14 Oct. 2004, Waeng, Narathiwat (BKF); 

Poopath-B135, 24 Aug. 2006, Waeng, Narathiwat 

(BKF); Poopath-B65, 10 Sept. 2006, Waeng, 


Note.— Some specimens from one tree 

exhibit woody fruiting calyx cup with more or less 

truncate lobes, thus a normal local variant. 

4. PARASHOREA 

Kurz., J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 39(2): 

65. 1870; Symington, Malayan Forest Rec. 16: 97. 

1943; Smitinand, Thai For. Bull. (Bot.) 12: 57. 

1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9(2): 

379. 1982. 

The genus Parashorea consists of 14 species, 

distributed from Burma, S. China, Indochina, and 

Malesia. Two species are recorded for Thailand of 

which one species is in the Hala-Bala forest 

complex. 

Parashorea stellata Kurz, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 39: 66. 1870; Craib, Fl. Siam. 1: 

144. 1925; Ridl., Fl. Mal. Pen. 1: 234. 1922; 




29: 154, f. 6. 2001.— Shorea stellata (Kurz) Dyer 

in Hook.f., Fl. Brit. Ind. 1: 304. 1874.— S. cinerea



Figure 8. Distribution maps of Vatica odorata, V. stapfi ana, V. umbonata and Parashorea stellata in Peninsular Thailand-Malaysia 

(♦ Pooma (2003), ▲ Symington (1943) and * Authors). 

73

74 

C.E.C. Fisch., Bull. Misc. Inform. Kew 1926: 460. 

1926.— Parashorea lucida (Miq.) Kurz, 


1. 1943.— P. poilanei Tardieu-Blot, Not. Syst. 

(Paris) 10(3): 136. 1942. Fig. 18. 

Thailand.— NORTHERN: Lamphun; SOUTH- 

WESTERN: Kanchanaburi, Ratchaburi, Prachuap 

Khiri Khan; PENINSULAR: Surat Thani, Phangnga, 

Phuket, Nakhon Si Thammarat, Trang, Satun, 

Songkhla, Yala (Bannang Sata, Than To, Betong 

districts), Narathiwat (Ban Bukit, Muang, Ba Cho, 

Waeng, Sukhirin districts). 

Distribution.— Burma, Laos, Vietnam, 


Ecology.— Very common, scattered or gregarious 

on ridges or hill slopes in lowland to upper 

evergreen dipterocarp forests; alt. 100–1,000 m. 

Flowering: May. Fruiting: July–October. 

Additional specimens.— Poopath-A4, 6 May 

2004, Than To, Yala (BKF); Poopath-A35, 18 July 

2004, Than To, Yala. (BKF); Poopath-B98, 13 

May. 2005, Sukhirin, Narathiwat (BKF); Poopath- 

B122, 11 July 2005, Sukhirin, Narathiwat (BKF); 


(BKF). 

5. NEOBALANOCARPUS 

P.S.Ashton, Gard. Bull. Singapore 31(1): 27. 

1978.— Balanocarpus King, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 62(2): 133. 1893; Ridl., Fl. Mal. 

Pen. 1: 247, 1922; Symington, Malayan Forest 

Rec. 16: 147, t. 70,71. 1943; Smitinand, Thai For. 

Bull. (Bot.) 12: 23. 1980. 

Monotypic genus found only in Peninsular 

Thailand and Malaysia. 

Neobalanocarpus heimii (King) P.S.Ashton, 

Gard. Bull. Singapore 31(1): 27. 1978; Fl.Mal. I, 9: 




(Bot.): 29, f. 5. 2001.— Balanocarpus heimii King, 

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62(2): 134. 

1893; Ridl., Fl. Mal. Pen. 1: 247. 1922; Symington, 

Malayan Forest Rec. 16: 147, t. 70,71. 1943; 

Smitinand, Thai For. Bull. (Bot.) 12: 23. 1980.— 

Balanocarpus wrayi King, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 62(2): 134. 1893.— Pierrea penangiana 

F.Heim ex Brandis, J. Linn. Soc., Bot. 31: 

110. 1895. Fig. 18. 

Thailand.— PENINSULAR: Narathiwat (Ba 

Cho, Rueso, Ban Tanyong Mas, Ban Bukit, 

Tanyong Mat, Sungai Padi, Waeng, Sukhirin 

districts). 


Ecology.— Rare, sporadic on ridges or hill 

slopes in lowland and hill evergreen dipterocarp 

forests; alt. 300–600 m. Flowering: July. Fruiting: 

Janualy–April. 


Oct. 2004, Waeng, Narathiwat (BKF); Poopath- 

B86, 12 March 2005, Waeng, Narathiwat (BKF); 

Poopath- B25, 10 July 2005, Waeng, Narathiwat 

(BKF). 

6. HOPEA 

Roxb., Pl. Coromandel 3(1): 7. 1811.— Neisandra 

Rafi n., Sylva tellur.: 163. 1838.— Hoppea Endl., 

Gen.: pl. 1014. 1840.— Hancea Pierre, For. Fl. 

Cochin. 4: sub t. 244. 1891.— Pierrea F.Heim, 

Bull. Mens. Soc. Linn. Paris 2: 958. 1891.— 

Dioticarpus Dunn, Bull. Misc. Inform. Kew 1920: 

337. 1920.— Pierreocarpus Ridl. ex Symington, 

Gard. Bull. Straits Settlem. 8(1): 30. 1934. 

The genus Hopea consists of 112 species, 

distributed from Sri Lanka, India, Bangladesh, 

Burma, S. China, Hainan, Indochina and Malesia. 

Nineteen species are recorded for Thailand of 

which nine species are in the Hala-Bala forest 

complex. 

1. Leaf lateral veins uniformly pinnate; venation obviously scalariform; leaf base symmetrical or asymmetrical. Nut pubescent or 

glabrous. Stilt roots present or lacking 

2. Leaf base symmetrically acute; domatia lacking. Corolla deep red. Fruiting calyx wings 8–10 cm long. Stilt roots present 

8. H. sublanceolata 

2. Leaf base more or less asymmetrical; domatia obvious. Corolla white, yellowish-white. Fruiting calyx wings 3–6 cm long. Stilt 

roots lacking



3. Young twigs glabrous. Leaves glabrous on both surfaces; domatia pocket-like, glabrous. Stamens 15. Nut ovoid, pubescent; 

fruiting calyx wings with 10–11 longitudinal veins 5. H. odorata 

3. Young twigs pubescent to puberulous. Leaves glabrous above, pubescent beneath; domatia with short tufted hairs. Stamens 10. 

Nut subglobose, glabrous; fruiting calyx wings with 7–9 longitudinal veins 7. H. sangal 

1. Leaf lateral veins dryobalanoid or subdryobalanoid-pinnate (Fig. 3), with long or short intermediate veins in between; venation 

obscurely scalariform or invisible; leaf base symmetrical. Nut glabrous. Stilt roots present (except in H. montana) 

4. Leaf lateral veins dryobalanoid-pinnate, with long intermediate veins in between; domatia present or lacking. Infl orescence 

bracts caducous 

5. Domatia densely white hairy. Ovary with prominent short-cylindric stylopodium 6. H. pedicellata 

5. Domatia glabrous. Ovary with slender stylopodium 

6. Three inner fruiting calyx developed into long lobes to c. 2 cm long, entirely enclosing nut body; base of nut prominent, c. 4 

mm long.Young leaves pinkish red or reddish mauve 9. Hopea sp. 

6. Three inner fruiting calyx developed into broad, coriaceous, shorter lobes, almost enclosing nut body; base of nut not prominent, 

less than 2 mm long. Young leaves pale green 

7. Leaves and fruiting calyx wings turning black to brownish black when dry, usually domatia. Fruiting calyx wings 5–7 cm 

long 2. H. dryobalanoides 

7. Leaves and fruiting calyx wings turning brown when dry, domatia absent. Fruiting calyx wings 3–4.5 cm long 

3. H. latifolia 

4. Leaf lateral veins subdryobalanoid-pinnate, with shorter intermediate veins in between; domatia present. Infl orescence bracts 

persistent 

8. Two outer calyx lobes developed into 2 large wings, 4.5–6 cm long. Young twigs peltate scaly. Leaves glabrous, shiny above, 

glaucous and usually with pale brown peltate scales beneath 4. H. montana 

8. All calyx lobes developed into 5 coriaceous, suborbicular lobes, entirely enclosing nut body. Young twigs puberulous. Leaves 

glabrous on both surfaces 1. H. bracteata 

1. Hopea bracteata Burck, Ann. Jard. Bot. 

Buitenzorg 6: 239. 1887; J. Linn. Soc., Bot. 31: 

111. 1895; Ashton, Fl. Mal. Ser. I, Spermatophyta 

9: 414. f. 70A–A4. 1982.— H. bracteata var. penangiana 

Symington, Malayan Forest Rec.16: 133, 

t. 62, f. 1. 1943.— H. bracteata var. perakensis, 

Symington; Malayan Forest Rec.16: 133, t. 62, f. 1. 

1943.— Balanocarpus curtisii King, J. Asiats. Soc. 

Bengal, Pt. 2, Nat. Hist. 62(2): 131. 1893; Ridl., Fl. 

Mal. Pen. 1: 246. 1922.— B. bracteatus Merr., 

Bibl. Enum. Born. Pl. 407. 1921.— Hopea minima 

Symington, Gard. Bull. Straits. Stettlem. 10: 337, 

pl. 11. 1939. Fig. 18, 25A. 


(Sukhirin). 

Distribution.— Peninsular Malaysia, Borneo, 


Ecology.— Very rare, gregarious on ridges or 

hill slopes in lowland evergreen dipterocarp forest; 

alt. 200–300 m. Flowering: April–May. Fruiting: 

June–July. 

Additional specimens.— Poopath-S2, 5 April 

2005, Sukhirin, Narathiwat (BKF); Poopath-B103, 

15 May 2005, Sukhirin, Narathiwat (BKF). 

Note.— A newly recorded species for Thailand. 

Hopea bracteata has many distinctive characteristics 

such as its small size as a small-medium tree 

6–26 m high, fl owers which are 25 mm diameter, 

sharp buttresses with stilt roots, and smooth bark. 

The leaves are ovate or oblong-ovate, thin and with 

a smooth surface. The leaf venation is ambigiously 

subdryobalanoid. The petals are dark red.The fruits 

are wingless and ovoid and the pink/red small protuberances 

distinguish this species from other 

Hopea spp. in Hala-Bala forest. 

2. Hopea dryobalanoides Miq., Fl. Ned. Ind. 

Eerste bijv.: 492. 1861; Symington, Malayan 

Forest Rec. 16: 123, t. 57, f. 1. 1943; Ashton, Fl. 

Mal. Ser. I, Spermatophyta 9: 402, f. 12, 13, 62 b-d. 

1982.— Hancea dryobalanoidea Pierre, Fl. Forest. 

Coch. 4, t. 244. 1891.— Hopea sarawakensis 

F.Heim, Bull. Mens. Soc. Linn. Paris 2: 971. 

1891.— H. borneensis F.Heim, Bull. Mens. Soc. 

Linn. Paris 2: 972. 1891.— H. micrantha auct. non 

Hook.f.: King, J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist. 62 (2): 126. 1893. Fig. 18, 24C. 


Sata & Than To districts). 


Borneo. (Fig. 9). 

Ecology.— Very rare, infrequently found on 

undulating areas, along stream courses, on ridges 

or hill slopes in lowland and hill evergreen 

75

76 

dipterocarp forests; alt. 150–600 m. Flowering: 

April. Fruiting: July–October. 



July 2005, Bannang Sata, Yala (BKF); Poopath- 

A104, 15 July 2005, Than To, Yala (BKF); 

Poopath-A131, 2 Oct. 2005, Bannang Sata, Yala 

(BKF). 


Generally, Hopea dryobalanoides is similar to other 

species in the genus in the dryobalanoid venation, 

the yellow colour of the leaves before falling, 

leaves which become brown to black when dried, 

and axillary domatia on the lower leaf surface. The 

stamen appendages are 2 times longer than the anthers. 

The fruits have wings (3.5–)5–6 x 0.5–1.5 

cm and the upper parts are black-brown when 

dried. H. H latifolia and H. H pedicellata of Hopea sect. 

Dryobalanoides are the most similar species. H. H 

latifolia has brown dried leaves with no domatia. 

The appendages are as long as the anthers. The 

ovary has no stylopodium. Its fruit has wings which 

are shorter and smaller (3–4.5 x 0.6–0.8 cm). H. H 

pedicellata has brown dried leaves with domatia. 

The appendages are 2–2.5 times as long as the anthers. 

The ovary has a distinct stylopodium. The 

fruit wing is a similar size to H. H latifolia. 

3. Hopea latifolia Symington, Gard. Bull. Straits 

Settlem. 10(2): 360. 1939; Malayan Forest Rec. 16: 

131, t. 61, f. 1. 1943; Smitinand, Thai For. Bull. 



Thai For. Bull. (Bot.) 29: 146. 2001.— H. intermedia 

auct. non King: Foxw., Malayan Forest Rec. 

10: 134. 1932.— H. beccarina auct. non Burck: 

Symington, Gard. Bull. Straits Settlem. 9: 325. 

1938. Fig. 18. 

Thailand.— PENINSULAR: Songkhla, Yala 

(Than To), Narathiwat (Waeng). 


(Brunei, Sarawak, Kalimantan). (Fig. 9). 

Ecology.— Very rare, infrequently found on 

ridges or hill slopes in lowland evergreen dipterocarp 

forest; alt. 100–300 m. Flowering: May–July. 




June 2005, Than To, Yala (BKF); Poopath-A102, 

15 July 2005, Than To, Yala (BKF); Poopath-B71, 

14 Oct. 2004, Waeng Narathiwat (BKF); Poopath- 

B111, 10 July 2005, Waeng, Narathiwat (BKF); 

Poopath-B116, 10 June 2005, Waeng, Narathiwat 

(BKF). 

4. Hopea montana Symington, J. Malayan Branch 

Roy. Asiat. Soc. 19(2): 141, pl. 1A. 1941; Malayan 

Forest Rec.16: 133, t. 62, f. 2. 1943; Ashton, Fl. Mal. 

Ser. I, Spermatophyta 9: 413. 1982. Figs. 18, 24D. 


Sata & Betong districts). 


(Sabah), Sumatra. (Fig. 10). 

Ecology.— Very rare, gregarious on ridges in 

hill and upper evergreen dipterocarp forests; alt. 

750–1,000 m. Fruiting: August. 


Oct. 2004, Bannang Sata, Yala (BKF); Poopath- 

A95, 14 July 2005, Bannang Sata, Yala (BKF). 


Thailand. The distinguishing characteristics of 

Hopea montana are the longitudinally cracked 

bark, the dark yellow-pink inner bark, and the lanceolate-ovate 

or lanceolate leaves. The leaves are 

5.5–8 cm long with scales on the midrib. The 

leaves are also carinate and have subdryobalanoid 

venation, the immature fruit has red-pink wings, 

and the tip of stylopodium is truncate. This species 

is found only in high montainous areas. Neither 

Symington (1943), Ashton (1982) nor in this study 

were fl owers collected so information on fl owers is 

lacking. However, it could be assumed that the 

fl owers are dark red like other species in Hopea 

subsect. Sphaerocarpae. 

5. Hopea odorata Roxb., Pl. Coromandel 3: 7, t. 

210. 1819; Craib, Fl. Siam. 1: 147. 1925; 




422, f. 73. 1982; Pooma & Newman, Thai For. 

Bull. (Bot.) 29:146. 2001.— H. faginea Hort. 

Calcuttenses ex A.DC., Prodr. 16 (2): 632. 1868. 

Fig. 18. 

Thailand.— NORTHERN: Chiang Mai, Lamphun,



Figure 9. Distribution maps of Neobalanocarpus heimii, Hopea bracteata, H. dryobalanoides and H. latifolia in Peninsular Thailand- 

Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors). 

77

78 

Lampang, Phayao, Tak, Sukhothai, Phichit, 

Nakhon Sawan; NORTH-EASTERN: Phetchabun, 

Loei; EASTERN: Chaiyaphum, Nakhon Ratchasima, 

Buri Ram, Surin; SOUTH-WESTERN: Uthai Thani, 

Kanchanaburi, Ratchaburi, Prachuap Khiri Khan; 

CENTRAL: Ang Thong, Bangkok; SOUTH-EASTERN: 

Prachin Buri, Chon Buri, Rayong, Trat; 

PENINSULAR: Chumphon, Surat Thani, Phangnga, 

Nakhon Si Thammarat, Trang, Satun, Songkhla, 

Pattani, Yala (Than To), Narathiwat (Ba Cho, Tak 

Bai, Sungai Padi districts). 

Distribution.— Bangladesh, Burma, Andaman 

& Nicobar Islands, Laos, Cambodia, Vietnam, 


Ecology.— Widespread throughout the country, 

but very rare and infrequently found in the 

Hala-Bala forest complex, in lowland evergreen 

dipterocarp forest, along stream courses; alt. 100– 

150 m. Flowering: April–May. Fruiting: June. 

Additional specimen.— Poopath-A13, 17 

May 2005, Than To, Yala (BKF). 

6. Hopea pedicellata (Brandis) Symington, Gard. 

Bull. Straits Settlem. 9(4): 327, pl. 19. 1938; 



1982, excl. syn. Hopea siamensis F.Heim; Pooma 

& Newman, Thai For. Bull. (Bot.) 29: 149. 2001.— 

H. micrantha King, J. Asiat. Soc. Bengal, Pt. 2, 

Nat. Hist. 62(2): 124. 1893; Ridl., Fl. Mal. Pen. 1: 

237. 1922, non Hook. f. 1860.— H. griffi thii var. 

pedicellata Brandis, J. Linn. Soc., Bot. 31: 69. 

1895; Ridl., Fl. Mal. Pen. 1: 238. 1922.— H. intermedia 

Brandis, J. Linn. Soc., Bot. 31: 67. 1895, 

non Miq. 1860.— H. mengerawan Brandis, J. 

Linn. Soc. Bot. 31: 70. 1895, non Miq. 1860.— H. 

pierrei Ridl., Fl. Mal. Pen. 1: 238. 1922. Fig. 18. 

Thailand.— PENINSULAR: Trang, Satun, Yala 

(Bannang Sata, Betong, Than To districts), 

Narathiwat (Sukhirin & Waeng districts). 

Distribution.— Cambodia, Peninsular Malaysia, 

Singapore, Borneo (Kalimantan, Sabah, Sarawak). 

(Fig. 10). 

Ecology.— Common, gregarious on ridges or 

hilltops in lowland and hill evergreen dipterocarp 

forests; alt. 150–600 m. Flowering: April–May. 




April 2005, Than to, Yala (BKF); Poopath-A96, 14 

July 2005, Bannang sata, Yala (BKF); Poopath- 

A100, 15 July 2005, Bannang sata, Yala (BKF); 

Poopath-A106, 3 Sept. 2005, Betong, Yala (BKF); 


(BKF); Poopath-B102, 15 May 2005, Sukhirin, 

Narathiwat (BKF); Poopath-B104, 15 May 2005, 

Sukhirin, Narathiwat (BKF); Poopath-B119, 11 

July 2005, Sukhirin, Narathiwat (BKF). 

7. Hopea sangal Korth., Verh. Nat. Gesch. Ned. 

Bezitt., Bot. 75. 1841; Symington, Malayan Forest 

Rec. 16: 141, t. 67, f. 2. 1943; Smitinand, Thai For. 

Bull. (Bot.) 12: 53. 1980; Ashton, Fl. Mal. Ser. I, 

Spermatophyta 9: 420; Pooma & Newman, Thai 

For. Bull. (Bot.) 29: 2001.— H. curtisii King, J. 

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62(2): 126. 

1893.— H. globosa Brandis, J. Linn. Soc., Bot. 31: 

61. 1895.— H. lowii Dyer ex Brandis, J. Linn. 

Soc., Bot. 31: 63. 1895.— H. minutifl ora C.E.C. 

Fisch., Bull. Misc. Inform. Kew 1927: 207. 1927. 

Fig. 18. 

Thailand.— PENINSULAR: Surat Thani, 

Phangnga, Nakhon Si Thammarat, Trang, Yala 


Narathiwat (Waeng). 

Distribution.— Burma, Peninsular Malaysia, 

Singapore, Borneo, Sumatra, Java, Lesser Sunda 

Islands. (Fig. 10). 

Ecology.— Rare, scattered along streams, 

undulating areas to ridged terrains in lowland 

and hill evergreen dipterocarp forests; alt. 150– 

550 m. Flowering: May–June. Fruiting: August 

–September. 



May 2005, Betong, Yala (BKF); Poopath-B23, 13 

May 2005, Waeng, Narathiwat (BKF); Poopath- 

B124, 12 July 2005, Waeng, Narathiwat (BKF); 

Poopath-B129, 2 Oct. 2005, Waeng, Narathiwat 

(BKF). 

8. Hopea sublanceolata Symington, Gard. Bull. 

Straits Settlem. 10: 341, pl. 13. 1939; Malayan 

Forest Records 16: 144, t. 69, f. 1. 1943; Ashton, 

Fl. Mal. Ser. I, Spermatophyta 9: 411. 1982.— H. H 

nervosa Foxw., Malayan Forest Rec. 10: 129.



Figure 10. Distribution maps of Hopea montana, H. odorata, H. pedicellata and H. sangal l in Peninsular Thailand-Malaysia ( ♦ 

Pooma (2003), ▲ Symington (1943) and * Authors) 

79

80 

1932, non King 1893. Figs. 18, 24B. 

Thailand.— PENINSULAR: Yala (Betong & 

Than To districts), Narathiwat (Waeng). 


Ecology.— Rare, scattered or gregarious on 

undulating areas along stream courses in lowland 

evergreen dipterocarp forest; altitude up to 150 m. 

Flowering: May–July. Fruiting: August–September. 



June 2004, Than To, Yala (BKF); Poopath-A48, 14 

Sept. 2004, Than To, Yala (BKF). 

9. Hopea sp. Fig. 18, 24A. 

This likely new Hopea sp. belonging to 

Hopea section Dryobalanoides subsection 

Dryobalanoides resembles the Malesian Hopea 

fl uviaris P.S.Ashton, H. H longirostrata P.S.Ashton 

and H. H sulcata Symington in general appearance. It 

differs from them in its smooth bark with deep red 

inner bark when cut; leaves with more lateral veins 

(14–18 per side), and base of the nut very thick, 

supported by remarkably swollen c. 4 mm long 

receptacle. 



Distribution.— Thailand: Narathiwat, Hala- 

Bala forest. (Fig. 11). 

Ecology.— Rare, scattered or gregarious on 

undulating areas or ridged terrains in lowland evergreen 

dipterocarp forest; altitude up to ca. 300 m. 

Flowering: March–May. Fruiting: June–August. 




July 2004, Waeng, Narathiwat (BKF); Poopath- 

B105, 15 May 2005, Sukhirin, Narathiwat (BKF); 

Poopath-B110, 10 July 2005, Waeng, Narathiwat 

(BKF); Poopath-B113, 10 June 2005, Waeng, 

Narathiwat (BKF); Poopath-B118, 11 July 2005, 

Sukhirin, Narathiwat (BKF); Poopath-B24, 22 

July 2005, Waeng, Narathiwat (BKF). 

Note.— Hopea sp. is probably a new species 

endemic to Peninsular Thailand. 

7. SHOREA 

Roxb. ex C.F.Gaertn., Suppl. Carp. 48. 1805; 



1982..— Doona Thwaites, Hooker’s J. Bot. Kew 

Gard. Misc. 4: 7. 1852.— Pentacme A.DC., Prodr. 

16(2): 626. 1886; Symington, Malayan Forest Rec. 

16: 104. 1943.— Isoptera Scheff. ex Burck, 

Meded. Lands Plantentuin 3: 27. 1886. — 

Ridleyinda Kunze, Rev. Gen. Pl. 1: 65. 1891.— 

Richetia F.Heim, Bull. Mens. Soc. Linn. Paris 2: 

975. 1891.— Anthoshorea Pierre ex F.Heim, Rech. 

Dipterocarp.: 41. 1892.— Parahopea F.Heim, 

Rech. Dipterocarp.: 66. 1892.— Pachychlamys 

Dyer ex Ridl., Fl. Mal. Pen. 1: 233. 1922. 

The genus Shorea consists of 196 species, 

widely distributed from Sri Lanka, India, Burma to 

Indochina and Malesia. Twenty fi ve taxa are recorded 

for Thailand of which fourteen species and 

one subspecies are in the Hala-Bala forest 

complex. 

1. Fruiting calyx wings erect, embracing and slightly exceeding nut body; nut ellipsoid, conical, or short-cylindric, 2.8–4.5 cm long. 

Corolla red 14. S. singkawang 

1. Fruiting calyx wings much exceeding nut body; nut ovoid, conical, subglobose to globose, not exceeding 2 cm in length (except 

in S. longisperma). Corolla white, yellowish white, pale yellow, occasionally red at upper part, pink or red at base 

2. Petals falling off separately. Stamens 25–35 

3. Leaves broadly elliptic, scabrid pubescent beneath. Corolla yellowish white in upper part, pinkish red at base. Stamens 

24–27 10. S. ochrophloia 

3. Leaves oblong, with sparse short hairs along main veins, or glabrescent beneath. Corolla yellow in upper part, red at base. 

Stamens 29–35 6. S. guiso 

2. Petals falling off in a rosette. Stamens 14–17(-25) 

4. Leaf venation loosely scalariform. Anthers with 2 pollen sacs 

5. Leaves lanceolate, oblong-lanceolate, glabrous on both surfaces; lateral veins 8–10 pairs; petioles 0.7–1 cm long. Young twig 

sparsely hairy. Nuts ellipsoid to conical, 1.5–1.8 cm long 4. S. faguetiana 

5. Leaves oblong, elliptic-oblong, glabrous above, pubescent to puberulous beneath; lateral veins 11–14 pairs; petioles 1.1–1.5 

cm long. Young twig densely hairy. Nuts broadly ovoid-conical-ellipsoid, 2.5–3 cm long 8. S. longisperma



4. Leaf venation densely scalariform. Anthers with 4 pollen sacs 

6. Leaves without minute red glands beneath. Anther ovoid-cylindric or short-cylindric, apical appendage usually much exceeding 

anther 

7. Leaf lateral veins 8–9 per side. Stamens 15; apical appendage twice to thrice as long as anther 13. S. paucifl ora 

7. Leaf lateral veins 12–20 per side. Stamens 14–25; apical appendage shorter or longer than anther 

8. Twig compressed, sparsely scaly or glabrescent. Stamens 25; apical appendage about twice as long as anther 

5. S. gratissima 

8. Twig terete, tomentose or glabrescent. Stamens 14–15 

9. Twig tomentose. Leaves broadly elliptic, 4–6.5 cm long, with short tufted hairs beneath; lateral veins 12–14 per side; 

petioles 4–7 mm long. Corolla white in upper part, pale pink or pinkish white at base. Apical appendage slightly shorter 

than anther r 1. S. assamica 

9. Twig glabrescent. Leaves oblong, elliptic-oblong, 8–16 cm long, pale green, densely white lepidote beneath; lateral veins 

16–18 pairs; petioles 12–16 mm long. Corolla white or yellowish white in upper part, red at base. Apical appendage 

about twice as long as anther 2. S. bracteolata 

6. Leaves with scattered minute red glands beneath. Anther subglobose-globose; apical appendage equalling to slightly exceeding 

anther 

10. Fruiting calyx wings broadly auriculate at base; both auricles slightly broader than oblong wings, not enclosing ovoid nut. 

Young leaves orange red. Leaves oblong, 9–15 cm long, glabrous on both surfaces 9. S. macroptera 

10. Fruiting calyx wings not auriculate at base. Young leaves pale green, yellowish green, or pinkish. Leaf blade variform, 

usually more or less hairy or scaly beneath 

11. Leaves ovate, 3–6(-9) cm long; petioles stout, 0.8–1.3 cm long. Corolla red in upper part, white at base 11. S. ovata 

11. Leaves oblong, elliptic, or lanceolate in outline, (6–)8–14 cm long; petioles slender, 1–2 cm long. Corolla pale yellow, 

yellowish white to yellowish green throughout 

12. Leaves dark green, glabrous, shiny above, white or brown tomentose to pubescent beneath. Young fruiting calyx 

wings yellowish green. Mature nut 1.4–1.8 cm long 7. S. leprosula 

12. Leaves dark green, glabrous, shiny above, pubescent to glabrescent, or lepidote beneath. Young fruiting calyx wings 

red. Mature nut 0.7–1.3 cm long 

13. Leaves lanceolate, lanceolate-elliptic, densely pale lepidote beneath; lateral veins (12–)14–16 per side; domatia 

lacking. Corolla yellowish white 3. S. curtisii 

13. Leaves oblong, elliptic, or ovate-elliptic, puberulent or glabrescent beneath; lateral veins 8–11(–12) per side; 

domatia hairy. Corolla white 12. S. parvifolia 

1. Shorea assamica Dyer subsp. globifera (Ridl.) 

Symington, Gard. Bull. Straits Settlem. 9(4): 331, 

pl. 20. 1938; Malayan Forest Rec. 16: 31, t. 15, f. 1. 



491, f. 93. 1982; Pooma & Newman, Thai For. 

Bull. (Bot.) 29: 158. 2001.— S. globifera Ridl., Fl. 

Mal. Pen. 1: 232. 1922.— S. sororia Slooten, Bull. 

Jard. Bot. Buitenzorg 18: 247, f. 9, 10. 1949. Fig. 19. 

Thailand.— PENINSULAR: Satun, Songkhla, 

Yala (Bannang Sata, Betong, Than To districts), 

Narathiwat (Sukhirin & Waeng districts). 


(Fig. 11). 


by streams, hill slopes or on ridges in lowland 

and hill evergreen dipterocarp forests; altitude 

up to ca. 600 m. Flowering: June–July. Fruiting: 

September–October. 


July 2004, Than To, Yala (BKF); Poopath-B16, 2 

May 2004, Sukhirin, Narathiwat (BKF); 

Poopath- B68, 10 Sept. 2004, Waeng, Narathiwat 

(BKF); Poopath-B75, 15 Oct. 2004, Sukhirin, 


2. Shorea bracteolata Dyer, Fl. Brit. Ind. 1(2): 

305. 1874; Ridl., Fl. Mal. Pen. 1: 229. 1922; 

Symington, Malayan Forest Rec. 16: 34, t. 16, f. 1. 

1943; Ashton, Man. Dipterocarp. Brunei 163, f. 15. 

1964; Fl. Mal. Ser. I, Spermatophyta 9: 496. 

1982.— S. foveolata Scort. ex Foxw., Malayan 

Forest Rec. 10: 183. 1932. Fig. 19, 26D. 

Thailand.— PENINSULAR: Narathiwat (Waeng). 


Sumatra, Borneo. (Fig. 11). 

Ecology.— Very rare, infrequently on undulating 

areas or ridges in lowland evergreen dipterocarp 

forest; altitude up to ca. 300 m. Flowering: 

March–April. Fruiting: May–June. 


2005, Waeng, Narathiwat (BKF); Poopath-B67, 10 

Sept. 2004, Waeng, Narathiwat (BKF); Poopath- 

B99, 14 May 2005, Waeng, Narathiwat (BKF). 

81

82 


Figure 11. Distribution maps of Hopea sublanceolata, Hopea sp., Shorea assamica and S. bracteolata in Peninsular Thailand- 

Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors).



3. Shorea curtisii Dyer ex King, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 62(2): 111. 1893; Ridley, 

Fl. Mal. Pen. 1: 223. 1922; Craib, Fl. Siam. 1: 142. 


33, f. 1. 1943; Ashton, Gard. Bull. Singapore 31(1): 




(Bot.) 29: 158. 2001. Fig. 19. 

Thailand.— PENINSULAR: Satun, Yala (Bannang 

Sata, Than To, Betong districts), Narathiwat 

(Sukhirin, Sungai Padi, Waeng districts). 


Borneo (Brunei, Sarawak). (Fig. 12). 

Ecology.— Very common, locally frequent 

on ridges or hilltops in stands in lowland and hill 


Flowering: May–July. Fruiting: August–September. 




B27, 8 June 2004, Waeng, Narathiwat (BKF); 

Poopath-B58, 8 Sept. 2004, Sukhirin, Narathiwat 

(BKF); Poopath-B112, 11 July 2005, Waeng, 


4. Shorea faguetiana F.Heim, Bull. Mens. Soc. 

Linn. Paris 2: 975. 1891; Symington, Malayan 

Forest Rec. 16: 50, t. 23, f. 1. 1943; Smitinand, 

Thai For. Bull. (Bot.) 12: 71. 1980; Ashton, Fl. 

Mal. Ser. I, Spermatophyta 9: 484. 1982; Pooma & 

Newman, Thai For. Bull. (Bot.) 29: 159. 2001; 

Pooma, Thai For. Bull. (Bot.) 30: 12. 2002.— S. 

ridleyana King, J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist. 62(2): 115. 1893.— S. dryobalanoides Dyer 

ex Brandis, J. Linn. Soc., Bot. 31: 95. 1895; Ridl., 

Fl Mal. Pen. 1: 226.1922. Fig. 19. 


Sata, Betong, Than To districts), Narathiwat 



(Sabah, Sarawak), Sumatra. (Fig. 12). 


areas or ridges in lowland and hill evergreen 

dipterocarp forests; altitude up to 600 m. Flowering: 

April–May. Fruiting: July–August. 




B115, 10 July 2005, Waeng, Narathiwat (BKF). 

5. Shorea gratissima (Wall. ex Kurz) Dyer in 

Hook. f., Fl. Brit. Ind. 1(2): 307. 1874; Ridl., Fl. 

Mal. Pen. 1: 226. 1922; Symington, Malayan 

Forest Rec.16: 36, t. 17, f. 1. 1943; Smitinand, Thai 

For. Bull. (Bot.) 12: 72. 1980; Ashton, Fl. Mal. Ser. 

I, Spermatophyta 9: 487. 1982; Pooma & Newman, 

Thai For. Bull. (Bot.) 29: 161, f. 7. 2001.— Hopea 

gratissima Wall. ex Kurz, J. Asiat. Soc. Bengal, Pt. 

2, Nat. Hist. 42: 61. 1873. Fig. 19. 

Thailand.— SOUTH-WESTERN: Prachuap Khiri 

Khan; PENINSULAR: Chumphon, Surat Thani, 

Phuket, Nakhon Si Thammarat, Phatthalung, 

Trang, Songkhla, Yala (Bannang Sata, Betong, 

Than To districts), Narathiwat (Sukhirin & Waeng 

districts). 

Distribution.— Burma (Tenasserim), Peninsular 

Malaysia, Singapore, Borneo (Sabah), Sumatra. 

(Fig. 12). 


on ridges or hilltops in lowland to upper 


Fruiting: October–November. 


Oct. 2003, Sukhirin, Narathiwat (BKF); Poopath- 

B62, 9 Sept. 2004, Sukhirin, Narathiwat (BKF); 


(BKF). 

Note.— New localities recorded from 

Narathiwat and Yala provinces. 

6. Shorea guiso (Blanco) Blume, Mus. Bot. 2: 34. 

1852; Symington, Malayan Forest Rec. 16: 16, t. 7, 

f. 2. 1943; Smitinand, Thai For. Bull. (Bot.) 12: 64. 


447.1982; Pooma & Newman, Thai For. Bull. 

(Bot.) 29: 163. 2001.— Mocanera guiso Blanco, 

Fl. Filip., ed. 1: 449. 1837.— Dipterocarpus guiso 

(Blanco) Blanco, Fl. Filip., ed. 2: 313. 1845.— 

Anisoptera guiso (Blanco) A.DC., Prodr. 16(2): 

616. 1868.— Shorea pierrei Hance, J. Bot. 16: 302. 

1878.— S. vulgaris Pierre ex Laness., Pl. Util. Col. 

Franc.: 301. 1886.— S. obtusa var. kohchangensis 

83

84 


Figure 12. Distribution maps of Shorea curtisii, S. faguetiana, S. gratissima and S. guiso in Peninsular Thailand-Malaysia (♦ Pooma 

(2003), ▲ Symington (1943) and * Authors).



F.Heim, Bot. Tidsskr. 25: 45. 1903.— S. robusta 

var. schmidtii F.Heim, Bot. Tidsskr. 25: 45. 1903.— 

S. longipetala Foxw., Malayan Forest Rec. 10: 174. 

1932. Fig. 19. 

Thailand.— NORTH-EASTERN: Nakhon Phanom; 

SOUTH-EASTERN: Prachin Buri, Chanthaburi, Trat; 

PENINSULAR: Trang, Satun, Songkhla, Yala 


Narathiwat (Rueso, Sukhirin, Waeng districts). 

Distribution.— Vietnam, Cambodia, Peninsular 

Malaysia, Sumatra, Borneo, Philippines. (Fig. 12). 

Ecology.— Common, scattered along streams 

courses to ridges in lowland and hill evergreen dipterocarp 

forests; altitude up to 600 m. Flowering: 

April. Fruiting: June–July. 


June 2005, Than To, Yala (BKF); Poopath-B2, 3 

April 2005, Waeng, Narathiwat (BKF); Poopath- 

B114, 10 July 2005, Waeng, Narathiwat (BKF);. 

7. Shorea leprosula Miq., Fl. Ned. Ind. Eerst bijv.: 

487. 1861; Ridl., Fl. Mal. Pen. 1: 222. 1922; 





(Bot.) 29: 166. 2001.— Hopea maranti Miq., Fl. 

Ned. Ind., Eerste bijv.: 489. 1860.— Shorea maranti 

(Miq.) Burck, Ann. Jard. Bot. Buitenzorg 6: 

217. 1887.— S. astrosticta Scort. ex Foxw., 

Malayan Forest Rec. 10: 220. 1932. Fig. 19. 

Thailand.— PENINSULAR: Songkhla, 

Pattani, Yala (Bannang Sata, Betong, Than To districts), 

Narathiwat (Sukhirin, Sungai Padi, Waeng 

districts). 


Borneo, Sumatra, Java. (Fig. 13). 

Ecology.— Very common, scattered by streams, 

undulating to ridged terrains or hilltops in lowland 

to upper evergreen dipterocarp forests; altitude up 

to 1,000 m. Flowering: July. Fruiting: October. 





(BKF). 

8. Shorea longisperma Roxb., Fl. Ind. 2: 618. 

1832; Ridl., Fl. Mal. Pen. 1: 143. 1922.— 

Parashorea longisperma Kurz, J. Asiat. Soc. 

Bengal, Pt. 2, Nat. Hist. 39: 66. 1870.— Shorea 

resina-negra Foxw., Malayan Forest Rec. 10: 205, 

pl. 16. 1932; Symington, Malayan Forest Rec. 16: 


Spermatophyta 9: 481. 1982. Figs. 19, 26C. 




(Brunei, Sarawak), Sumatra. (Fig. 13). 

Ecology.— Very rare, infrequently on hill 

slopes in lowland evergreen dipterocarp forest; alt. 

ca. 300 m. Fruiting: July–October. 



B45, 23 July 2004, Sukhirin, Narathiwat (BKF); 




9. Shorea macroptera Dyer in Hook. f., Fl. Brit. 

Ind. 1(2): 308. 1874; Ridl., Fl. Mal. Pen. 1: 225. 


38, f. 1. 1943; Smitinand, Thai For. Bull. (Bot.) 12: 

79. 1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 


(Bot.) 29: 167. 2001.— S. auriculata Scort. ex 

Foxw., Malayan Forest Rec. 10: 195. 1932. Fig. 19. 

Thailand.— PENINSULAR: Yala (Betong). 


Sumatra, Borneo. (Fig. 13). 

Ecology.— Very rare, often gregarious on 

undulating areas or hill slopes in hill evergreen dipterocarp 

forest; alt. ca. 500 m. Fruiting: 

June–August. 


June 2005, Betong, Yala (BKF); Poopath-A40, 19 

July 2005, Betong, Yala (BKF). 

10. Shorea ochrophloia Strugnell ex Symington, 

Gard. Bull. Straits Settlem. 8: 268, pl. 17. 1935; 

Malayan Forest Rec. 16: 112. 1943; Ashton, Fl. 

Mal. Ser. I, Spermatophyta 9: 447. 1982. Figs. 20, 

25C–D. 

85

86 


Figure 13. Distribution maps of Shorea leprosula, S. longisperma, S. macroptera and S. ochrophloia in Peninsular Thailand- 

Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors).



Thailand.— PENINSULAR: Narathiwat (Waeng). 


(Fig. 13). 

Ecology.— Very rare, infrequently on undulating 

terrain in lowland evergreen dipterocarp 

fores; alt. ca. 150 m. Flowering: April. Fruiting: July. 



B96, 5 April 2005, Waeng, Narathiwat (BKF). 


Thailand. Shorea ochrophloia is similar to S. guiso 

in the characetrs that defi ne the section. The differences 

are that S. ochrophloia is tomentose on the 

lower surface of the leaves and young twigs whereas 

S. guiso is sparsely pubescent. Also S. ochrophloia 

has relatively larger and thicker leaves compared to 

S. guiso. The leaves of S. ochrophloia are relatively 

thick and slightly coriaceous and undulating along 

the secondary veins but the leaves of S. guiso are 

thinner and smooth. The tips of the petals are light 

yellow or whitish yellow in S. ochrophloia and 

bright yellow in S. guiso. 

11. Shorea ovata Dyer ex Brandis, J. Linn. Soc. 

Bot. 31: 91. 1895; Symington, Malayan Forest 

Rec. 16: 82, t. 39, f.. 2. 1943; Ashton, Man. 

Dipterocarp. Brunei 230, f. 16. 1964; Fl. Mal. Ser. 

I, Spermatophyta 9: 545, f. 114. 1982.— S. parvifolia 

King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 

62(2): 112. 1893, non Dyer 1874.— S. plagata 

Foxw., Philipp. J. Sci., C. 13: 192. 1918.— S. agsaboensis 

W.L.Stern, Brittonia 17: 36, f. 1,2. 1965.— 

S. parvifolia subsp. velutinata auct. non P.S.Ashton: 

Pooma & Newman, Thai For. Bull. (Bot.) 29: 169. 

2001. Fig. 20, 26B. 


KEY TO THE SUBSPECIES 

Than To district), Narathiwat (Sukhirin). 


Borneo, Philippines (Mindanao). (Fig. 14). 

Ecology.— Rare, gregarious on dry ridges or 

hilltops in hill and upper evergreen dipterocarp forests; 

alt. 500–900 m. Flowering: May–October. 

Fruiting: September–December. 



Sept. 2004, Than To, Yala (BKF); Poopath-A66, 

10 April 2005, Than To, Yala (BKF); Poopath-A85, 

22 May. 2005, Betong, Yala (BKF, spirit specimen); 

Poopath-A94, 20 June 2005, Than To, Yala; 

Poopath-A108, 10 Sept. 2005, Betong, Yala (BKF). 


Shorea ovata is very similar to S. parvifolia but S. 

ovata has dark yellow inner bark, the leaves are 

usually small and 3–6–(9) cm long, and the base of 

the leaves is cuneate or obtuse. In S. parvifolia the 

inner bark is dark red, the leaves are relatively larger 

(6–13 cm), and the leaf base is truncate or cordate. 

Other distinct characteristics of S. ovata are the red 

fl owers, petals 5–6 mm long, margin entire, and 

fruits globose or broadly ovoid. S. parvifolia has 

white fl oweres, petals 8 mm long, margin dentate 

and contorted and fruits ovoid. 

12. Shorea parvifolia Dyer in Hook. f., Fl. Brit. 

Ind. 1(2): 305. 1874; Ridl., Fl. Mal. Pen. 1: 224. 

1922; Foxw., Malayan Forest Rec. 3: 31. 1927; 


1943; Ashton, Gard. Bull. Singapore 20: 278: 

1963; Man. Dipt. Brun.: 206, f. 16. 1964; 




29: 169. 2001. 

1. Leaves sparsely pubescent or glabrescent beneath; base obtuse or slightly cordate; margin not revolute; domatia not prominent 

S. parvifolia subsp. parvifolia 

1. Leaves scabrous hairy beneath; base acute or cuneate; margin often narrowly revolute; domatia prominent 

S. parvifolia subsp. velutinata 

subsp. parvifolia.— Shorea sutulata King, J. 

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 62(2): 110. 

1893; Ridl., Fl. Mal. Pen. 1: 222. 1922. Foxw., 

Malayan Forest Rec. 10:201. 1932.— S. gentilis 

Parijs, Repert. Spec. Nov. Regni Veg. 33: 224. 

1933. Fig. 20. 

Thailand.— PENINSULAR: Songkhla, Yala 


87

88 

Narathiwat (Cho Ai Rong, Sungai Padi, Sukhirin, 

Waeng districts). 


Borneo (Brunei, Kalimantan, Sabah, Sarawak), 


Ecology.— Very common, scattered on 

undulating to ridged terrains or by streams in lowland 

and hill evergreen dipterocarp forests; altitude 

up to ca. 700 m. Flowering: April–July. Fruiting: 

September–October. 


June 2004, Than To, Yala (BKF); Poopath-B46, 23 

July 2004, Sukhirin, Narathiwat (BKF); Poopath- 




Narathiwat (BKF); Poopath-B88, 3 April 2005, 

Waeng, Narathiwat (BKF). 

subsp. velutinata P.S.Ashton, Gard. Bull. 

Singapore 20: 278. 1963; Fl. Mal. Ser. I, 

Spermatophyta 9: 547. 1982. Fig. 20, 26A. 

Thailand.— PENINSULAR: Yala (Betong). 


(Brunei, Kalimantan, Sabah), Sumatra. (Fig. 13). 

Ecology.— Very rare, infrequently on un 

dulating terrain in hill evergreen dipterocarp forests; 

alt. c. 500 m. Flowering: April. Fruiting: June 

–July. 


April 2005, Betong, Yala (BKF); Poopath-A93, 11 

June 2005, Betong, Yala (BKF); Poopath-A93-1, 

11 June 2005, Betong, Yala (BKF). 

Note.— A newly recorded subspecies for 

Thailand. Shorea parvifolia subsp. velutinata is 

distinct in the following characters: large leaves 

with obtuse or acute base, lower leaf surface with 

tomentose hairs along the midrib, prominent secondary 

and tertiary veins. S. parvifolia subsp. parvifolia 

normally has smaller leaves with obtuse or 

subcordate base, the lower surface is glabrescent 

and the secondary veins are rather fl at. S. parvifolia 

subsp. velutinata was recorded by Pooma & 

Newman (2001) but their material (Niyomdhum 

( 

4837, 20 Oct. 1996, Sukhirin, Narathiwat (BKF); 

Niyomdhum 5541, 21 Aug. 1998, Sukhirin, 

Narathiwat (BKF) Niyomdhum 5546, 21 Aug. 1998, 


Sukhirin, Narathiwat (BKF)) is, in fact, S. ovata. 

13. Shorea paucifl ora King, J. Asiat. Soc. Bengal, 

Pt. 2, Nat. Hist. 62(2): 116. 1893; Ridl., Fl. Mal. 

Pen. 1: 228. 1922; Symington, Malayan Forest 

Rec. 16: 87, t. 41, f. 1. 1943; Ashton, Man. 

Dipterocarp. Brunei 207, f. 17, pl. 46. 1964; Fl. Mal. 

Ser. I, Spermatophyta 9: 511. 1982. Fig. 20, 25B. 


Than To districts). 

Distribution.— Peninsular Malaysia, 

Singapore, Sumatra, Borneo. (Fig. 14). 

Ecology.— Rare, scattered on undulating to 

ridged terrains in lowland and hill evergreen dipterocarp 

forests; alt. 200–600 m. Flowering: May. 



June 2005, Than To, Yala (BKF); Poopath-A70, 10 


17 May 2005, Than To, Yala (BKF); Poopath-A99, 

15 July 2005, Than To, Yala (BKF). 


Thailand. Shorea paucifl ora is distinct in its very 

brown-red dried leaves, tertiary venation densely 

diagonal, and with tomentose domatia in secondary 

vein axils. The fruits have very long wings (6– 

10 cm). The bark is longitudinally scaly and with a 

reddish pink inner bark. 

14. Shorea singkawang (Miq.) Miq., Ann. Mus. 

Bot. Lugd.-Bat. 3: 84. 1867; Symington, Malayan 

Forest Rec. 16: 92, t. 43, f. 1. 1943; Ashton, Gard. 

Bull. Singapore 31(1): 47. 1978; Smitinand, Thai 

For. Bull. (Bot.) 12: 80. 1980; Ashton, Fl. Mal. I. 9: 


(Bot.) 29: 176. 2001.— Hopea singkawang g Miq., 

Sum. 1: 489. 1860.— Shorea thiseltonii King, J. 

Asiat. Soc. Bengal 62(2): 122. 1893.— S. forbesii 

King ex Brandis, J. Linn. Soc., Bot. 31: 77. 1895.— 

Pachychlamys thiseltonii (King) Ridl., Fl. Mal. 

Pen. 1: 233. 1922.— P. beccarianus auct. non Dyer 

ex Brandis: Ridl., Fl. Mal. Pen. 1: 233. 1922.— 

Shorea hemsleyana auct. non (King) King ex 

Foxw.: Smitinand, Thai For. Bull. (Bot.) 12: 78. 

1980, p.p. Fig. 20.



Figure 14. Distribution maps of Shorea ovata, S. parvifolia ssp. parvifolia, S. parvifolia ssp. velutinata and S. paucifl ora in 

Peninsular Thailand-Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors). 

89

90 




(Fig. 15). 

Ecology.— Rare, infrequently on undulating 

to ridged terrains in lowland and hill evergreen dipterocarp 

forests; alt. 200–600 m. Flowering: April– 

May. Fruiting: July. 


April 2005, Sukhirin, Narathiwat (BKF); 

Poopath-B61, 3 May 2004, Sukhirin, Narathiwat 

(BKF); Poopath-B107, 15 May 2005, Sukhirin, 

Narathiwat (BKF); Poopath-B121, 11 July 2005, 

Sukhirin, Narathiwat (BKF). 

Figure 15. Distribution maps of Shorea singkawang g in 

Peninsular Thailand-Malaysia (♦ Pooma (2003), ▲ Symington 

(1943) and * Authors). 


ACKNOWLEDGMENTS 

The authors would like to thank Dr Weerachai 

Nanakorn for valuable suggestions. Sincere thanks 

are due to all staff at Hala-Bala Wildlife Research 

Station and the 7th Chulabhorn Development 

Project for their very kind fi eld work support. We 

also thank Dr Rachun Pooma for his support for the 

fi eld surveys and the provision of valuable relevant 

research information. We would like to extend our 

thanks and appreciation to Dr David Middleton for 

his comments on the manuscript. Thanks also to all 

staff members at the BKF herbarium for the herbarium 

and library facilities. And thanks to the 

Graduate School of Kasetsart University for fi nancial 

support the fi rst author’s research study. 

REFERENCES 

Ashton, P.S. (1982). Dipterocarpaceae. Flora 

Malesiana Ser. I, 9: 237–552. 

Craib, W.G. (1925). A list of the plants known from 

Siam with records of their occurrence. Florae 

Siamensis Enumeratio. Siam Society, 

Bangkok: 133–148. 

Foxworthy, F.W. (1932). Dipterocarpaceae of the 

Malay Peninsula. Malayan Forest Records 10: 

1–289. 

Pooma, R. & M. Newman. (2001). Checklist of 

Dipterocarpaceae in Thailand. Thai For. Bull. 

(Bot.) 29: 110–187. 

______. (2003). Dipterocarpaceae in Thailand: 

Taxonomic and Biogeographical Analysis. 

Thesis, Doctor of Philosophy, Kasetsart 

University. 

Smitinand, T., T. Santisuk & C. Phengklai. (1980). 

The Manual of the Dipterocarpaceae of 

Mainland South East Asia. Thai For. Bull. 

(Bot.) 12: 1–133. 

Symington, C.F. (1943). Forester’s Manual of 

Dipterocarps. Malayan Forest Records No.16, 

2nd ed (2004): 1–519.



Figure 16. Species of Anisoptera and Dipterocarpus in the Hala-Bala forest complex: leaves and fruits. 

91

92 


Figure 17. Species of Dipterocarpus and Vatica in the Hala-Bala forest complex: leaves and fruits.



Figure 18. Species of Parashorea, Neobalanocarpus and Hopea in the Hala-Bala forest complex: leaves and fruits. 

sp. 

93

94 


Figure 19. Species of Shorea in the Hala-Bala forest complex: leaves and fruits.



Figure 20. Species of Shorea in the Hala-Bala forest complex: leaves and fruits. 

95

96 


A B 

C D 

Figure 21. Anisoptera laevis Ridl.: A. infructescence; Dipterocarpus acutangulus Vesque: B. fruit; D. crinitus Dyer: C. stipular shoot 

and leaves, D. fruit.



A B 

C D 

Figure 22. Vatica bella Slooten: A. fruit; V. nitens King: B. infructescences; V. stapfi ana (King) Slooten: C. fruit; V. umbonata (Hook.f.) 

Burck: D. fruit. 

97

98 


A B 

C D 

Figure 23. Vatica maingayi Dyer: A. infl orescences, B. infructescences; V. lowii King: C. fruit; V. cuspidata (Ridl.) Symington: D. 

fruits.



A B 

C D 

Figure 24. Hopea sp.: A. infructescences; H. sublanceolata Symington: B. infructescences; H. dryobalanoides Miq.: C. leaves & 

fruits; H. montana Symington: D. leaf & fruits. 

99

100 


A B 

C D 

Figure 25. Hopea bracteata Burck: A. fruit; Shorea paucifl ora King: B. fruit; S. ochrophloia Strugnell ex Symington: C. infl orescence, 

D. infructescence.



A B 

C D 

Figure 26. Shorea parvifl ora ssp. velutinata P.S. Ashton: A. infructescences; S. ovata Dyer ex Brandis: B. fruits; S. longisperma 

Roxb.: C. infructescence; S. bracteolata Dyer: D. infructescence. 

101


A new species of Rhachidosorus (Rhachidosoraceae), a genus new to Thailand 

STUART LINDSAY*, DAVID J. MIDDLETON* & PIYAKASET SUKSATHAN** 

ABSTRACT. The genus Rhachidosorus is newly recorded for Thailand and the species Rhachidosorus siamensis S.Linds. is 

described. 

KEY WORDS: Rhachidosorus, Rhachidosoraceae, fern, Thailand. 

Rhachidosorus is a genus of about seven 

species from East and Southeast Asia, from Japan 

to Sumatra and the Philippines (Christenhusz et al., 

2011). The genus has been variously placed in 

Athyriaceae (e.g., Chu et al., 1999), Woodsiaceae 

(e.g., Smith et al., 2006, 2008) and, most recently, 

in Rhachidosoraceae (Christenhusz et al., 2011; 

Rothfels et al., 2012). On a joint expedition of the 

Royal Botanic Garden Edinburgh and the Queen 

Sirikit Botanic Garden in 2009 a fern was collected 

in Doi Pha Hom Pok National Park in Chiang Mai 

Province which proved not only to be a new record 

of the genus Rhachidosorus for Thailand but a new 

species in the genus. 

Descriptions of the genus and the new species 

are given below. 

RHACHIDOSORUS 

Ching, Acta Phytotax. Sin. 9: 73. 1964; Chu et al., 

Fl. Reipubl. Popularis Sin. 3(2): 267. 1999. 

Terrestrial or lithophytic, medium-sized to 

large ferns. Rhizome creeping, ascending or erect, 

scaly; scales brown, clathrate or only weakly so, 

sometimes of quite disparate shapes and sizes but 

generally narrowly triangular to linear, margin 

entire, basally attached. Frond d with distinct stipe 

and lamina. Stipes not thickened at base, not winged. 

Laminae bipinnate to tripinnate or quadripinnatifi 

d, rarely pinnate; pinnae usually not opposite, 

anadromic. Veins free, forked or pinnate. Sori 

oblong, narrowly elliptic or elongate, mostly on 

acroscopic veins in ultimate segments; indusia 

membranous, oblong or elongate, attached on one 

side along veins. Spore surface muriform or with 

warty projections. 

About seven species from East and Southeast 

Asia, from Japan to Sumatra and the Philippines. 

Rhachidosorus siamensis S.Linds., sp. nov. Similar 

to Rhachidosorus blotianus Ching but differing in 

darker stipe and rachis, lamina much less divided 

and sori more remote from costule/costulet. Typus: 

Thailand, Chiang Mai, Fang District, Doi Pha Hom 

Pok National Park, trail from Kew Lom Ranger 

Substation to Pang Mong Kon, 1750 m altitude, 

20º3’56’’ N, 99º8’6’’ E, in shady gully of small 

stream in lower montane forest on granite bedrock, 

2 October 2009, D.J. J Middleton, S. Lindsay & P. 

Suksathan 5051 (holotype QBG; isotypes BKF, E, 

P). Fig. 1. 

Terrestrial. Rhizome ascending, 2–3 cm diameter 

(when fresh), producing a mass of fi brous 

roots; scales on rhizome and stipe very variable in 

shape and size, brown or golden brown, weakly 

clathrate with elongate cells, narrowly triangular to 

linear, often rather crisped, 2–14 x 0.1–2.3 mm. 

Fronds to 113 cm long. Stipe and rachis mid to 

dark green above, very dark green to black beneath 

* Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, Scotland, U.K. Email: s.lindsay@rbge.ac.uk 

** Herbarium, Queen Sirikit Botanic Garden, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand.

A NEW SPECIES OF RHACHIDOSORUS (RHACHIDOSORACEAE), A GENUS NEW TO THAILAND (S. LINDSAY , D.J. MIDDLETON1 & P. SUKSATHAN) 103 

(distinction less obvious when dry), stipe 20–36 

cm long, densely scaly at base, becoming sparser 

towards rachis and into rachis. Lamina mid to dark 

green and slightly shiny above, much paler beneath, 

47–78 x 18–36 cm, bipinnate to tripinnate, ovate in 

outline, apex acuminate, formed from progressively 

smaller and closer pinnae without a distinct terminal 

pinna; rachis ± terete but with a narrow wing on 

either side visible from above, each with a thickened 

margin, with linear scales and short glandular 

hairs; 18–22 pairs of free pinnae, 3.5–12 cm apart 

near base, triangular, apex acuminate, largest pinnae 

towards base but not basal, 9.8–20 x 2.9–8.7 cm, 

stalked for 4–8 mm; axes winged, wings with 

thickened margins which are sometimes also slightly 

inrolled, lowest pinnae refl exed or not; ultimate 

segments ovate to elliptic, dentate, apex obtuse to 

acute, glabrous above, with many scattered short 

glandular hairs on veins and lamina beneath; venation 

clearly visible, veinlets not reaching margin 

and with enlarged endings visible above. Sori 

oblong or narrowly elliptic, mostly on acroscopic 

veins, one per pinnule lobe, basal lobes often larger 

and then with sori on both acroscopic and basiscopic 

veins and opening towards each other, 2–4 mm 

long, more or less equidistant from costule/costulet 

(depending on whether frond is bipinnate or tripinnate) 

and segment margin; indusia pale green, often 

translucent, margin slightly darker and minutely 

irregular to fi mbriate. Spore surface muriform. 

Distribution.— Only known from the type 

locality. 

Ecology.— Terrestrial on clay bank in shady 

gully of small stream in lower montane forest on 

granite bedrock at 1750 m. 

Note.— In the key to fern families in the 

Flora of Thailand (Tagawa & Iwatsuki, 1979) 

Rhachidosorus siamensis would key out to 

Athyriaceae, a family in which the genus was 

previously included. See Rothfels et. al (2012) for 

the differences that are now recognised between 

Rhachidosoraceae, Athyriaceae and Woodsiaceae. 


We thank the Leverhulme Trust and Queen 

Sirikit Botanic Garden for the funding for this 

work; the director and staff of Doi Pha Hom Pok 

National Park for their logistical support in the 

fi eld; and Dr Harald Schneider for helpful 

discussions. 

REFERENCES 

Christenhusz, M.J.M., Zhang, X-C. & Schneider, 

H. (2011). A linear sequence of extant families 

and genera of lycophytes and ferns. Phytotaxa 

19: 7–54. 

Chu Wei-ming, Wang Zhong-ren, Hsieh Yin-tang 

& He Zhao-rong (1999). Athyriaceae. In: Chu 

Wei-ming, ed., Fl. Reipubl. Popularis Sin. 3(2): 

32–504. Science Press, Beijing 

Rothfels, C.J., Sundue, M.A., Kuo, L.-Y., Larsson, 

A., Kato, M., Schuettpelz, E. & Pryer, K.M. 

(2012). A revised family-level classifi cation 

for eupolypod II ferns (Polypodiidae: 

Polypodiales). Taxon 61: 515–533. 


P., Schneider, H. & Wolf, P.G. (2006). A classifi 

cation of extant ferns. Taxon 55: 705–731. 


P., Schneider, H. & Wolf, P.G. (2008). Fern 

Classifi cation. Pp. 417–467 In: Ranker, T.A. & 

Haufl er, C.H. (eds), Biology and Evolution of 

Ferns and Lycophytes, CUP, Cambridge. 

Tagawa, M. & Iwatsuki, K. (1979). In: Smitinand, 

T. & Larsen, K. (eds), Flora of Thailand 3 (1). 

Royal Forest Department, Bangkok.

104 

E 


C 

A B 

Figure 1. Rhachidosorus siamensis S.Linds. A. Habit; B. Whole frond from above; C. Rachis and pinna above; D. The same rachis 

and pinna beneath; E. Pinnule with sori; F. Scales on crozier. All photos from Middleton et al. 5051. A by Piyakaset Suksathan, B–F 

by David J. Middleton. 

D 

F


Orchipedum Breda (Orchidaceae, subfam. Orchidoideae), a new generic record for Thailand 

INTRODUCTION 

The genus Orchipedum Breda belongs to the 

subtribe Goodyerinae, tribe Cranichideae in the 

subfamily Orchidoideae (Pridgeon et al., 2003), 

previously placed in other subfamilies, e.g. 

Neottioideae Lindl. (Seidenfaden & Wood, 1992) 

and Spiranthoideae Dressler (Dressler, 1993). The 

genus contains only three species including two 

Malesian species, O. plantaginifolium Breda (Breda, 

1827–1829; Seidenfaden & Wood, 1992; Comber, 

2001) and O. wenzelii i (Ames) J.J. Sm. (Smith, 1934), 

and one species from South Vietnam, O. echinatum 

Aver. & Averyanova., described by Averyanov & 

Averyanova (2006). Although the subfamily 

Orchidoideae has recently been treated in the Flora 

of Thailand, including 4 tribes, 7 subtribes, 30 genera 

and a total of 146 species (Pedersen et al., 2011), 

the genus Orchipedum has not been recorded from 

Thailand until now. During a fl oristic inventory of 

native orchids in the southernmost part of Peninsular 

Thailand by the author, O. plantaginifolium, previously 

known only from Peninsular Malaysia, 

Sumatra and Java, was discovered. 

ORCHIPEDUM 

Breda, Gen. Sp. Orchid. Asclep. 2: t. 10. 1827 

(1829); Seidenf. & J.J. Wood, Orchids Pen. Mal. & 

Sing.: 79. 1992; J.B.Comber, Orchids of Sumatra: 

71. 2001. Type species: O. plantaginifolium Breda 

JAREARNSAK SAE WAI* 

ABSTRACT. The genus Orchipedum is recognized as a new generic record for Thailand with one species, Orchipedum plantaginifolium 

Breda. The genus and species are described and illustrated. 

KEY WORDS: Orchipedum plantaginifolium, Orchidaceae, new generic record, Thailand. 

Terrestrial, perennial herb; rhizome creeping, 

succulent, rooting at the nodes; erect stem severalleaved, 

with a terminal infl orescence. Leaves spirally 

arranged, scattered along the stem, persistent, glabrous, 

convolute, green; petiole grooved, basally 

expanded into an amplexicaul sheath; lamina usually 

obliquely elliptic-ovate or elliptic-lanceolate. 

Infl orescences erect, racemose, many-fl owered; 

peduncle with a few sheathing scale-leaves. Flowers 

sessile, resupinate. Sepals free, hairy; dorsal sepal 

connivent with petals forming a hood; lateral sepals 

obliquely spreading, base concave, enclosing the 

labellar spur. Petals thin, membranous. Labellum 

3-lobed, adnate to column base, spurred; spur short, 

subglobose, inner surface with a group of fl eshy 

dendritic appendages on each side; base of labellum 

with 2 erect lateral lobes and 2 parallel keels; midlobe 

with a short claw, anchoriform. Column arcuate; 

anther 2-loculate; pollinia 2, sectile; caudicle and 

viscidium present; stigma lobes connate. Ovary 

hairy. Capsule and Seeds not seen. 

Three species distributed in southern Vietnam, 

Peninsular Malaysia, Sumatra, Java, and the 

Philippines. One species in Thailand. The description 

below is based on Thai material. 

This genus can be easily distinguished from 

other putatively related genera in the subtribe 

Goodyerinae (e.g. Herpysma Lindl. and Hylophila 

Lindl.) by having a short, pouch-like labellar spur 

* Herbarium (PSU), Princess Maha Chakri Sirindhorn National History Museum & Centre for Biodiversity of Peninsular Thailand 

(CBIPT), Department of Biology, Faculty of Science, Prince of Songkla University, Songkhla 90112, Thailand. Email: Jareansak.s@ 

psu.ac.th

106 

containing a group of fl eshy dendritic appendages 

on each side. 

Orchipedum plantaginifolium Breda, Gen. Sp. 

Orchid. Asclep. 2: t. 5. 1827 (1829); Seidenf. & 

J.J.Wood, Orchids Pen. Mal. & Sing.: 79, fi g. 29. 

1992; J.B. Comber, Orchids of Sumatra: 71. 

2001.— Queteletia plantaginifolia (Breda) Blume, 

Coll. Orchid.: 117. 1859; Holttum, Rev. Fl. Malaya 

1: 130. 1953. Type: Indonesia, Java, Kuhl 2012 

(holotype L). Fig. 1. 

Erect part including infl orescence to 1 m 

high; rhizome creeping, succulent, green, glabrous, 

8–12 mm in diam.; each node usually with 4–5 

roots; internodes 2.5–6.5 cm long. Leaves 6–12, 

glabrous; petiole (including sheath) 3–5 cm long, 

amplexicaul sheath ca 1 cm long; lamina obliquely 


elliptic-ovate to elliptic-lanceolate, 6–15 by 2–7 

cm, apex acuminate, base cuneate, margin straight 

or slightly undulate, midrib impressed above and 

prominent underneath. Infl orescence dense, 10- to 

22-fl owered; peduncle 8–9 cm long, glandularhairy, 

lower part almost glabrous; scale-leaves 

sheathing, lanceolate, 2.5–3.3 cm long, apex acuminate, 

sparsely glandular-hairy or subglabrous; 

rachis 4–10 cm long, glandular-hairy; fl oral bracts 

lanceolate, greenish turning pale reddish brown, 

15–25 by 5–7 mm, apex acuminate, glandularhairy. 

Flowers 12–15 mm in diam. Sepals greenish, 

5-veined, outer surfaces glandular-hairy; dorsal 

sepal connivent with petals forming a hood, dorsal 

sepal strongly concave, narrowly elliptic-ovate, 

10–11 by 4–5 mm; lateral sepal spreading, obliquely 

ovate, 10–11 by 5.5–6 mm. Petals thin, membranous, 

obliquely oblanceolate to oblong-oblanceolate, 

Figure 1. Orchipedum plantaginifolium Breda: A–B. Habit and habitat; C. Infl orescence; D. Flower. Photographed by J. Wai.

ORCHIPEDUM BREDA (ORCHIDACEAE, SUBFAM. ORCHIDOIDEAE), A NEW GENERIC RECORD FOR THAILAND (J. WAI) 107 

10–11 by 3.5–4 mm, 1-veined, outer surfaces 

sparsely glandular-hairy or subglabrous. Labellum 

3-lobed, with a median longitudinal groove, 10–11 

mm long, white; spur subglobose, ca 3 mm in 

diam., inner surface with a group of fl eshy dendritic 

appendages on each side; lateral lobes erect, semicircular, 

3.5–4 mm high, 5–6 mm long, forward 

edges thickened, with many small lobulated warts 

on the underside; keels 2, erect, semi-circular, extending 

in parallel from the base to near the middle 

of labellum; midlobe bilobed, with a short claw ca 

1.5 mm long , anchoriform, 3–4 by 5–5.5 mm, 

apex obtuse to subacute; lobules slightly erose. 

Column arcuate, with 2 bidentate rostellar arms, 

8–9 mm long; anther narrowly ovoid, ca 6 mm 

long; pollinia 2, sectile, narrow, clavate; stigma 

ovate, ca 1 mm long. Ovary 8–10 mm long, glandular 

hairy. Capsule and Seeds not seen. 

Thailand.— PENINSULAR: Yala [Betong district, 

Khlong Mu Bo, 3 April 2005, J. J Wai 187 7 ( PSU); 

same loc., 12 April 2010, J. J Wai 1887 7 ( PSU)]. 


Java. 

Ecology.— In deep shade along streamside 

areas in primary lowland evergreen forest at 500– 

600 m alt. Flowering in March to June. 

Conservation status.— Its provisional status 

is considered to be Least Concern (LC) according 

to IUCN Red List Criteria Version 3.1 (IUCN, 

2001). It is not under immediate threat, because of 

its wide geographical distribution and existence in 

well-protected habitats. 

Notes.— Although the few specimens collected 

from Thailand, this species seems to be 

locally fairly common. During fi eld observations 

from 2005–2012, a lot of sterile plants were found 

in Betong, especially in the lowland forest of Bang 

Lang National Park. 


The author would like to thank Assoc. Prof. 

Dr. Kitichate Sridith for his advice and two 

anonymous referees for their valuable comments 

on my manuscript and Mr. Nopadol Sae Wai for his 

kind support in the fi eld. This work was supported 

by the Higher Education Research Promotion and 

National Research University Project of Thailand, 

Offi ce of the Higher Education Commission. 

REFERENCES 

Averyanov, L.V. & Averyanova, A.L. (2006). New 

orchids from Vietnam. Komarovia 4: 1–39. 

Breda, J.G.S. Van. (1827–1829). Genera et species 

Orchidearum et Asclepiadarum quas in itinerere 

per insulam Java collegerunt Dr. H. Kuhl et Dr. 

J.C. van Hasselt. Ghent. Folio. 15 fol. & 15 tab. 

col. 

Comber, J.B. (2001). Orchids of Sumatra. Natural 

History Publications (Borneo) in association 

with the Royal Botanic Gardens, Kew and 

Singapore Botanic Gardens, Singapore. 

Dressler, R.L. (1993). Phylogeny and Classification 

of the Orchid Family. Dioscorides Press, 

Portland, Oregon. 


Criteria: Version 3.1. IUCN Species Survival 



Pedersen, H.Æ., Kurzweil, H., Suddee, S. & Cribb, 

P.J. (2011). Orchidaceae 1 (Cypripedioideae, 

Orchidoideae,Vanilioideae). In T. Santisuk & 

K. Larsen (eds.), Flora of Thailand. Vol. 12(1), 

pp. 1-302. Prachachon Co., Ltd., Bangkok. 

Pridgeon, A.M., Cribb, P.J., Chase, M.W. & 

Rasmussen, F.N. (eds.). (2003). Genera 

Orchidacearum vol. 3: Orchidoideae part 2, 

Vanilloideae. Oxford University Press, Oxford. 

Seidenfaden, G. & Wood, J.J. (1992). The Orchids 

of Peninsular Malaysia and Singapore. Olsen 

& Olsen, Fredensborg. 

Smith, J.J. (1934). Artificial key to the Orchid genera 

of the Netherlands Indies, together with 

those of New Guinea, the Malay Peninsula and the 

Philippines. Blumea 1: 195–215.


INTRODUCTION 

The remarkably rich orchid fl ora of Myanmar 

is one of the most poorly known in Asia, and is currently 

the subject of ongoing inventory work 

(Kress, 2003; Ormerod & Kumar, 2003, 2008; 

Ormerod & Wood, 2010; Tanaka et al., 2011; 

Kurzweil & Lwin, 2012). Comprehensive studies 

by the authors of this paper, by Paul Ormerod, 

Sathish Kumar and Jin Xiaohua are aimed at producing 

a complete checklist of all species recorded 

in the country so far. During two fi eldtrips in the 

northernmost part of Kachin State in 2007 and 

2009 we collected material of fi ve species of 

Orchidaceae which are not known in Myanmar as 

yet, and these new distribution records are presented 

below. 

1. Calanthe alismifolia Lindl., Fol. Orchid. 6: 8. 

1855; Hook.f., Fl. Brit. India 5: 849. 1890; Seidenf., 

Opera Bot. 114: 94, fi g. 47. 1992; N. Pearce & P.J. 

Cribb, Fl. Bhutan 3(3): 283. 2002; S.C. Chen et al. 

in Z.Y. Wu et al. (eds), Fl. China 25: 302. 2009 [as 

alismatifolia]; Z.Y. Wu et al. (eds), Fl. China 

Illustrations 25: 431.1–2, 432.1–17. 2010.— 

Alismorkis alismifolia (Lindl.) Kuntze, Revis. Gen. 

Pl. 2: 650. 1891.— Type: India, Sikkim, Hooker 

239A (syntype K-LINDL); India, Khasia, Hooker 

& Thomson 239B (syntype K-LINDL). [fi de 

Pearce & Cribb, 2002]. 

New records in the orchid fl ora of Myanmar 

HUBERT KURZWEIL* & SAW LWIN** 

ABSTRACT. Five orchid species, Calanthe alismifolia Lindl., Ceratostylis radiata J.J. Sm., C. subulata Blume, Cleisostoma linearilobatum 

(Seidenf. & Smitinand) Garay and Panisea tricallosa Rolfe are newly recorded for Myanmar. While the occurrence of 

Ceratostylis radiata is a remarkable extension of the geographical range of this species, the remaining four species were previously 

known to occur in most of the surrounding countries and their discovery in Myanmar is therefore not surprising. 

KEY WORDS: Myanmar, new records, Orchidaceae. 

Distribution.— Eastern Himalayas to Vietnam, 

China and southern Japan. 

Ecology.— Our collection was made in undisturbed 

mountain forest at about 1115 m. In the 

eastern Himalayas the species is found among 

grass at the base of rocks between 660 and 1600 m 

(Pearce & Cribb, 2002), and in China in evergreen 

broad-leaved forest between 700 and 2100 m 

(Chen & al., 2009). 

Myanmar.— Kachin State: Mountains west 

of Putao, near the village of Ziyadan, undisturbed 

mountain forest, 1115 m, 23 March 2009, Kurzweil 

& Lwin 2693 (SING [spirit]). 

Notes.— A widespread species with white 

fl owers and typically three-lobed lip. A characteristic 

feature are the rather wide ovate-elliptic leaves 

to which the species epithet alludes. Due to its general 

distribution the occurrence in Myanmar is not 

surprising. 

2. Ceratostylis radiata J.J. Sm., Orch. Java: 295. 

1905; Seidenf., Opera Bot. 89: 115, fi g. 66, pl. 9c. 

1986; Seidenf. & J.J. Wood, Orchid. Penins. 

Malaysia Singap.: 311, fi g. 137c–e, pl. 19c. 1992; 

J.J. Wood & P.J. Cribb, Checklist Orch. Borneo: 

205. 1994. Type: Java, Soekaboemi, Garoet and 

south Preangen, Raciborski s.n. (holotype BO). [fi de 

Wood & Cribb, 1994]. Fig. 1A–B.— Ceratostylis 

* Singapore Botanic Gardens, 1 Cluny Road, 259569 Singapore. Email: HUBERT_KURZWEIL@nparks.gov.sg 

** Myanmar Floriculturist Association, Ahlone Road, Yangon, Myanmar.

NEW RECORDS IN THE ORCHID FLORA OF MYANMAR (H. KURZWEIL & S. LWIN) 109 

linearifolia Ridl., Fl. Malay Penins. 4: 110. 1924. 

Type: Malaysia, Kedah, Langkawi, Gunung Raja, 

13 Nov. 1921, Haniff 7103 (holotype SING!). 

Distribution.— Vietnam and Peninsular 

Thailand to West Malesia. 

Ecology.— Our collection was made in secondary 

mountain forest. The few published habitat 

records in other parts of the distribution area indicate 

that in Java Ceratostylis radiata grows as a 

trunk epiphyte in shade at altitudes of between 750 

and 1070 m (Comber, 1990). Populations in Borneo 

are found in lower montane forest, podsol forest 

and oak-laurel forest at altitudes of between 1100 

and 1900 m (Wood & Cribb, 1994). 


of Putao, between the villages of Wasadan and 

Ziyadan, secondary mountain forest, 20 March 

2009, Kurzweil & Lwin 2609 (SING, SING 

[spirit]). 

Notes.— This species is unmistakable with 

its large white stellate fl owers with densely hairy 

sepal dorsal surfaces and a characteristic lip shape. 

The reticulate leaf sheaths make the species recognisable 

in the vegetative state. It appears that an 

immature specimen was illustrated by Seidenfaden 

(1986: fi g. 66 on page 114) as it shows a specimen 

with connate lateral sepals, i.e. it was drawn before 

the lateral sepals had opened up. In contrast, the 

colour photo in the same publication (plate 9c) 

shows fully open fl owers. 

Ceratostylis radiata was previously only 

known to occur in Vietnam, Peninsular Thailand, 

Peninsular Malaysia, Sumatra, Borneo and Java. 

The nearest occurrence to the Myanmar locality is 

in Peninsular Thailand (in the provinces of 

Phangnga and Nakhon Si Thammarat) which is 

about 2000 km away, and this is a surprising range 

extension. 

3. Ceratostylis subulata Blume, Bijdr.: 306. 1825; 

Seidenf., Opera Bot. 89: 109, fi g. 62. 1986; 

Seidenf., Opera Bot. 114: 161. 1992; Seidenf. & 

J.J. Wood, Orchid. Penins. Malaysia Singap.: 310, 

fi g. 136a–d, pl. 29b. 1992; J.J. Wood & P.J. Cribb, 

Checklist Orch. Borneo: 205. 1994; N. Pearce & 

P.J. Cribb, Fl. Bhutan 3(3): 363. 2002; S.C. Chen & 

J.J. Wood in Z.Y. Wu et al. (eds), Fl. China 25: 361. 

2009; Z.Y. Wu et al. (eds), Fl. China Illustrations 

25: 485.4–6. 2010. Type: Java, Salak, Blume s.n. 

(syntype, herbarium unknown); Pantjar, Blume s.n. 

(syntype, herbarium unknown). Fig. 1C.— 

Ceratostylis teres (Griff.) Rchb.f., Bonplandia 

(Hannover) 2: 89. 1854; Hook.f., Fl. Brit. India 5: 

825. 1890.— Appendicula teres Griff., Not. Pl. 

Asiat. 3: 35. 1851. Type: India, Upper Assam, 

Negrigam, Griffi th s.n. (holotype K-LINDL) [fi de 

Pearce & Cribb, 2002].— Ceratostylis cepula 

Rchb.f., Bonplandia (Hannover) 5: 53. 1857. Type: 

Java, Prov. Bandong, Zollinger 3185 (holotype, 

herbarium unknown).— Ceratostylis malaccensis 

Hook.f., Fl. Brit. India 5: 825. 1890. Type: Perak, 

Scortechini s.n. (syntype K); Batang Padung, 4900 

ft., Wray s.n. (syntype K); Johor, Mount Ophir, 

Griffi th s.n. (Kew Distr. 5213) (syntype K). 

Distribution.— North-eastern India and eastern 

Himalaya to southern China, and throughout 

the whole of Malesia as far east as Vanuatu. 

Ecology.— Our specimen was epiphytic in 

secondary forest. In China the plants are epiphytic 

on trees or lithophytic on rocks in forest (Chen & 

al., 2009). In Peninsular Malaysia the species is a 

common epiphyte in montane as well as lowland 

forest (Seidenfaden & Wood, 1992). Plants in 

Borneo grow in lowland forest, hill dipterocarp 

forest and lower montane oak-laurel forest at altitudes 

of between 400 and 1700 m (Wood & Cribb, 

1994). According to Comber (1990) Javanese 

plants grow in tufts on high branches and are therefore 

usually only seen on fallen trees, and this may 

be the case in other parts of the distribution area as 

well. 

Myanmar.— Kachin State: Mountains northeast 

of Putao, epiphytic in secondary forest, 560 m, 

4 March 2007, Kurzweil & Lwin 2379 (SING 

[spirit]). 

Notes.— The species, in its current delimitation, 

is very widespread in tropical and subtropical 

Asia. Thus its discovery in Myanmar is quite natural. 

In the past, this collection would have been referred 

to Ceratostylis teres (Griff.) Rchb.f. which 

was described from north-eastern India, but is now 

regarded as a synonym (Ormerod, pers. comm.). 

Most recent authors consider C. subulata and C. 

teres as conspecifi c, although Seidenfaden (1986, 

1992) raised doubts about their conspecifi ty.

110 


Figure 1. A–B. Ceratostylis radiata J.J. Sm.: A. Flowers; B. Habit. Kurzweil & Lwin 2609. Note the reticulate leaf sheaths; 

C. C. subulata Blume, habit. Kurzweil & Lwin 2379.


The Myanmar specimen cited corresponds in 

fl ower, stem and leaf structure as well as in the 

fl ower colour well with Ceratostylis subulata 

(Seidenfaden, 1986). However, the column-foot is 

unusually long, measuring nearly 2 mm. 

4. Cleisostoma linearilobatum (Seidenf. & 

Smitinand) Garay, Bot. Mus. Leafl . 23: 172. 1972; 

Seidenf., Opera Bot. 124: 60. 1995; N. Pearce & 

P.J. Cribb, Fl. Bhutan 3(3): 509, fi g. 113, pl. 27. 

2002; S.C. Chen & J.J. Wood in Z.Y. Wu et al. 

(eds), Fl. China 25: 459. 2009; Z.Y. Wu et al. (eds), 

Fl. China Illustrations 25: 606.5–7. 2010.— 

Sarcanthus linearilobatus Seidenf. & Smitinand, 

Orch. Thail. (Prelim. List): 684. 1965.— Ormerodia 

linearilobata (Seidenf. & Smitinand) Szlach., Ann. 

Bot. Fenn. 40: 68. 2003.— Type: Thailand, Chiang 

Mai, Kawng He, 960 m, Kerr 363 (holotype K). 

Fig. 2A. 

Cleisostoma sagittiforme Garay, Bot. Mus. 

Leafl . 23: 174. 1972; Seidenf., Dansk Bot. Ark. 

29(3): 19, fi g. 5. 1975.— Sarcanthus sagittatus 

King & Pantl., J. Asiat. Soc. Bengal, Pt. 2, Nat. 

Hist. 66: 595. 1897.— Ormerodia sagittata (King 

& Pantl.) Szlach., Ann. Bot. Fenn. 40: 68. 2003.— 

Type: India, Khasia, Pantling 629 (holotype CAL). 

Cleisostoma sikkimense Lucksom, Indian J. 

Forest. 15: 27. 1992.— Type: India, Sikkim, 

Leeving, Lucksom 204A (holotype BHSC) [fi de 

Pearce & Cribb, 2002]. 

Distribution.— India (Sikkim) to south-western 

China and Peninsular Malaysia. 

Ecology.— Our plants grew in evergreen 

mountain forest, which also corresponds well with 

populations in Yunnan (Chen & al., 2009). Eastern 

Himalayan plants grow in shade on Castanopsis 

tribuloides at altitudes of between 500 and 2000 m 

(Pearce & Cribb, 2002). 


of Putao, undisturbed evergreen forest, 22 March 

2009, Kurzweil & Lwin 2684 (SING [spirit]); 

mountains west of Putao, 1000 m, 25 March 2009, 

Kurzweil & Lwin 2772 (SING [spirit]). 

Notes.— Among the species with fl at and 

apically bilobed leaves and fl owers with a linear 

stipe, Cleisostoma linearilobatum is well characterised 

having fl owers with a longitudinal spur septum 

and a truncate operculum front edge. 

Characteristic features of the species are also the 

two prominent rostellum lobes and the outwardbulging 

distal parts of the lip side lobes. The species 

is rather widespread in continental Asia, and 

its discovery in Myanmar falls into a gap in its 

known distribution. 

5. Panisea tricallosa Rolfe, Bull. Misc. Inform. 

Kew 1901: 148. 1901; Seidenf., Opera Bot. 89: 86, 

fi g. 46, pl. 6c. 1986; Lund, Nord. J. Bot. 7: 520, 

fi gs. 10–17. 1987; Seidenf., Opera Bot. 114: 196. 

1992; N. Pearce & P.J. Cribb, Fl. Bhutan 3(3): 347. 

2002; S.C. Chen & J.J. Wood in Z.Y. Wu et al. 

(eds), Fl. China 25: 333. 2009.— Sigmatogyne tricallosa 

(Rolfe) Pfi tzer in H.G.A. Engler (ed.), 

Pfl anzenr., IV, 50(32): 133. 1907.— Type: India, 

Assam, cult. RBG Glasnevin (holotype K!). Fig. 

2B–C. 

Panisea pantlingii (Pfi tzer) Schltr., Orchideen: 

155. 1914.— Sigmatogyne pantlingii Pfi tzer in 

H.G.A. Engler (ed.), Pfl anzenr., IV, 50(32): 134. 

1907.— Type: India, Assam, Watt 571 (holotype K!). 

Sigmatogyne bia Kerr, J. Siam Soc., Nat. 

Hist. Suppl. 9: 236. 1933.— Panisea bia (Kerr) 

Tang & F.T. Wang, Acta Phytotax. Sin. 1: 79. 

1951.— Type: Laos, Phu Bia, 1500 m, 12 April 

1932, Kerr 0971 (holotype K). 

Panisea unifolia S.C. Chen, Acta Bot. 

Yunnan. 2: 304. 1980.— Type: China, Yunnan, 

Fong Chin, 2100 m, T. T T. T Yü 16321 (holotype PE). 

Panisea tricallosa var. garrettii I.D. Lund, 

Nordic J. Bot. 7: 524. 1987.— Type: Thailand, Doi 

Inthanon, 1550 m, Garrett 623 (holotype BKF). 

Distribution.— Nepal to Indochina and 

south-western China. 

Ecology.— Our specimens were growing in 

mountain forest. Eastern Himalayan plants were 

cited as growing on rocks in open oak forest at an 

altitude of 1700–2160 m (Pearce & Cribb, 2002). 

Myanmar.— Kachin State: Mountains northeast 

of Putao, in primary mountain forest, 6 March 

2007, Kurzweil & Lwin 2442 (SING [spirit]); 

mountains west of Putao, on fallen tree in a clearing 

in secondary forest, 1325 m, 19 March 2009, 

Kurzweil & Lwin 2573 (SING). 

Notes.— A widespread species in the subtropical 

parts of continental Asia, and its occurrence 

in Myanmar was therefore expected.

112 


Figure 2. A. Cleisostoma linearilobatum (Seidenf. & Smitinand) Garay, fl ower. Kurzweil & Lwin 2772; B–C. Panisea tricallosa 

Rolfe. Kurzweil & Lwin 2573.



Most of all our thanks go to Mr Paul Ormerod 

for his advice on the species reported here. We 

would also like to acknowledge the help of the 

Myanmar Forest Department for arranging our 

fi eldwork, and Dr Jin Xiaohua for checking the 

type of Panisea unifolia in the herbarium PE. The 

second author would also like to acknowledge the 

fi nancial assistance provided by a Singapore 

Botanic Gardens Research Fellowship. 

REFERENCES 

Chen Sing-Chi, Liu Zhongjian, Zhu Guanghua, 

Lang Kai-Yung, Tsi Zhan-Huo, Luo Yibo, Jin 

Xiaohua, Cribb, P.J., Wood J.J., Gale, S.W., 

Ormerod, P., Vermeulen, J.J., Wood, H.P., 

Clayton, D. & Bell, A. (2009). Orchidaceae. 

In: Wu Zhengyi, Raven, P.H. and Hong Deyuan 

(eds.), Flora of China, vol. 25, pp. 1–570. 

Science Press, Beijing & Missouri Botanical 

Garden Press, St. Louis. 

Comber, J.B. (1990). Orchids of Java. Bentham- 

Moxon Trust, Royal Botanic Gardens, Kew. 

Kress, W.J., DeFilipps, R., Farr, E. & Yin-Yin-Kyi. 

(2003). A checklist of the trees, shrubs, herbs 

and climbers of Myanmar. Contributions from 

the US National Herbarium 45: 1–590. 

Kurzweil, H. & Lwin, S. (2012). First record of 

Taeniophyllum (Orchidaceae) in Myanmar. 

Gardens’ Bulletin Singapore 64: 133–137. 

Ormerod, P. & Sathish Kumar, C. (2003). 

Orchidaceous additions to the Flora of Burma 

(Myanmar). Rheedea 13: 43–50. 

Ormerod, P. & Sathish Kumar, C. (2008). 

Orchidaceous additions to the Flora of 

Myanmar 2. Rheedea 18: 75–80. 

Ormerod, P. & Wood, E.W. (2010). A new species 

of Pinalia (Orchdaceae: Eriinae) from 

Myanmar. Harvard Papers in Botany 15(2): 

349–351. 

Pearce, N.R. & Cribb, P.J. (2002). The orchids of 

Bhutan. Royal Botanic Gardens Edinburgh & 

Royal Government of Bhutan. 

Seidenfaden, G. (1986). Orchid genera in Thailand 

XIII. Thirty-three epidendroid genera. Opera 

Botanica 89: 1–216. 

Seidenfaden, G. (1992). The orchids of Indochina. 

Opera Botanica 114: 1–502. 

Seidenfaden, G. & Wood, J.J. (1992). The Orchids 

of Peninsular Malaysia and Singapore. Olsen 

& Olsen, Fredensborg. 

Tanaka, N., Yukawa, T., Khin Myo Htwe, Koyama, 

T. & Murata, J. (2011). New or noteworthy 

plant collections from Myanmar (7): Fourteen 

additional species of Orchidaceae. Acta 

Phytotax. Geobot. 61(3): 161–165. 

Wood, J.J. & Cribb, P.J. (1994). A check-list of the 

orchids of Borneo. Royal Botanic Gardens, 

Kew.


A new species record of Sciaphila (Triuridaceae) for Thailand 

SAHUT CHANTANAORRAPINT & AMONRAT CHANTANAORRAPINT** 

ABSTRACT. Sciaphila tenella Blume, a myco-heterotrophic plant, is newly recorded for Thailand. A description and illustrations 

are provided. 

KEY WORDS: achlorophyllous, myco-heterotrophic, Sciaphila tenella, Thailand, Triuridaceae 

INTRODUCTION 

The genus Sciaphila Blume (including Andruris 

Schltr.) comprises 37 species (Govaerts et al., 2012) 

and is the largest genus of the family Triuridaceae, 

mainly distributed in tropical Asia and America 

with a few species in subtropical and temperate regions 

(Maas van de Kamer & Weustenfeld, 1998). 

The highest species diversity of the genus is in 

South-East Asia including the Malay Peninsula, 

Sumatra, Java and Borneo (Meerendonk, 1984; 

Mass & Rübsamen, 1986). Taxa of this genus are 

characterized by being small and delicate, achlorophyllous 

herbs, monoecious, with erect stems and 

scale-like leaves; infl orescences are terminal, with 

bisexual or unisexual fl owers; male fl owers possess 

2–6 stamens usually concentrated in the apical part 

of the infl orescence; female and bisexual fl owers 

have 10−80 ovaries mostly concentrated in the 

basal part of the infl orescence. 

DESCRIPTION 

Most Sciaphila species are small in size with 

highly reduced vegetative morphology and are not 

easily discerned in the fi eld. Five species were previously 

known in Thailand (Larsen, 1972; Triboun 

& Larsen, 1999; Maneenoon & Sirirugsa, 2002; 

Chantanaorrapint & Thaithong, 2004): S. arfakiana 

Becc., S. maculata Miers, S. nana Blume, S. secundifl 

ora Thwaites ex Benth. and S. thaidanica K. 

Larsen, although Govaerts et al. (2012) record just 

two. More new records of species are expected in 

the unexplored areas, especially in the lower peninsular 

part of the country. 

During the botanical surveys in the peninsular 

of Thailand, Sciaphila tenella Blume was discovered 

which is a new record for Thailand. The description 

and illustration below are based on the Thai 

specimens cited alone. 

Key to the species of Sciaphila in Thailand (modifi ed from Chantanaorrapint and Thaithong 2004). 

1. Flowers bisexual together with male fl owers. Apex of perianth lobes long-bearded 

2. Male and bisexual fl owers with 3 stamens 1. S. maculata 

2. Male and bisexual fl owers with 6 stamens 2. S. tenella 

1. Flowers unisexual. Apex of perianth lobes not as above 

3. Male perianth lobes with a claviform appendage at the apex. Style awl-shaped, smooth 

4. All perianth lobes with a claviform appendage 3. S. arfakiana 

4. Only the 3 smaller perianth lobes with a claviform appendage; the 3 larger without appendages 4. S. nana 

3. Male perianth lobes without a claviform appendage at the apex. Style clavate, papillose 

5. Stems ca. 5 cm tall, branched from the base. Infl orescences 1–1.5 cm long. Flowers 4–5 mm in diam. Male fl owers with 6 

perianth segments 5. S. thaidanica 

5. Stems ca. 15 cm tall, fi rst branching ca. 4.5 cm from the base. Infl orescences 5 cm long. Flowers 8–10 mm in diam. Male fl owers 

with 4–8 perianth segments 6. S. secundifl ora 

* PSU-Herbarium, Department of Biology, Faculty of Science Prince of Songkla University, Hat Yai, Songkhla, 90112, Thailand. 

Email: sahut.c@psu.ac.th 

** Faculty of Natural Resources, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand. Email: amonrat.b@psu.ac.th

A NEW SPECIES RECORD OF SCIAPHILA (TRIURIDACEAE) FOR THAILAND (S. CHANTANAORRAPINT & A. CHANTANAORRAPINT) 115 

Sciaphila tenella Blume, Bijdr. Fl. Ned. Ind. 10: 

515. 1825; Meerendonk, Fl. Males. Ser. I. 10: 117. 

1984; Weeras. in Dassan. & Clayton (eds.), Revised 

Handb. Fl. Ceylon 14: 287. 2000; H.Ohashi et al., 

J. Jap. Bot. 83: 32. 2008; Y.H.Guo & Cheek in Z.Y. 

Wu et al. (eds.), Fl. China 23: 125. 2010.— Type: 

Indonesia, Java, Unknown s.n. (L, digital image 

L0050321!). Figs. 1–2. 

Monoecious, myco-heterotrophic, achlorophyllous 

herbs, reddish purple. Stem delicate, erect, 

to 15 cm tall, simple or 1–2-branched, glabrous. 

Leaves scale-like, entire, appressed to stem, ovate 

to lanceolate, acute to acuminate, 1–3 by 0.7–2 

mm. Infl orescence terminal, racemose, erect, of 12 

to 25 fl owers with basal bisexual fl owers and male 

ones in apical part; pedicels 3–6 mm; bracts lanceolate, 

2–4 mm long. Bisexual fl owers ca. 2‒3 

mm in diameter; perianth with 6 unequal lobes alternating 

larger and smaller lobes; larger lobes 

1.1–2.0 by 0.3–0.7 mm; smaller lobes 0.8–1.6 by 

0.3–0.6 mm; all lobes completely refl exed when 

opened; apex of lobes acute, bearded; stamens 6, 

easily broken off, fi laments short, anthers 2–3 locules; 

ovaries 10–25(30), obovate, the upper half tuberculate; 

style short, lateral; stigma papillate. Male 

fl owers 1.5‒2 mm in diameter; similar to the bisexual 

ones, but smaller and ovaries not well developed or 

reduced. Fruits aggregate, suborbicular; fruitlets 

dehiscent, ovoid, 0.9–1.2 mm in diameter. 

Thailand.— PENINSULAR: Trang: [Chao Pa 

waterfall, 17 Jan 2012, S. Chantanaorrapint, J. 

Inuthai & C. Promma 801 (BCU, BKF, PSU, spirit); 

Satun [Tarutao Island, 6°37’26’’N 99°38’27.1’’E, 24 

May 2008, S. Chantanaorrapint 2029 9 ( PSU, spirit)]. 

Distribution.— Widely distributed in tropical 

and subtropical Asia: Sri Lanka, Peninsular 

Malaysia, Singapore, Sumatra, Java, Borneo, The 

Philippines, New Guinea, Pacifi c islands, China 

and Japan. 

Ecology.— The Thai specimens were found 

growing amongst other mycoheterotrophic plants 

such as Epirixanthes sp., Gymnosiphon aphyllus 

Blume, Sciaphila secundifl ora Thwaites ex Benth. 

and Thismia alba Holttum ex Jonker, in sandy soil 

covered by leaf litter over sandstone or limestone 

rocks, under shade in primary lowland evergreen forest, 

ca. 90 m altitude. Flowering in the rainy season. 

Notes.— Sciaphila tenella is readily distinguished 

from all other species of Sciaphila by the 

presence of hairs at the apex of the perianth lobes 

and the 6 stemens with 2–3 locules. This species is 

most similar to S. maculata in having hairs at the 

apex of the perianth lobes, however the stamen 

number differs. 


The authors would like to thank Assoc. Prof. 

Dr Obchant Thaithong (BCU) and Prof. Dr 

Kitichate Sridith (PSU) for their valuable comments 

on the fi rst draft of the manuscript. Sincerely 

thank also to the anonymous readers who reviewed 

this manuscript. Thanks also due to the staff of 

Tarutao National Park for their cooperation during 

fi eld surveys. This work was supported by the 

Faculty of Science, Prince of Songkla University. 

REFERENCES 

Chantanaorrapint, S. & Thaithong, O. (2004). 

Sciaphila nana Blume (Triuridaceae), a new 

record for Thailand. Thai Forest Bulletin 

(Botany). 32: 12‒14. 

Govaerts, R, Maas-van der Kamer, H. & Maas, P. 

(2012). World Checklist of Triuridaceae. 

Facilitated by the Royal Botanic Gardens, 

Kew. Published on the internet; http://apps. 

kew.org/wcsp/. Retrieved 2012-09-05. 

Larsen, K. (1972). Triuridaceae. In: T. Smitinand 

& K. Larsen (eds), Flora of Thailand Vol. 2(1), 

pp. 175–176. Applied Scientifi c Research 

Cooperation of Thailand, Bangkok. 

Maas, P. J. M. & Rübsamen, T. (1986). Triuridaceae. 

Flora Neotropica Monograph 40: 1–55. 

Maas van de Kamer, H. & Weustenfeld, T. (1998). 

Triuridaceae. In: Kubitzki, K. The Families and 

Genera of Vascular Plants. 3. Flowering Plants. 

Monocotyledons Lilianae (except Orchidaceae), 

pp. 452–458. Spinger. Berlin, Heidelberg, Germany. 

Maneenoon, K. & Sirirugsa, P. (2002). Two species 

of Sciaphila Blume (Triuridaceae), new record 

for Thailand. Thai Forest Bulletin (Botany) 30: 

39‒42. 

Meerendonk, J. M. P. van de. (1984). Triuridaceae. 

In: C. G. G. J. van Steenis (ed), Flora Malesiana 

Ser. I. Vol. 10(1), pp. 109–121. Martinus 

Nijhoff Publishers, Netherlands. 

Triboun, P. & Larsen, K. (1999). Sciaphila secundifl 

ora Thwaites ex Benth. (Triuridaceae), a 

new record for Thailand. Thai Forest Bulletin 

(Botany) 27: 47‒51.

116 


Figure 1. Sciaphila tenella Blume: A. Habit; B. Young bisexual fl ower; C.–D. Young fruits; E. Stamen; F. Carpel; G. Fruitlet. From 

S. Chantanaorrapint 2029. Drawn by S. Chantanaorrapint.

A NEW SPECIES RECORD OF SCIAPHILA (TRIURIDACEAE) FOR THAILAND (S. CHANTANAORRAPINT & A. CHANTANAORRAPINT) 117 

Figure 2. Sciaphila tenella Blume: A. Natural habitat; B. Upper part of infl orescence; C. Male fl ower with 6 stamens; D. Young 

fruits. Photographed by S. Chantanaorrapint.


Cephalantheropsis longipes (Orchidaceae), a New Record for Peninsular Thailand 

PAUL ORMEROD*, SAHUT CHANTANAORRAPINT** & HUBERT KURZWEIL*** 

ABSTRACT. Cephalantheropsis longipes (Hook.f.) Ormerod is newly recorded from Peninsular Thailand. Notes are supplied on the 

phytogeographic signifi cance of this discovery, variation of the species and its synonymy. 

KEY WORDS: Thailand, Cephalantheropsis longipes, new distribution record 

INTRODUCTION 

Cephalantheropsis Guillaumin is an orchid 

genus of about four Southeast Asian terrestrial, 

caulescent, Calanthe-like species. It differs from 

Calanthe e R.Br. in having a caulescent habit combined 

with axillary and pseudoterminal infl orescences, 

and fl owers with a spurless labellum that is free 

from the column. In 1998 the fi rst author reviewed 

the genus, accepting fi ve species (including one 

new taxon from Peninsular Malaysia). Since then 

Averyanov (2001) has reduced the type species 

Ceph. lateriscapa Guillaumin to a synonym of the 

widespread Ceph. longipes (Hook.f.) Ormerod, a 

view that is accepted here. 

A specimen found by the second author during 

fi eldwork in Peninsular Thailand was identifi ed as 

Cephalantheropsis longipes which was not known 

before to occur in the country. This new distribution 

record is presented below. 

DESCRIPTION 

Cephalantheropsis longipes (Hook.f.) Ormerod, 

Orch. Digest 62: 156. 1998.— Calanthe longipes 

Hook.f., Fl. Brit. Ind. 6: 195. 1890.— Alismorkis 

longipes (Hook.f.) Kuntze, Rev. Gen. Pl. 2: 650. 

1891.— Phaius longipes (Hook.f.) Holttum, Gard. 

Bull. Singapore 11: 286. 1947. Type: India, Sikkim, 

G. King s.n. (holotype CAL?).— P. mindorensis 

Ames, Philipp. J. Sci., Bot. 2: 324. 1907. Type: 

Philippines, Mindoro, Mt Halcon, 1310 m, 13 Nov. 

1906, E.D. Merrill 5612 (holotype AMES!).— 

Calanthe dolichopoda Fukuy., Bot. Mag. Tokyo 

49: 296. 1935. Type: Taiwan, Daitun, near Shinten, 

10 Nov. 1933, N. Fukuyama 4536 6 (holotype 

TAI?).— Cephalantheropsis lateriscapa Guillaumin, 

Bull. Mus. Natl. Hist. Nat., II, 32: 189. 1960.— 

Gastrorchis lateriscapa (Guillaumin) Averyanov, 

Prelim. List Vietnam Orch. 1: 204. 1988. Type: 

Vietnam, Lam Dong Province, Dalat, Manline, P. 

Tixier 12/59 (holotype P).— Calanthe gracilis 

Lindl. var. sumatrana J.J. Sm., Blumea 5: 704. 

1945. Type: Indonesia, Sumatra, Aceh Province, 

above Takengon, 1290 m, Aug. 1934, C.G.G.J. van 

Steenis 6041 (holotype BO; fl oral sketch L, copy K). K 

Terrestrial herb; stem terete, many-noded, 

laxly 5-leaved in upper half, 30–40 cm long, 0.4– 

0.8 cm thick; sheaths in basal half 4, tubular, upper 

sheath sometimes with a small foliose blade. 

Leaves elliptic-lanceolate, subacuminate, glabrous, 

7.5–15.0 by 2.5–4.5 cm; leaf sheaths 4–6 cm long. 

Infl orescences up to 3 per stem, pseudoterminal 

(this one longest) or axillary (these two shorter), 

laxly many-fl owered, to 15 cm long; fl oral bracts 

narrowly lanceolate, long-acuminate, caducous, 

* P.O. Box 8210, Cairns 4870, Queensland, Australia. Email: wsandave1@bigpond.com 

** Herbarium, Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand. Email: 

chantanaorrapint@gmail.com 

*** Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569. Email: Hubert_Kurzweil@nparks.gov.sg

CEPHALANTHEROPSIS LONGIPES (ORCHIDACEAE), A NEW RECORD FOR PENINSULAR THAILAND 

(P. ORMEROD, S. CHANTANAORRAPINT & H. KURZWEIL) 

3-veined, 1.5–1.7 by 0.5–0.7 cm. Flowers not 

widely opening, shortly and laxly pubescent externally, 

yellow or orange-yellow. Sepals: dorsal sepal 

oblong-lanceolate, acute, 5-veined, ca. 10 by 

3.5 mm; lateral sepals oblong-lanceolate, subacuminate, 

5-veined, ca. 10 by 3.75 mm. Petals oblong-lanceolate 

from a broadly clawed base, acute, 

3-veined with branched lateral veins, ca. 9 by 3.3 

mm. Labellum trilobed, medially with 2 lamellate 

keels extending from above the base of the lip onto 

the middle of the epichile, ca. 8.5 by 7.3 mm; hypochile 

fl abellate with irregularly erose side lobes, 

concave medially, ca. 6.2 by 7.3 mm; claw ca. 0.5 

mm long; epichile reniform, ca. 1 by 2 mm. Column 

short, stout, semiterete, shortly pubescent (especially 

dorsally), base tumid, ca. 3.4 mm long (including 

anther cap). Ovary pedicellate ovary narrowly 

clavate, densely shortly pubescent, 18–23 

mm long. [Description after the Thai specimen]. 

Thailand.— PENINSULAR: Nakhon Si 

Thammarat [Khao Nan National Park, Sanyen Mt, 

montane forest, 1200 m, 25 July 2010, S. 

Chantanaorrapint SC2228 (PSU).] 

Distribution.— India (type), Myanmar, China, 

Vietnam, Peninsular Malaysia, Indonesia (Sumatra), 

Taiwan, Philippines. 

Other specimens examined: 

India.— Sikkim [without locality, G. King 

s.n. (BM); 1876, ex Major Mant s.n., icon G. King 

s.n. (K); 1879, G. King s.n. (BM); T. Anderson s.n. 

(MEL); Darjeeling area, 1220 m, cult. RBG 

Glasnevin, ex R. Pantling s.n. (K); Rungbee, 1830 

m, Oct. 1893, R. Pantling 2 (AMES, BM, K)]; 

Assam [Mawryngkneng, 1220 m, 1 Oct. 1951, 

Chand 4934 (K)]. 

Myanmar.— Kachin [Kachin Hills, 1898, S. 

Mokim s.n. (BM, K); Nam Tamai Valley, 1220– 

1525 m, 7 Nov. 1937, F. Kingdon-Ward 13443 

(AMES, BM)]. 

China.— Yunnan [Mengtze, 1220 m, A. 

Henry 11819 (K)]; Guangxi [SE of Shangsze, Nam 

She Village, 15 Dec. 1934, W.T. Tsang 24792 

(AMES)]. 

Vietnam.— Kon Tum Province [Dak Gley 

District, valley of Dak Poko River, 6–7 km to S of 

Dak Gley (near Dak Tung Village), 9 Dec. 1995, 

L.V. Averyanov et al. VH 2404 (GH)]; Lam Dong 

119 

Province [Dalat, ravine to the S of Langbian Palace, 

14 Nov. 1924, F. Everard 1767 7 ( AMES); Langbian, 

1500 m, 8 Dec. 1903, W. Micholitz s.n. (K)]. 

Philippines.— Luzon, Laguna Province [C.B. 

Robinson BS 17115 (PNH, destroyed; AMES 

drawing); Mt Maquiling, 1000 m, 17 March 1906, 

E.D. Merrill 5131 (AMES)]; Bataan Province [Mt 

Cayapo, 1450 m, 16 Dec. 1947, G.E. Edano PNH 

3199 (AMES)]. Panay, Capiz Province [Mt 

Madiaas, M. Ramos & G.E. Edano BS 30673 

(AMES)]. 

Ecology.— In lower montane forest at 1200 m 

elevation. 

Notes.— 1. Averyanov (2001) united 

Cephalantheropsis lateriscapa with the earlier C. 

longipes, pointing out that the plant is a polymorphic 

taxon in regard to the shape of the labellum. 

At fi rst it seemed possible to recognise C. lateriscapa 

by its almost sessile (claw 0.5 mm long) 

labellum epichile but material of C. longipes varies 

in claw length (0.5–1.0 mm long). For example the 

Chinese collection Henry 11819 has a 0.5 mm long 

claw whilst a Philippine specimen (Merrill 5131) 

had an asymmetric labellum with the claw 0.5 mm 

wide on one side and 1 mm wide on the other. 

Some forms of C. longipes have a poorly developed 

labellum epichile and in this regard resemble 

C. halconensis (Ames) S.S. Ying. The latter taxon 

however is a smaller plant (to 30 cm tall) with narrower 

leaves (to 2.3 cm wide). These vegetative 

character states are given in the description of the 

Taiwanese Calanthe kooshunensis Fukuy. and for 

this reason the fi rst author still maintains that it 

should be treated as a synonym of C. halconensis 

rather than C. longipes. 

2. The distribution of Cephalantheropsis 

longipes is partly Transhimalayan in that the plant 

is recorded in an arc from Sikkim to Guangxi and 

Northern Vietnam, then across to Taiwan. It then 

spreads south into the Philippines. In Vietnam it is 

found in the north, middle and south (type locality 

for C. lateriscapa). The collection localities from 

southern Vietnam (Dalat/Langbian area), southern 

Thailand and northern Sumatra form an almost 

straight line when plotted on a map. Recently however 

C. longipes was found in Peninsular Malaysia, 

showing that it has spread south into the Peninsula 

(see Ong et al., 2011). Floral details (e.g. ellipticlanceolate, 

acute petals) of the Sumatran and Thai

120 

plants seem to agree quite well. The south 

Vietnamese plants however have more oblong, 

blunter petals as evidenced by the drawings of 

Seidenfaden (1992) and confi rmed after study of 

Everard 1767 7 (AMES). Further collecting is likely 

to uncover C. longipes in Bhutan, northern 

Thailand and Laos. 


The authors wish to thank Mr Andre 

Schuiteman for a copy of J.J. Smith’s drawing of 

Calanthe gracilis var. sumatrana in Leiden and the 

late Dr Gunnar Seidenfaden for copies of his drawings 

of types. Herbarium and library staff at the 

Harvard University Herbaria (A, AMES, GH), BM 


and K are thanked for their help and hospitality 

during the fi rst author’s visits. 

REFERENCES 

Averyanov, L.V. (2001). New Orchids in the Flora 

of Vietnam 1. Vietnam National University 

Journal of Science 17, 2:1–6. 

Ong, P.T., O’Byrne, P., Yong, W.S.Y. & Saw, L.G. 

(2011). Wild Orchids of Peninsular Malaysia. 

Forest Research Institute Malaysia. 

Ormerod, P.A. (1998). Review of Cephalantheropsis. 

Orchid Digest 62: 155–159. 

Seidenfaden, G. (1992). The Orchids of Indochina. 

Opera Botanica 114: 1–502.


A reassessment of the identity and rarity of Clerodendrum chlorisepalum (Lamiaceae) 

in Thailand and Vietnam 

INTRODUCTION 

The genus Clerodendrum L. (Lamiaceae) 

numbers some 150 species globally, of which 28 

are found in Thailand (Leeratiwong et al., 2011) 

and between 30 (Govaerts, 2011) and 34 (Phuong, 

2007) have been recognised in Vietnam. During 

IUCN Red List research on Clerodendrum by the 

fi rst author and the preparation of an account of the 

genus in Thailand by the second author, a discrepancy 

was noticed concerning the distribution of 

Clerodendrum chlorisepalum Merr. ex Moldenke. 

This species has been accepted previously as native 

in Thailand and Vietnam (Moldenke, 1963, 1980, 

1985; Govaerts, 2011). However, we have found 

that the concept of this elegant species had been 

confused by Moldenke, in its original description 

(Moldenke, 1963) and in subsequent reviews of its 

status (Moldenke, 1980, 1985). Although C. chlorisepalum 

had been included in the 1997 IUCN 

Red List (Walter & Gillett, 1998) as threatened in 

Thailand, with the additional category ‘I’ (recorded 

in nation but status not available), it had been omitted 

from later IUCN lists due to data defi ciencies. 

It was also omitted from recent checklists of threatened 

plants in Thailand (Pooma et al., 2005; 

Santisuk et al., 2006). 

JAMES A. WEARN* & CHARAN LEERATIWONG** 

ABSTRACT. Re-examination of the type material of Clerodendrum chlorisepalum Merr. ex Moldenke, and additional, previously 

cited collections from Thailand and Vietnam, has necessitated a clarifi cation of the identity and rarity of the species. This is due to 

confusion with the much more common plant C. nutans Wall. ex Jack. Clerodendrum chlorisepalum has been assessed as Vulnerable 

(VU B1ab(iii)+D) using the IUCN Red List categories due to redetermination of specimens and reappraisal of its distribution. 

Clerodendrum chlorisepalum is endemic in Vietnam and, therefore, hereby excluded from the Flora of Thailand. The Thai specimens 

are C. nutans. 

KEY WORDS: Clerodendrum, distribution, Lamiaceae, Red List, taxonomy. 

REASSESSMENT 

Despite having been included in the fi rst 

global Red List of threatened plants (Walter & 

Gillett, 1998), the distribution of C. chlorisepalum 

in Thailand was believed to be geographically wide 

ranging (specimens from Northern, Central and 

Southwestern fl oristic regions had been determined 

in herbaria as this species and cited by Moldenke 

(1963: 87; 1985: 452)), but with few individuals 

present in each region. The protologue of C. chlorisepalum 

was published in the Studies of the Flora 

of Thailand series (Moldenke, 1963). In doing so, 

Moldenke took up a name which had been coined 

more than two decades earlier by Merrill (who had 

never published it), then relating to a single specimen 

from Vietnam (Pételot 6401, see the revised species 

description, below). Nevertheless, ‘Clerodendrum 

chlorisepalum Merr.’ was a name which Moldenke 

had cited under Indo-China in his geographical 

summaries prior to its validation by him (Moldenke, 

1942: 59; Moldenke 1949: 136). Later, Moldenke 

thought he had seen material from Thailand and, 

using Merrill’s epithet, along with a second Pételot 

specimen ( (Pételot 8485) 

as the type, he described 

the species, adding Thailand to its distribution 

(Moldenke, 1971: 294). The trigger for Moldenke’s 

* Herbarium, Library, Art & Archives, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK. Email: j.wearn@kew.org 

** Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand.

122 

description of C. chlorisepalum was a then newly 

collected plant from Sai Yok in Southwest Thailand 

(Larsen ( 8515), 

which Moldenke believed to be 

conspecifi c with the Vietnamese collections of 

Pételot). However, this determination was erroneous, 

and has led to subsequent confusion of the identity 

and distribution of the species. 

Pételot 8485 does represent material of C. 

chlorisepalum as understood by Merrill, and that 

Following communications with staff at herbaria 

holding Thai specimens identifi ed as C. chlorisepalum 

and examination of that material, as well 

as investigation of undetermined collections and 

discussions with botanists local to collection localities, 

we are able to conclude that all material 

collected from Thailand, and previously perceived 

to be C. chlorisepalum, is in fact C. nutans. No 

collections of C. chlorisepalum from Laos have 

been found by us, although botanical collecting 

more generally has not been abundant in that country 

(Schuiteman & de Vogel, 2000; Newman et al., 

2007). These assertions reduce the known distribution 

of C. chlorisepalum signifi cantly – to Vietnam 

only (Le Trong Cuc (2005: 289); Phuong (2007: 

106); and see below). 

It is worth noting that, C. nutans (syn. C. wallichii 

Merr.) has also been frequently confused 

with C. laevifolium Blume, a species with larger 

calyces and subsessile leaves (see Wearn & 

Mabberley, 2011). 

It is rather ironic that a report from a project 

which had focused on the fl ora of Thailand contains 

the protologue of a species which was never 

actually found there. This situation is not unique 


collection was chosen by Moldenke as the type. We 

have found that the additional Thai collection, included 

at the end of the protologue (i.e. Larsen 

8515), and other putative material from Thailand, 

which was later cited by Moldenke (1985: 452), 

represent material of C. nutans Wall. ex Jack. 

Several characters clearly separate these collections 

from C. chlorisepalum (see Table 1). 

Table 1. Comparison of C. chlorisepalum with the species with which it is most commonly-confused, 

C. nutans, using revised concepts. 

C. chlorisepalum Merr. ex Moldenke C. nutans Wall. ex Jack 

Infl orescence Nodding, lax/spreading. Distinctly pendulous. 

Corolla (at anthesis) Tube 1.9–2.2 cm long, longer than 

lobes; lobes ovate-elliptic, each 3–4 

mm wide. 

Tube 0.8–1.2 cm long, equal in length 

to, or shorter than lobes; four lateral 

lobes obovate and 6–7 mm wide, 

upper one broadly elliptic and 4–4.5 

mm wide. 

and is reminiscent of Thunberg’s description of 

plants from Japan (often including the epithet 

‘japonica ‘ ’) which were, in fact, originally from 

China. More importantly, without critical reassessment 

of this material, the narrower ecological 

breadth and geographic distribution of C. chlorisepalum 

would not have been understood. 

Clerodendrum chlorisepalum Merr. ex Moldenke, 

Dansk Bot. Arkiv 23: 87. 1963, only type, not additional 

specimen cited. Type: Vietnam, Lao Cai 

Province, route from Laokey [Lao Cai] to Chope 

[Sa Pa], 1943, Pételot 8485 (holotype NY [image 

seen]; isotypes P, VNM [image seen]). 

Small shrub, 1–2 m tall; branchlets slender, 

pale brown, quadrangular, glabrous or subglabrous. 

Leaves decussate; blade ± elliptic, 8–15 cm long 

by 2.5–7 cm wide, 2.5–3.2 times as long as wide; 

margin subentire-repand; base acute; apex long 

acuminate; surfaces subglabrous, mid-green adaxially, 

paler abaxially; venation arcuate, 5–7 

pairs of secondary veins; petioles 1.5–3 cm long. 

Infl orescence a terminal thyrse, 13–30 cm long by 

8–18 cm wide, fi nely branched, lax and nodding 

with cymes borne laterally from main axis;

A REASSESSMENT OF THE IDENTITY AND RARITY OF CLERODENDRUM CHLORISEPALUM (LAMIACEAE) 

IN THAILAND AND VIETNAM (J.A. WEARN & C. LEERATIWONG) 

peduncles 4 [distally] –41 [proximally] mm long, 

glabrous to minutely puberulent. Calyx green 

(sometimes with pale reddish tinge to lobe apices), 

± campanulate, (8–)9–10.5 mm long at anthesis; 

lobes 5, united near to the base, lanceolate, ± glabrous, 

lacking glands or with some minute (diameter 

< 50 μm) glands adaxially. Corolla white, slightly 

scented, glabrous; tube 1.9–2.2 cm long; lobes 

ovate-elliptic, 1.3–1.4 cm long by 3–4 mm wide. 

Stamens 4, exserted, extending c. 15 mm from corolla 

tube. Ovary globose, glabrous. Style shortly 

bifi d apically, extending to c. 8 mm beyond the stamens. 

Fruit t not seen. 

Distribution.— The revised distribution of this 

species includes only three confi rmed localities in 

northern Vietnam, one in each of three provinces: 

Lao Cai [route from Laokey (Lao Cai) to Chope 

(Sa Pa), 25 Sept. 1943, Pételot 8485 (NY, P, 

VNM); Hoa Binh [Mai Chau District, Pa Co, 20 

Sept. 2005, Vu Xuang Puong et al. HNK 580 (HN, 

K)]; and Vinh Phuc [Soc Son Pagoda, Oct. 1937, 

Pételot 6401 (NY, P, VNM)]; as well as an additional, 

unconfi rmed record from Ha Tay Province 

[Ba Vi, no material seen]). 

Ecology.— Found in open forests and in 

semi-open grassland; on clay soils and limestone; 

958–1800 m altitude. Flowering August–October 

(at least). 

IUCN Conservation Status.— Since the type 

material was collected in 1943, there has been much 

regional unrest, causing destruction of habitats and 

buildings alike (Lao Cai being razed to the ground 

during the late 1970s). In 1986 the Hoang Lien Son 

Nature Reserve was created to protect parts of the 

natural landscape to the south and west of Sa Pa, 

although the Reserve only covers a relatively small 

(c. 30 km2 ) area. Today, this species is threatened 

by habitat destruction as land clearance continues 

at an alarming pace in Vietnam, especially for terraced 

agriculture on the hills where C. chlorisepalum 

is native. 

The total confi rmed provincial area of occupancy 

in Vietnam is less than 15,000 km2. The 

presence of C. chlorisepalum within this area is 

clearly sporadic. Hence, the area of actual occupancy 

is likely to be much smaller. We have therefore 

categorised C. chlorisepalum as Vulnerable 

(VU B1ab(iii)+D) using IUCN (2001) categories. 

123 

Although there may be some undiscovered populations, 

the distribution of this species appears to be 

fragmented within a restricted geographical region. 

Also, C. chlorisepalum has to our knowledge, so 

far been represented by single individuals at all 

known localities (established via personal communications), 

and therefore, may not be able to spread 

via root-suckers, unlike several other Clerodendrum 

species (e.g. C. chinense (Osbeck) Mabb., C. japonicum 

(Thunb.) Sweet, and C. paniculatum L.), 

though its biology is little known. 

Clarifi cation of the identity and distribution 

of this species is essential to avoid over-collection. 

It is critical that fecundity is not hampered by future 

collecting activities. We recognise that botanical 

exploration in the regions where this species is native 

has been rather low during the past 50+ years, 

but, considering the data available and discussions 

with in-country botanists, we believe that this assessment 

refl ects the knowledge of the species at 

this time. Clerodendrum chlorisepalum is not 

known to be cultivated, and so ex situ stocks are 

unlikely to exist. 

Etymology.— The epithet means ‘having 

green sepals’ (i.e. calyx lobes). 

Notes.— Clerodendrum chlorisepalum is a 

high altitude species unlike C. nutans. Clerodendrum 

chlorisepalum is also similar to C. disparifolium 

Blume, although the latter has smaller calyces and 

narrower corolla tubes, and is also much larger in 

stature (i.e. a large shrub to small tree). That species 

is common in Malesia and has been collected 

from southern Thailand but is absent from Vietnam. 

In 2005, collaborative fi eldwork between the 

Royal Botanic Gardens, Kew (K) and the Vietnamese 

Institute of Ecology and Biological Resources 

(IEBR) produced a signifi cant recent set of material 

from Vietnam. Among the specimens collected 

from northern Vietnam, was one which appears to 

be new material of C. chlorisepalum (Phuong ( et al. 

HNK 580). 

ACKNOWLEDGMENTS 

We are grateful to staff at the following herbaria 

for aiding our quest for material of C. chlorisepalum: 

AAU, BK, BKF, C, CMU, E, K, HNU, 

LL, NY, QBG, US and VNM. Eric Danell (Dokmai

124 

Botanical Garden, Chiang Mai) facilitated contact 

with botanists local to focal sites in Thailand. 

Particular thanks is also due to David Mabberley 

(Royal Botanic Gardens & Domain Trust, Sydney), 

Alan Paton (K), Steve Bachman (K), Kurt Keller, 

Peter Williams and James Maxwell (CMU) for 

valuable discussions. 

REFERENCES 

Govaerts, R. (2011). World checklist of selected 

plant families. Royal Botanic Gardens, Kew. 

Online, continually updated version URL 

http://apps.kew.org/wcsp/ [accessed 18 December 

2011]. 


Criteria Verson 3.1, IUCN Species Survival 

Commission, International Union for Conservation 

of Nature and Natural Resources. 

Information Press, Oxford. 

Le Trong Cuc. (2005). Danh luc các loài thuc vat 

Viet Nam [Checklist of plant species of 

Vietnam, in Vietnamese]. Tap 3, Nganh moc 

lan: Magnoliophyta. Hà Noi: Nhà Xuât Ban 

Nông Nghiep. 

Leeratiwong, C., Chantaranothai, P. & Paton, A. 

(2011). A synopsis of the genus Clerodendrum 

L. (Lamiaceae) in Thailand. Tropical Natural 

History 11: 177–211. 

Moldenke, H.N. (1942). The known geographic 

distribution of the members of the Verbenaceae 

and Avicenniaceae. Privately published, New 

York. 

________. (1949). The known geographic distribution 

of the members of the Verbenaceae, 

Avicenniaceae, Stilbaceae, Symphoremaceae, 

and Eriocaulaceae. Privately published, New 

York. 

________. (1963). Studies in the Flora of Thailand 

22. Avicenniaceae, Symphoremaceae, Verbenaceae. 

Dansk Botanisk Arkiv 23: 83–92. 

________. (1971). A fi fth summary of the Verbenaceae, 

Avicenniaceae, Stilbaceae, Dicrastylidaceae, 

Symphoremaceae, Nyctanthaceae, and 

Eriocaulaceae of the world as to valid taxa, 

geographic distribution, and synonymy. 

Volume 1. Phytologia Memoirs I. New Jersey. 


Moldenke, H.N. (1980). A sixth summary of the 

Verbenaceae, Avicenniaceae, Stilbaceae, Chloanthaceae, 

Symphoremaceae, Nyctanthaceae, and 

Eriocaulaceae of the world as to valid taxa, 

geographic distribution and synonymy. Phytologia 

Memoirs II. New Jersey. 

________. (1985). Notes on the genus 

Clerodendrum (Verbenaceae) XI. Phytologia 

58: 432–462. 

Newman, M., Ketphanh, S., Svengsuksa, B., 

Thomas, P., Sengdala, K., Lamxay, V. & 

Armstrong, K. (2007). A checklist of the vascular 

plants of Lao PDR. Royal Botanic 

Gardens, Edinburgh. 

Phuong, V.X. (2007). Thuc vât chí Viet Nam – 

Flora of Vietnam. Volume 6. Ho cơ roi ngưa – 

Verbenaceae [in Vietnamese]. Hà Noi: Nhà 

Xuât Ban Khoa Hoc Và Ky Thuât. 

Pooma, R. (2005). A preliminary check-list of 

threatened plants in Thailand. Forest Herbarium 

(BKF), National Park, Wildlife and Plant 

Conservation Department, Bangkok. 

Santisuk, T., Chayamarit, K., Pooma, R. & Suddee, 

S. (2006). Thailand red data: plants. ONEP 

Biodiversity Series, Volume 17. Offi ce of 

Natural Resources and Environmental Policy 

and Planning, Bangkok. 

Schuiteman, A. & de Vogel, E. (2000). Orchid genera 

of Thailand, Laos, Cambodia, and Vietnam. 

Vietnamese-English edition. Nationaal Herbarium 

Nederland, Leiden. 

Walter, K.S. & Gillett, H.J. (1998). 1997 IUCN red 

list of threatened plants. International Union 

for Conservation of Nature and Natural 

Resources, Gland. 

Wearn, J.A. & Mabberley, D.J. (2011).Clerodendrum 

(Lamiaceae) in Borneo. Systematic Botany 36: 

1050–1061.


Coelogyne phuhinrongklaensis (Orchidaceae), a new species for Thailand 

PAKORN TIPPAYASRI* & CHATCHAI NGERNSAENGSARUAY* 

ABSTRACT. Coelogyne phuhinrongklaensis Ngerns. & P. Tippayasri (Orchidaceae), a new species from northern Thailand is described, 

illustrated and presented. It is an epiphyte or lithophyte in montane forest at 1100–2100 m altitude. 

KEY WORDS: Coelogyne phuhinrongklaensis, Orchidaceae, new species, Thailand. 

INTRODUCTION 

Coelogyne Lindl. belongs to the tribe 

Arethuseae, subtribe Coelogyninae, subfamily 

Epidendroideae (Orchidaceae), and comprises 4 

subgenera and 19 sections (Pridgeon et al., 2005). 

It is a large genus of about 200 species, widely distributed 

in South and South-East Asia reaching 

across to the south-west Pacifi c Islands (Clayton, 

2002; Pridgeon et al., 2005). Thaithong (1999) recorded 

27 species of Coelogyne in Thailand based 

on the work of Seidenfaden (1975, 1995). 

During fi eld work in August 2006 and 2007, 

a new species of Coelogyne was discovered at Phu 

Hin Rong Kla National Park in Phitsanulok province, 

northern Thailand. This species belongs to 

Coelogyne section Elatae. The peduncle of all species 

in this section has sterile, imbricate bracts at 

the junction with the rachis. The rachis produces a 

single set of simultaneously opening fl owers 

(Pridgeon et al., 2005). 

DESCRIPTION 

Coelogyne phuhinrongklaensis Ngerns. & P. 

Tippayasri, sp. nov., Typus: Thailand, Phitsanulok, 

Phu Hin Rong Kla National Park, at the beginning 

of Man Daeng nature trail, near Man Daeng Forest 

Protection Unit, 1650 m altitude, 25 Aug. 2007, P. 

Tippayasri, C. Ngernsaengsaruay, O. Phueakkhlai 

& N. Anuraktrakoon 183 (holotype BKF, spirit 

specimen; isotypes BK, BKF, QBG, dried specimens). 

Figs. 1–2. 

Epiphyte or lithophyte, sympodial, glabrous. 

Rhizome creeping, terete, 5–6 mm in diam., woody, 

covered with papery, persistent overlapping scales, 

with roots at the nodes. Pseudobulbs distant, 3–7 

cm apart, (elongated) ovoid, 4.7–5.5 by 1.5–1.7 

cm, consisting of a single internode, fl eshy, with 

4–5 ribs; base enclosed by chartaceous, overlapping 

enlarged, later eroding scales. Leaves 2, subcoriaceous; 

petiole channeled, c. 2–7 by 0.3 cm; blade 

elliptic-oblong to lanceolate, 11.5–27 by 3–5 cm, 

apex acute or acuminate, margin entire, 5-nerved, 

midrib prominent beneath. Raceme hysteranthous 

(arising from apex of mature pseudobulb), curved, 

18–33 cm long; peduncle slender, terete, 11.5–23.5 

cm by 1–2 mm; rachis arched, 2–7 cm long, its 

base enclosed by 9–13 persistent, imbricate, leathery 

scale leaves, internodes 3.5–7 cm. Floral bracts 

dropped at anthesis, concave, ovate, c. 3.5 by 2 cm, 

apex acute, pale green turning pale green-yellowish 

to yellow before falling, caducous. Flowers (2–)3– 

4, showy, most open simultaneous, resupinate, 4–5 

by c. 5 cm. Median sepal l usually curved to the 

front, lanceolate, 3.5–4 by 1.3–1.4 cm, apex acute, 

3- or 5-nerved. Lateral sepals patent, lanceolate, 

3.5–4 by 1.3–1.4 cm, apex acute, 3- or 5-nerved. 

Petals recurved, linear, 3.4–3.8 by 0.5–0.6 cm, 

apex acute, 1-nerved. Lip 3-lobed, when fl attened 

pandurate, 2.7–3.2 by 2.5–3 cm; lateral lobes erect, 

margin entire, distinctly projecting in front, sinus 

1–1.3 cm, its top acute to rounded; keels 3, median 

keel at base of hypochile, short, swollen, conical, 

5–7 by 1.5–2 mm; lateral keels from base of lip to 

* Department of Botany, Faculty of Science, Kasetsart University, Chatuchak, Bangkok 10900, Thailand, email: fsciccn@ku.ac.th

126 

more than a halfway the epichile, 2.4–2.6 cm long, 

their basal part white, with entire, brown margin, 

terminal part of keels dark brown, with entire margin, 

with white median groove of lip between 

keels, with 2 brown longitudinal parallel bands 

from near tip of hypochile to more than a half of 

epichile between keels, 1–1.6 cm long, background 

of lip white, the areas from keels to lip margins 

with scattered brown small patches running from 

base of lip to more than a half of epichile; midlobe 

quadrangular, 1.6–1.8 by 1.2–1.6 cm, widening 

gradually to apex, apex emarginate, margin undulate 

and erose, terminal part of midlobe white; side 

lobes erect, broadly ovate, 0.8–1 by c. 0.8 cm, apex 

obtuse, margin entire, with brown veins. Column 

curved, spathulate in outline, c. 2.2 cm long, top 

winged, margin uneven dentate, terminal part 

white, basal part creamy white; anther short conical, 

c. 4 by 4 mm, apex bilobed ; pollinia 4 in 2 pairs, 

obovate, c. 2.5 by 1.5 mm. Ovary including pedicel 

1.5–2.3 cm. Capsule not seen. 

Colour notes: Sepals and petals white, inside 

at base yellowish, pale green in bud. Lip white, hypochile 

inside around the median keel slightly orange. 

Lateral lobes with irregular shaped spots of 

various size, fading to small spots on the base of 

the epichile; front half of hypochile to halfway the 

epichile with two brown bands between the keels. 

Median keel brown; lateral keels white with brown 

margin, the top third all brown. 

Thailand.— NORTHERN: Phitsanulok [Phu Hin 

Rong Kla National Park, Lan Hin Taek, 1100 m altitude, 

19 Aug. 2006, P. Tippayasri & C. Kasetluksamee 

33 (Herb. of the Department of Botany, Kasetsart 

University, spirit specimen); Phu Hin Rong Kla 

National Park, at the beginning of Man Daeng 

nature trail, near Man Daeng Forest Protection 

Unit, 1650 m altitude, 25 Aug. 2007, P. Tippayasri, 

C. Ngernsaengsaruay, O. Phueakkhlai & N. Anuraktrakoon 

183 (BK, BKF, QBG); Phu Soi Dao 

National Park, Chattrakan District, 2100 m altitude, 

17 Sept. 2008, P. Suksathan 4611 (QBG)]. 

Distribution.— Known only from northern 


Ecology.— Epiphyte on trees and lithophyte 

on moss-covered sandstone rock platforms in montane 

forest, 1100–2100 m altitude. Flowering: 

August–September. 


Vernacular.— Ueang thian pak si nam tan 

(เอื้องเทียนปากสีนําตาล). 

Etymology.— The generic name, Coelogyne 

from the Greek koilos, hollow, cavity, and gyne, 

female, which refers to the concave stigma (Pridgeon 

et al., 2005). The specifi c epithet is named after 

Phu Hin Rong Kla National Park where the authors 

found and collected the type specimens. 

IUCN Conservation Status.— In Thailand 

the species is known from small populations in Phu 

Hin Rong Kla National Park and Phu Soi Dao 

National Park. This species grows on trees and on 

moss-covered sandstone rock platforms, which are 

quite close to nature trails. Every year, many tourists 

visit these protected areas, which might impact 

on the species habitat. Fruit development in 

Coelogyne phuhinrongklaensis was not observed 

during several fi eld visits to the sites in 2006 and 

2007, which means that growth of population size 

is limited at most. The species has a very restricted 

distribution in northern Thailand. We therefore 

suggest the conservation status ‘Vulnerable’ (VU) 

(IUCN, 2001). 

Notes.— Coelogyne fi lipeda differs from the 

Thai Coelogyne phuhinrongklaensis in its dimensions 

of plant and fl ower parts (peduncle 1.8–2 cm 

against 11.5–23.5 cm, size of sepals 2 cm against 

3.5–4 cm, epichile of lip fl at against plicate, absence 

of median keel against a short swollen keel at 

the base of the hypochile, and base color of fl ower 

light green or yellow-green with purple veins on lip 

(Seidenfaden, 1992; Clayton, 2002) against white 

with brown markings on the lip). 


The authors would like to thank Dr E.F. de 

Vogel, the World expert on Thai Coelogyne and 

Professor Dr John Parnell for their kind improvements 

to the manuscript. Thanks also to Miss 

Orporn Phueakkhlai, Miss Nongnoot Anuraktrakoon 

and Mr Charin Kasetluksamee for their assistance 

in the fi eld works. This work was supported by 

Faculty of Science, Kasetsart University (Science 

Research Fund, ScRF 2006) and the Graduate 

School, Kasetsart University.

COELOGYNE PHUHINRONGKLAENSIS (ORCHIDACEAE), A NEW SPECIES FOR THAILAND (P. TIPPAYASRI & C. NGERNSAENGSARUAY) 127 

Figure 1. Coelogyne phuhinrongklaensis Ngerns. & P. Tippayasri: A. habit; B. fl ower; C. dorsal sepal (inside and outside); D. lateral 

sepal (inside and outside); E. petal (inside and outside); F. lip (lower and upper surfaces); G. longitudinal section of lip (side view); 

H. column, ovary and pedicel; I. lip and column; J. anther (below and above); K. pollinia. Drawn by P. Tippayasri.

128 


Figure 2. Coelogyne phuhinrongklaensis Ngerns. & P. Tippayasri: A. habit; B–C. infl orescence with fl ower buds and fl oral bracts; 

D–E. fl ower (front view); F. fl ower (side view); G. Lateral view of fl ower, sepals and petals removed); H. lip. Photographed by P. 

Tippayasri (A–C, G–H) and C. Ngernsaengsaruay (D–F). 

A 

D G 

E F H 

B 

C

COELOGYNE PHUHINRONGKLAENSIS (ORCHIDACEAE), A NEW SPECIES FOR THAILAND (P. TIPPAYASRI & C. NGERNSAENGSARUAY) 129 

REFERENCES 

Clayton, D.A. (2002). The Genus Coelogyne: A 

Synopsis. Natural History Publications 

(Borneo) Sdn. Bhd., Kota Kinabalu, Sabah, 

Malaysia. 

IUCN (2001). IUCN Red List Categories and 

Criteria: Version 3.1. Prepared by the IUCN 

Species Survival Commission. IUCN, Gland, 

Switzerland and Cambridge, UK. 

Pridgeon, A.M., Cribb, P.J., Chase, M.W. & 

Rasmussen, F.N. (eds.). (2005). Genera 

Orchidacearum Vol. 4. Oxford University 

Press, Oxford. 

Seidenfaden, G. (1975). Orchid genera in Thailand 

3 Coelogyne Lindl. Dansk Bot. Ark. 29(4): 

1–94. 

________. (1992). The orchids of Indochina 

Coelogyne Lindl. Opera Bot. 114: 106–120. 

________. (1995). Contributions to the orchid fl ora 

of Thailand 12: Coelogyne. Opera Bot. 124: 

17–18. 

Thaithong, O. (1999). Orchids of Thailand. Offi ce 

of Environmental Policy and Planning, Bangkok, 

Thailand.


Tripogon purpurascens (Chloridoideae: Poaceae): a native Thai grass recently recognized 

ATCHARA TEERAWATANANON 1, 2, 3 & SARAWOOD SUNGKAEW 2, 3, 4 * 

ABSTRACT. Tripogon purpurascens Duthie is reported here as a new record for Thailand. The species is described and illustrated. 

A key to Tripogon occurring in Thailand is provided. 

KEY WORDS: conservation status, new record, taxonomy, Thailand. 

INTRODUCTION 

Tripogon Roem. & Schult. is a genus of approximately 

40 species, most of which are native to 

the Old World tropics, especially Asia and Africa 

(Phillips & Chen, 2002, Clayton et al., 2006 onwards, 

Newmaster et al., 2008). The genus can be described 

as a natural group based mainly on the presence of 

a single terminal raceme and by having lateral 

nerves of the glabrous lemmas (Phillips & Launert, 

1971; Clayton & Renvoize, 1986; Veldkamp & 

Phillips, 2003; Newmaster et al., 2008). 

1 Natural History Museum, National Science Museum, Technopolis, Pathum Thani 12120, Thailand. 

2 Center for Advanced Studies in Tropical Natural Resources, Kasetsart University. 

3 Kasetsart University Center of excellence on bamboos, Kasetsart University. 

4 Department of Forest Biology, Faculty of Forestry, Kasetsart University, Bangkok 10900, Thailand. 

* Corresponding author 

KEY TO THAI TRIPOGONN SPECIES 

In Asia, the major taxonomic studies of 

Tripogon were prepared by Bor (1960), Phillips & 

Chen (2002), Veldkamp & Phillips (2003) and 

Newmaster et al. (2008), in which four species 

were recognised for Thailand. During preparation 

of the taxonomic treatment of Poaceae for the Flora 

of Thailand, specimens of Tripogon purpurascens 

Duthie, which is new to Thailand, were collected 

from Nong Bau Lam Phu Province (north-eastern 

Thailand), constituting a new record for the 

country. 

1. Central awn as long as or longer than the lemma body, lateral veins extended T. trifi dus 

1. Central awn shorter than the lemma body, lateral veins not extended 

2. Lemma midvein extended into a small point, 0.2−0.5 mm long T. purpurascens 

2. Lemma midvein extended into an awn, 1−2.5 mm long 

3. Leaf sheath glabrous on the shoulders. Lower glume 4−5 mm long T. siamensis 

3. Leaf sheath with dense woolly hairs on the shoulders. Lower glume 1.7−2.5 mm long 

4. Spikelet 12−15 mm long (without the awn). Upper glume c. 5 mm long. Leaf blade fl at, c. 3 mm wide T. larsenii 

4. Spikelet 4.5−15 mm long (without the awn). Upper glume 3−3.8 mm long. Leaf blade involute, 0.5−0.8 mm wide T. thorelii 

Tripogon purpurascens Duthie, Ann. Hort. Bot. 

Calc. 9: 74, t. 92. 1901; Bor, Grasses Burma, Ceyl., 

Ind. & Pakist.: 522. 1960; Noltie, Fl. Bhutan 3: 

655. fi gs. 32/k-m. 2000; S.M. Phillips & S.L. Shen, 

Fl. China 57(4): 913. 2002. Type: India, Uttarakhand, 

Tehri Garhwal, Tons Valley, alt. 4,000 ft, 5 May 

1900, Duthie 23532 (holotype K! (K245023)).— 

Festuca fi liformis Nees ex Steud. Syn. Pl. Glumac. 

1: 302. 1854. non Pourr. 1788.— Tripogon abyssinicus 

auct. non. Nees. ex Steud., Hook.f., Fl. 

Brit. India 7: 287. 1896.— T. jacquemontii var. 

submuticus Hook.f., Fl. Brit. India 7: 287. 1897.

TRIPOGON PURPURASCENS (CHLORIDOIDEAE: POACEAE): A NATIVE THAI GRASS RECENTLY RECOGNIZED 

(A. TEERAWATANANON & S. SUNGKAEW) 

Type: India, Simla, alt. 7,000−8,000 ft, Aug. 1889, 

Duthie 10149 (holotype K! (K245021)).― T. submuticus 

Hack. ex Hook.f., Fl. Brit. India 7: 287. 

1896. nom. invaidl. pro syn. T. jacquemontii var. 

submuticus.― T. hookerianus Bor, Grasses Burma, 

Ceyl., Ind. & Pakist.: 522. 1960. Type: India, 

Hazara, alt. 8,000 ft, 24 July 1896 (lectotype K! 

(K245025), designated here). 

Perennial, caespitose. Culms 5−20 (−35) cm 

tall, erect; sheaths 1−2 cm long, margins scarious, 

basal sheaths becoming fi brous; ligule a ciliate 

membrane. Leaves: blades fi liform, 3−15 × 0.3−1 cm, 

margins scabrous, upper surface scabrous-hispidulous, 

lower surface glabrous. Racemes solitary, 

(2−) 4−7 (−17) cm long; rhachis glabrous. Spikelet: 

laterally compressed, oblong, 4−7 × 1 mm. Florets 

(2−) 4−6 (−8), imbricate; rhachilla c. 0.4 mm long; 

lower glume narrowly lanceolate, 1.5−2.5 (−3) mm 

long, acuminate, 1-nerved, scabrous on nerve. 

Glumes: upper glume lanceolate, (2.2−) 3−4.5 mm 

long, margins scarious, apex acute to acuminate, 

1-nerved; lemma oblong-lanceolate, 2−4 mm long, 

bifi d, 3-nerved, median nerve reaching the upper 

margin and exerted as a small point, 0.2−0.5 mm 

long, lateral nerves not extended; palea slightly 

shorter than lemma, apex obtuse, keels narrowly 

winged or wingless, ciliolate; callus hairy, hairs 

0.3−1 mm long. Stamens 3, anthers 0.6−2 mm 

long; stigmas c. 4 mm long. Caryopsis oblong, c. 1 

mm long. 

Thailand.— NORTH-EASTERN: Nong Bua Lam 

Phu [Muang, 8 July 2001, Teerawatananon & 

Sungkaew 2001-290 0 ( AAU, Thailand Natural History 

Museum)]. 

India.— Himachal Pradesh: Simla, Duthie 

10149 (K: K245021); Simla, Kotgurh, Thomson 

s.n. (K: K245022); Juansar, Tonse Valley near 

Thadyar, Duthie 19784 (K: K245024); Jammu and 

Kashmir: Pahlgan, Lidder, Stewart 21625 (K). 

Pakistan.— Northwest Frontier Province: 

Hazara Kagan Valley, Inayat 20364 (K: K245025); 

l.c., Inayat 20364a (K: K245026); Punjab: 

Rawalpindi, Stewart 29220 (K). 

Nepal.— Micha, near Jumla, Polunin, Sykes 

& Williams 4486 6 ( K). 

Distribution.— Saudi Arabia, Yemen, 

Afghanistan, Pakistan, India (type), Nepal, Bhutan, 

China. 

131 

Ecology.— Open areas on sandstone rock 

formations. 

Notes.— Tripogon purpurascens is confi ned 

to a type of habitat which is commonly found in the 

north-eastern Thailand. The species is distributed 

from the Arabian Peninsula to Asia, especially in 

arid places and in open stony or rocky areas at 

moderate altitudes in the Himalaya from India and 

China to Pakistan (Phillips & Chen, 2002). The occurrence 

of this species in Thailand is thus an extension 

of its geographical range. The discontinuous 

geographical range of T. hookerianus from 

Myanmar and Indo-China may simply be due to 

insuffi cient collecting. 

Tripogon hookerianus was published by Bor 

(1960) based on Wingate s.n. (K) and Inayat 20364 

(K). K We designate Inayat 20364 4 here as the lectotype. 


The authors are grateful to the curators and 

the staff of K and TCD for the use of specimens. 

Thanks to Drs Simon Laegaard and Sylvia M. 

Phillips for helpful suggestions. 

REFERENCES 

Bor, N.L. (1960). The Grasses of Burma, Ceylon, 

India and Pakistan (Excluding Bambuseae),vol. 

1. International Series of Monographs on Pure 

and Applied Biology, Division: Botany. 

Pergamon Press, Oxford, London. 

Clayton, W.D. & Renvoize, S.A. (1986). Genera 

Graminum: grasses of the world. – Kew 

Bulletin Additional Series 13: 1–389. 

Clayton, W.D, Vorontsova, M.S, Harman, K.T & 

Williamson. H. (2006 onwards). GrassBase: 

The Online World Grass Flora. http://www. 

kew.org/data/grasses¬db.html. [accessed 2 

August 2011; 15:30 GMT]. 

Holmgren, P.K. & Holmgren, N.H. (1998 [continuously 

updated]). Index Herbariorum: a global 

directory of public herbaria and associated 

staff. New York Botanical Garden’s Virtual 

Herbarium. Published on the internet http:// 

sweetgum.nybg.org/ih/. [accessed 2 August 

2011; 08:00 GMT].

132 


Figure 1. Tripogon purpurascens: A, B. habit; C. spikelet; D. fl oret; E. lower glumes; F. upper glumes; G. lemmas; H. palea. Drawn 

by A. Teerawatananon from A. Teerawatananon & S. Sungkaew 2001-290.

TRIPOGON PURPURASCENS (CHLORIDOIDEAE: POACEAE): A NATIVE THAI GRASS RECENTLY RECOGNIZED 

(A. TEERAWATANANON & S. SUNGKAEW) 

McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, 

V., Hawksworth, D.L., Marhold, K., Nicolson, 

D.H., Prado, J., Silva, P.C., Skog, J.E., 

Wiersema, J.H. & Turland, N.J. (2006). 

International Code of Botanical Nomenclature 

(Vienna Code) adopted by the Seventeenth 

International Botanical Congress Vienna, 

Austria, July 2005. A. R. G. Gantner Verlag, 

Liechtenstein. 

Newmaster, S.G., Velusamy, B., Murugesan, M. & 

Ragupathy, S. (2008). Tripogon cope, a new 

species of Tripogon (Poaceae: Chloridoideae) in 

India with a morphometric analysis and synopsis 

of Tripogon in India. Systematic Botany. 

33: 695–701. 

Phillips, S.M. & Chen, S.L. (2002). The genus 

Tripogon (Poaceae) in China. Kew Bulletin 57: 

911–924. 

Phillips, S.M. & Launert, E. (1971). A revision of 

the African species of Tripogon Roem. & 

Schult. Kew Bulletin 25: 301–322. 

133


Two new records of Xyris L. (Xyridaceae) for Thailand 

PHONGSAK PHONSENA*, PRANOM CHANTARANOTHAI*, 1 & AMORNRAT MEESAWAT* 

ABSTRACT. Xyris borneensis Rendle and X. linifolia P.Royen, are newly recorded for Thailand. The petals, capsules and seeds of 

X. linifolia are described for the fi rst time. A provisional conservation status for each species is given. 

KEY WORDS: Conservation status, taxonomy, Thailand, Xyris, Xyridaceae 

INTRODUCTION 

The family Xyridaceae (with fi ve genera, but 

only one genus, Xyris L. in Asia) was published in the 

Flora of Thailand comprising 11 species (Hansen, 

1987). In the process of taxonomic revision of 

Xyridaceae in Thailand and SE Asia, recently 

additional material was collected, and among these 

collections two new records for Thailand were 

found. The fi rst species, X. X borneensis Rendle, was 

hitherto included in X. X bancana Miq. by Hansen 

(1979a; 1983; 1987). However, we examined type 

material of both names and found that X. X borneensis 

is different from X. X bancana. Also the anatomy of 

leaves and micromorphological details of the 

fertile bract confi rm their distinction. The second 

species, X. X linifolia P.Royen, discovered in Bueng 

Kan Province, NE Thailand was not known from 

Thailand even though it was recorded from a nearby 

places in Laos (Hansen, 1979b: 1983; 1987; 

Van Royen, 1954). Hansen (1987) included this 

species in his key, at the time only known from 

Laos assuming that it was likely to occur in 

Thailand. He did not include a description because 

no Thai material of X. X linifolia had yet been 

collected. 

MATERIALS AND METHODS 

The descriptions and illustrations below are 

based on Thai collections. Measurements of 

morphological characters were taken from living 

mature plants in fl ower and/or in fruit. Leaf width 

was measured in the median part of the blade. 

Sepal, petal, stamen and pistil morphology was 

measured from fully opened fl owers. Seed and 

fertile bract morphology was investigated by scanning 

electron microscopy (SEM) and light microscopy 

(LM). 

DESCRIPTION OF TWO NEW RECORDS 

FROM THAILAND 

1. Xyris borneensis Rendle, J. Bot. 37: 506. 1899; 

Malme, Bull. Jard. Bot. Buitenzorg, III, 10: 387. 

1929; P.Royen in Steenis, Fl. Males., Ser. 1, 

Spermat. 4: 370. 1953.— Type: Malaysia, Sabah 

(North Borneo), Burbidge 1877– 7 78 (holotype 

BM!; isotype K!). Fig. 2. 

Tufted perennial herb, 20–40(–75) cm tall. 

Leaves subterete or linear, wiry, twisted, 6–30(–40) 

cm by 0.4–1.2 mm, forming a ligule 0.5–1 mm 

long; blade basally rugulose with short transverse 

ridges, margin entire, apex bluntly-oblique. Scape 

terete or subterete with 7–8 low ridges below spike, 

20–40(–75) cm by 0.5–1 mm. Spike ellipsoid, 5–7 

by 3–4 cm. Sterile bracts 4–8, greenish- to reddishbrown, 

elliptic to ovate, 4–6 by 2–3 mm, margin 

entire, lacerate at apex. Fertile bracts greenish- to 

reddish-brown, ovate to elliptic, 6–8 by 4–6 mm, 

* Applied Taxonomic Research Center, Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen 40002, 


1 Corresponding author. Email: pranom@kku.ac.th

TWO NEW RECORDS OF XYRIS L. (XYRIDACEAE) FOR THAILAND (P. PHONSENA, P. CHANTARANOTHAI & A. MEESAWAT) 135 

margin lacerate near apex; stomate fi eld ovate to 

elliptic, 3–6 by 2.5–5 mm, with papillose epidermis. 

Lateral sepals pale brown, 3–4 by 0.7–1 mm, 

crest ciliate at least in the upper half. Petal limbs 

obovate, yellow, 4–8 by 2–4 mm, margin distally 

entire. Staminodes bibrachiate, branches elongatepenicillate, 

ca. 1.5 mm long. Anthers oblong, 1.2– 

1.5 mm long, shallowly bifi d, deeply sagittate; fi laments 

ca 1 mm long. Style terete, 3–3.5 mm long, 

distally 3-branched, branches ca. 1.5 mm long. 

Capsules ellipsoid to obovoid, dark brown, 5–6 by 

2.5–3 mm. Seeds brown, translucent, ovoid, 0.4– 

0.5 mm long, 0.2–0.3 mm in diam, with 14–16 longitudinal 

ridges and with 6–9 transverse ridges. 

Thailand.— PENINSULAR: Trang [Sikao, Hat 

Chao Mai NP, 21 Aug. 2010, Phonsena & 

Sutthisaksopon 6548 (BK!, BKF!, KKU!, L!)]; 

Songkhla [Thepha, 14 June 1992, Laesen et al. 

4291 (A!, PSU!)]; Pattani, 17 March 1923, Kerr 

7882 (K!). 

Distribution.— Peninsular Malaysia, Indonesia 

(Banka, Billiton), Borneo (Kalimantan, Sarawak, 

Brunei, Sabah). 

Ecology.— On open, sandy soil, wet places 

in strand vegetation, at sea level up to 150 m, except 

a specimen of de Vogel et al. 8134 ( (L!), 

which 

was collected from Nabawan, Sabah at 700 m altitude. 

Flowering August to September; the fl owers 

usually open mid-morning (ca. 10.00 a.m.) and 

persist until mid-afternoon (ca. 2.00 p.m.). 

Vernacular.— Kathin sai (กระถินทราย). 

Conservation status.— Although this species 

is found in strand vegetation, which is under threat 

from human disturbance, its wide distribution 

means a conservation status of Least Concern (LC) 

(IUCN, 2001) is most appropriate. 

Notes.— Xyris borneensis was placed by 

Hansen (1979a) in the synonymy of X. X bancana 

Miq. along with X. X ridleyi Rendle, X. X glaucella 

Malme, X. X chinensis Malme and X. X subcomplanata 

Malme, reporting that X. X borneensis, from Borneo, 

has subterete leaves but that it is otherwise similar. 

Many specimens were misidentifi ed as X. X bancana. 

However, we hold a distinct view: for example, the 

stomate fi eld of X. X borneensis is covered with papillae 

(versus non-papillose in X. X bancana), the 

epidermal cells at the base of the leaf of X. X 

borneensis possess striate papillae (versus a row of 

minute papillae in X. X bancana), the leaf of X. X 

borneensis possesses a large air space at the central 

of section (versus many dispersed air spaces between 

vascular bundles in X. X bancana) and the leaf 

of X. X borneensis possesses six compound vascular 

bundles and two single vascular bundles (versus 11 

compound vascular bundles and three single vascular 

bundles in X. X bancana) (Table 1 and Figure 

4E–I). Therefore, we regard X. X borneensis as a 

species in its own right. 

2. Xyris linifolia P.Royen, Blumea 7: 477. 1954; 

B.Hansen, Acta Phytotax. Geobot. 30, 4–6: 192: 

1979; in Leroy, Fl. Cambodge, Laos & Viet-Nam 

20: 157, Pl. 31, 8. 1983; in Smitinand & Larsen, Fl. 

Thailand 5, 1: 131. 1987.— Type: Laos, Mt Kan, 

Smiles s.n. (holotype K!). Fig. 3. 

Solitary perennial herb, 25–55(–65) cm tall, 

base abruptly dilated and bulbous. Bulb ovoid, 1.5– 

3 by 0.8–1.4 cm. Leaves 1(or 2) per plant, subterete 

to terete, twisted, without a ligule; blade smooth, 

20–40 cm by 0.8–1.5 mm, margin entire, apex 

bluntly oblique to acute. Scape terete, twisted, 25– 

55(–65) cm by 0.7–1.5 mm. Spike ovoid to ellipsoid, 

0.9–1.4 by 0.6–1.1 cm. Sterile bracts 4–8, 

reddish- or dark brown, elliptic to ovate, 4–6 by 

3–5 mm, margin entire, lacerate at apex. Fertile 

bracts reddish- or dark brown, ovate, 5–7 by 4–5 

mm, margin entire; stomate fi eld ovate to elliptic, 

1.8–2 by 1–1.2 mm. Lateral sepals hyaline, 5.5–7 

by 1–1.5 mm, crest entire or coarsely serrate. Petal 

limbs yellow, obovate, 10–12 by 8–10 mm, margin 

distally lacerate. Staminodes bibrachiate, branches 

elongate-penicillate, 3–3.5 mm long. Anthers oblong, 

2.5–3 mm long, shallowly bifi d, deeply sagittate; 

fi laments 1–1.5 mm long. Style terete, 3.5–4.5 

mm long, distally 3-branched, branches ca 2.5 mm 

long. Capsules brown, obovoid to ellipsoid, 4–6 by 

3–4 mm. Seeds yellowish-brown or brown, translucent, 

ellipsoid, 0.9–1 mm long, ca. 0.3 mm in 

diam., with 9–11 dark brown, longitudinal ridges 

and with 1 or 2 transverse ridges. 

Thailand.— NORTH-EASTERN: Bueng Kan 

[(Si Wilai, Na Sing, Ban Na Sai, 22 Sept. 2009, 

Phonsena & Boonsuk 6426 6 ( BKF!, L!); 25 Sept. 

2009, Phonsena & Boonsuk 6626 6 ( KKU!); 28 

Nov. 2010, Phonsena & Sutthisaksopon 6540 

(KKU!, QBG!); Phon Charoen, Ban Nong Paen,

136 

26 July 2009, Phonsena 6394 (BKF!, KKU!); 

Bueng Khong Long, Pho Mak Khaeng, Ban Dong 

Chomphu, 8 Aug. 2010, Phonsena & Sutthisaksopon 

6547 7 ( KKU!)]. 

Distribution.— Laos. 

Ecology.— On open, grassy places, sandy 

soil in deciduous dipterocarp forest, at 180–200 m 

altitude. Flowering June to October; the fl owers 

open during late morning (ca. 10.30 a.m.) and persist 

until mid-afternoon (ca. 2.00 p.m.). 

Vernacular name.— Kathin nam khong 

(กระถินนําโขง). 

Conservation status.— Xyris linifolia is 

found in a small area in deciduous dipterocarp forest 

of the Mekong basin of Laos and Thailand. 

Only three localities in Thailand are known, each 

consisting of a small population. Its habitat is 

changing rapidly due to disturbance by logging 


Table 1. Main differences between Xyris borneensis Rendle and X. X bancana Miq. 

Characters X. borneensis X. bancana 

Habitat lowland plant, in strand vegetation 

at sea level up to 150 m alt. 

(except a specimen of de Vogel 

et al. 8134, L) 

lowland or lower montane plant, in 

various forest types (include strand 

vegetation) from sea level up to 

1300 m alt. 

Leaf: shape subterete or linear, wiry fl at, linear 

size 6–30(–40) cm by 0.4–1.2 mm 30–70 cm by (1–)4–9 mm 

indumentum rugulose, striate when dry not rugulose and striate when dry 

Leaf anatomy: 

epidermal cells at 

the median part of leaf 

without papillae with a row of minute papillae 

epidermal cells at 

the base of leaf 

air space with a large air space at the central 

of section 

vascular bundles 6 compound vascular bundles, 2 

simple vascular bundles 

with striate papillae with a row of minute papillae 

with many dispersed air spaces 

between vascular bundles 

11 compound vascular bundles, 3 

simple vascular bundles 

Stomate fi eld: 

size 3–6 by 2.5–5 mm 1.5–2 by 1–2 mm 

indumentum papillose non-papillose 

activities and various types land uses. An 

Endangered (EN), rating (IUCN, 2001) is merited. 

Notes.— Van Royen (1954) in his original 

description of X. X linifolia, indicated that the single 

known collection at the foothill of Mt Kau, Siam 

was made by Smiles in 1893. Hansen (1983) 

argued that Mt Kau should be Mt Kan, not in Siam 

(Thailand) but in Laos, where Smiles travelled 

along the Mekong River at the time. However, 

Hansen (1987) included this species in his key for 

the treatment in the Flora of Thailand, without a 

description, assuming that it was likely to occur in 

Thailand also. 


The fi rst author would like to thank the directors, 

curators and staff of A, BK, BKF, BM, C, K, 

L, P, PSU and QBG for their assistance during my 

visits. Financial support from the Graduate School,


Figure 1. Distribution of Xyris bancana ( ), X. X borneensis ( ) and X. X linifolia ( ). 

Khon Kaen University, is acknowledged. Thanks 

are due to Dr Brigitta Duyfjes (L) for valuable help 

in preparing the manuscript, to Mr Phanom 

Sutthisaksopon and Mr Boonchuang Boonsuk for 

assistance in fi eldwork and to Mr Theerawat Srisuk 

and Mr Woranart Thammarong (Department of 

Biology, Faculty of Science, Khon Kaen University) 

for the beautiful drawings. 

REFERENCES 

Hansen, B. (1979a). Notes on Asian species of 

Xyris. Botanisk Tidsskrift 74 (1):22. 

________. (1979b).The genus Xyris in Thailand 

and Indochina. Acta Phytotaxonomica et 

Geobotanica 30 (4–6): 192. 

Hansen, B. (1983). Xyridaceae. In: J.S. Leroy 

(ed.), Flore du Cambodge du Laos et du 

ViêtNam 20: 151–164. Muséum National 

d’Histoire Naturelle, Paris. 

________. (1987). Xyridaceae. In: T. Smitinand & 

K. Larsen (eds.), Flora of Thailand 5(1): 130– 

138. The Forest Herbarium, Royal Forest 

Department, Bangkok. 

IUCN (2001). IUCN Red List Categories and 

Criteria Version 3.1. IUCN Species Survival 



Van Royen, P. (1954). Some new Australasian species 

of Xyris. Blumea 7: 477. 1954.

138 

A 

1 cm 

B 


Figure 2. Xyris borneensis Rendle: A. Habit; B. leaf; C. junction of leaf showing a ligule; D. spike; E. fertile bract; F. lateral sepal; 

G. petal limb and stamen; H. staminode; I. style; J. seed. All from Phonsena & Sutthisaksopon 6548 (KKU). Drawn by W. Thammarong. 

1 cm 

D 

G 

I 

1 mm 

H 

1 cm 

E 

1 mm 

F 

C 

J 

1 mm 

1 mm m 

1 mmm 

1 mm 

100 µm

A 


1 cm 

B 

I 

C 

D 

1 cm 

Figure 3. Xyris linifolia P.Royen: A. habit; B. leaf lamina and leaf apex; C. junction of leaf and sheating; D. spike; E. fertile bract; F. 

lateral sepal; G. petal limb and stamen; H. staminode; I. style; J. seed. A–I from Phonsena & Sutthisaksopon 6540 (KKU); J from 

Phonsena & Boonsuk 6426 6 (KKU). Drawn by T. Srisuk. 

E 

1 mm 

4 mm 

5 mm 

J 

H 

G 

F 

2 mm 

3 mm 

100 μm 

5 mm 

1 mm

140 


Figure 4. Xyris linifolia P.Royen: A. habit; B. infl orescence.— X. X borneensis: C. infl orescence; D. habit; E. vascular bundle in transverse 

section of leaf.— X. X bancana: F. vascular bundle in transverse section of leaf; G. epidermal cells and stoma at the median part 

of leaf showing rows of minute papillae.— X. X borneensis: H. epidermal cells and stomata at the median part of leaf; I. epidermal 

cells and stoma at the base of leaf showing striate papillae. A from P. Phonsena 6394 (KKU); B from P. Phonsena & P. Sutthisaksopon 

6540 (KKU); C, D, E, H & I from P. Phonsena & P. Sutthisaksopon 6548 (KKU): F & G from P. Phonsena 6698 (KKU). 

Photographed by P. Phonsena (A, B, D–I) and P. Sutthisaksopon (C).


Fimbristylis pubisquama (Cyperaceae), a new record for the Flora of Thailand 

KAMOLHATHAI WANGWASIT*, KHANIT WANGWASIT** & PRANOM CHANTARANOTHAI*** 

ABSTRACT. Fimbristylis pubisquama Kern is newly recorded for Thailand. A description illustration of this taxon are provided. An 

emended portion of the key to Fimbristylis in the Flora of Thailand account of this genus is proposed. 

KEY WORDS: Fimbristylis, Flora of Thailand, key, taxonomy 

INTRODUCTION 

During fi eldwork in north-eastern Thailand, 

Fimbristylis sieboldii Miq. ex Franch. & Sav. and a 

closely allied species were collected. The former 

was misidentifi ed in the Flora of Thailand 

Cyperaceae account (Simpson & Koyama, 1998) 

as F. ferruginea (L.) Vahl (Zhang et al., 2010). The 

latter was identifi ed as F. pubisquama Kern and is 

new to Thailand. With the discovery of F. pubisquama 

and an earlier discovery of Fimbristylis 

alata E.-G. Camus by J F Maxwell (Maxwell 2002) 

the total number of Fimbristylis species in Thailand 

now stands at 62. 

TAXONOMY 

Fimbristylis pubisquama Kern, Blumea 8: 131. 

1955.— F. compressa Boeck. in Linnaea 38: 387. 

1874. (non Roem. et Schult. 1817).— Type: India, 

Wight 2902 (holotype P; isotype L). Figs. 1–2. 

Annual. Culms tufted, 20–80 cm by 0.8–3 

mm, ± biconcex, smooth. Leaves basal, blade narrowly 

linear, 2–30 cm by 0.5–2 mm, abruptly obtuse 

to acute, base canaliculate, glabrous; sheath up 

to 20 cm long, pale brown; ligules with a fringe of 

short hairs ca. 0.5 mm long. Involucral bracts 3–4, 

leaf-like, longest 6–13 cm long, as long as or longer 

than infl orescence. Infl orescence simple to 

compound, open, 5–10 by 2–10 cm; primary 

branches 4–8, 0.5–7 cm long. Spikelets 12–100 per 

infl orescence, usually solitary or clusters of 2, ellipsoid 

to ovoid, 4–8 by 1.5–2.5 mm. Glumes many 

per spikelet, spirally arranged, 1.8–2 mm by 0.8– 

1.2 mm, acute or mucronate, densely pubescent, 

upper part dark reddish- (rusty) brown, margin ciliate, 

1-nerved. Stamens 2; anthers 0.5–0.6 mm long. 

Stigmas 2. Nutlets obovoid, biconvex, 0.7–0.9 mm 

by 0.6–0.8 mm mature dark brown, ±smooth, with 

indistinct hexagonal or polygonal epidermal cells. 

Thailand.— NORTH-EASTERN: Maha Sarakham 

[Kantarawichai, 7 Sept. 2010, Ka. Wangwasit t & K. 

Wangwasit 028 (KKU!); Nadoon, 7 Sept. 2010, 

Ka. Wangwasit t & K. Wangwasit 042 (KKU!)]; 

EASTERN: Roi Et [Pathumrat, 7 Sept. 2010, Ka. 

Wangwasit t & K. Wangwasit 053 (KKU!)]. 

Distribution.— Southern India, Sri Lanka, 

Myanmar, Cambodia, Vietnam, northern Australia. 

Ecology.— Open wet grassy fi elds, spreading 

into paddy fi elds, on saline soil. 

Note.— Fimbristylis pubisquama Kern 

closely resembles F. sieboldii Miq. ex Franch. & 

Sav. Nevertheless, it is distinguishable from the 

latter by its annual habit, the absence of a shortly 

creeping woody rhizome and smaller nutlets. 

* Department of Biology, Faculty of Science, Mahasarakham University, Kantarawichai District, Maha Sarakham 44150, Thailand. 

Email: K_Phulphong@yahoo.com 

** Muang Phon Khon Kaen Botanic Garden,The Botanical Gardens Organization, Phon District, Khon Kaen 40120, Thailand. 

*** Applied Taxonomic Research Center, Faculty of Science, Khon Kaen University, Khon Kaen 40002, Thailand.

142 


A 

D 

G 

1 mm 

E 

F 

C 

B 

1 mm 

1 mm 

1 mm 

1 mm 

Figure 1. Fimbristylis pubisquama Kern: A. habit; B. spikelet; C. glume; D. stamen and nutlet with style and stigma; E. style and 

stigma; F. nutlet; G. nutlet surface. Scale bars = 1 mm. All from Ka. Wangwasit & K. Wangwasit 028 (KKU). Drawn by K. Wangwasit.

FIMBRISTYLIS PUBISQUAMA (CYPERACEAE), A NEW RECORD FOR THE FLORA OF THAILAND 

(K. WANGWASIT, K. WANGWASIT & P. CHANTARANOTHAI) 

An emended portion of the key to Fimbristylis in the Flora of Thailand (Simpson & Koyama, 1998) 

is proposed below. 

41. Glumes hairy at least in the apical part 

41 a. Culms usually crowded along a somewhat creeping rhizome. Leaf blades up to 10 cm long. Involucral bract usually shorter 

than infl orescence. Glumes sparsely pubescent F. sieboldii 

41 b. Culms tufted. Leaf blades up to 30 cm long. Lowest involucral bract usually overtopping the infl orescence. Glumes densely 

pubescent F. pubisquama 

41. Glumes glabrous 

Figure 2. SEM micrographs of nutlet of Fimbristylis pubisquama Kern: A. shape; B. nutlet surface. All from Ka. Wangwasit & K. 

Wangwasit 028 (KKU). 


We would like to thank Dr David A. Simpson 

for his valuable comments on manuscript preparation. 

The project was supported by the Faculty of 

Science, Mahasarakham University. 

REFERENCES 

Kern, J. H. (1955). Florae Malesianae precursors 

10. Notes on Malaysian and some S.E. Asian 

Cyperaceae 3. Blumea 8: 110–169. 

A B 

143 

Maxwell, J.F. (2002). Fimbristylis alata E.-G. 

Camus (Cyperaceae): a new record for 

Thailand. Natural History Bulletin of Siam 

Society 50(1): 115–116. 

Simpson, D. A., and Koyama, T. (1998). 

Cyperaceae. In: T. Santisuk & K. Larsen (eds), 

Flora of Thailand 6(4): 247–485. 

Shuren, Z., Songyun, L., Koyama, T. and Simpson 

D. A. (2010). Fimbristylis Vahl. In: Z.Y. Wu, 

P.H. Raven & D.Y. Hong (eds), Flora of China 

23: 200–218.

Thai Forest Bulletin

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