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
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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
THAI FOREST BULLETIN (BOTANY) 40
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.

THAI FOR. BULL. (BOT.) 40: 14–16. 2012.
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
THAI FOREST BULLETIN (BOTANY) 40
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.
Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall,
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.

THAI FOR. BULL. (BOT.) 40: 17–19. 2012.
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
THAI FOREST BULLETIN (BOTANY) 40
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
ACKNOWLEDGEMENTS
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

THAI FOR. BULL. (BOT.) 40: 20–25. 2012.
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.
KEY TO THE SPECIES
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.
THAI FOREST BULLETIN (BOTANY) 40
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

REVISION OF CYCLOCODON GRIFF. EX HOOK.F. & THOMSON (CAMPANULACEAE) (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 23
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.
ACKNOWLEDGEMENTS
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
THAI FOREST BULLETIN (BOTANY) 40
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).

REVISION OF CYCLOCODON GRIFF. EX HOOK.F. & THOMSON (CAMPANULACEAE) (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 25
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.

THAI FOR. BULL. (BOT.) 40: 26–30. 2012.
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,
THAI FOREST BULLETIN (BOTANY) 40
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
3 Green 15 Green
4 Green 16 Green
5 Green 17 Green
6 Purple 18 Green
7 Purple 19 Green
8 Green 20 Green
9 Purple 21 Green
10 Green 22 Green
11 Purple 23 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.
ACKNOWLEDGEMENTS
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.
THAI FOREST BULLETIN (BOTANY) 40
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.

THAI FOR. BULL. (BOT.) 40: 31–37. 2012.
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
THAI FOREST BULLETIN (BOTANY) 40
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)].
Distribution.— Endemic to Thailand.
Ecology.— Shaded or slightly open area
along streams or waterfalls in evergreen forest.
Flowering and fruiting; March to July.
THAI FOREST BULLETIN (BOTANY) 40
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)].
Distribution.— Endemic to Thailand.
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.
ACKNOWLEDGEMENTS
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

THREE NEW SPECIES OF PIPER (PIPERACEAE) FROM THAILAND (C. SUWANPHAKDEE, D.A. SIMPSON & P. CHANTARANOTHAI) 35
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
THAI FOREST BULLETIN (BOTANY) 40
1 cm
Figure 3. Piper smitinandianum Suwanph. & P. Chantar.: Branch with an infructescence.

THREE NEW SPECIES OF PIPER (PIPERACEAE) FROM THAILAND (C. SUWANPHAKDEE, D.A. SIMPSON & P. CHANTARANOTHAI) 37
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).

THAI FOR. BULL. (BOT.) 40: 38–56. 2012.
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.
THAI FOREST BULLETIN (BOTANY) 40
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

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 41
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
THAI FOREST BULLETIN (BOTANY) 40
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.

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 43
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
THAI FOREST BULLETIN (BOTANY) 40
KEY TO THE VARIETIES
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

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 45
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.
THAI FOREST BULLETIN (BOTANY) 40
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.

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 47
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
KEY TO THE VARIETIES
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
THAI FOREST BULLETIN (BOTANY) 40
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).

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 49
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;
THAI FOREST BULLETIN (BOTANY) 40
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
long; anthers all bearded at apex. Fruit t a capsule,
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

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 51
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
long; anthers all bearded at apex. Fruit t a capsule,
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).
Ecology.— Between 150–200 m altitude.
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
THAI FOREST BULLETIN (BOTANY) 40
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).
Ecology.— Between 2000–2100 m altitude.
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.

THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 53
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
long; anthers all bearded at apex. Fruit t a capsule,
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.
THAI FOREST BULLETIN (BOTANY) 40
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
THE LESSER-SIZED LOBELIAS OF ASIA AND MALESIA (W.J.J.O. DE WILDE & B.E.E. DUYFJES) 55
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)
dopatrioides var. cantonensis
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
dopatrioides var. cantonensis
griffi thii
heyneana
nummularia
reinwardtiana var. reinwardtiana
ACKNOWLEDGEMENTS
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
THAI FOREST BULLETIN (BOTANY) 40
zeylanica
New Guinea
archboldiana
brachyantha
conferta
dioica
donanensis
dopatrioides var. cantonensis
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.

THAI FOR. BULL. (BOT.) 40: 57–101. 2012.
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
THAI FOREST BULLETIN (BOTANY) 40
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,
THAI FOREST BULLETIN (BOTANY) 40
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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.
KEY TO THE SPECIES
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,
Malayan Forest Rec. 16: 218, t. 104, f. 2. 1943;
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
THAI FOREST BULLETIN (BOTANY) 40
Figure 4. Distribution maps of Anisoptera curtisii, A. laevis, A. scaphula and Dipterocarpus acutangulus in Peninsular Thailand-
Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors)

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
(Betong & Than To districts), Narathiwat (Sukhirin
& Waeng districts).
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.
Additional specimens.— Poopath-A80, 22
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
KEY TO THE SPECIES
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.
Additional specimens.— Poopath-A32, 15
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.
1982; Pooma & Newman, Thai For. Bull. (Bot.)
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.
Additional specimens.— Poopath-A36, 18
July 2004, Than To, Yala (BKF); Poopath-A1, 20
March 2005, Than To, Yala (BKF); Poopath-A68,10
April 2005, Than To, Yala (BKF); Poopath-A74,
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 FOREST BULLETIN (BOTANY) 40
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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.
Fruiting: July–August.
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,
Spermatophyta 9: 317. 1982; Pooma & Newman,
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),
THAI FOREST BULLETIN (BOTANY) 40
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.
Additional specimens.— Poopath-B86, 3 April
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,
Malayan Forest Rec. 16: 180, t. 84, f. 2. 1943;
Smitinand, Thai For. Bull. (Bot.) 12: 37. 1980;
Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 306.
1982; Pooma & Newman, Thai For. Bull. (Bot.)
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.
Fruiting: July–August.
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.
(Bot.) 12: 38. 1980; Ashton, Fl. Mal. Ser. I,
Spermatophyta 9: 305. 1982; Pooma & Newman,
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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.
1823; Symington, Malayan Forest Rec. 16: 186, t.
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).
KEY TO THE SPECIES
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.
Additional specimens.— Poopath-B18, 2
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
THAI FOREST BULLETIN (BOTANY) 40
Figure 6. Distribution maps of Dipterocarpus hasseltii, D.kerrii, D. retusus and Vatica bella in Peninsular Thailand-Malaysia (♦
Pooma (2003), ▲ Symington (1943) and * Authors)

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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
(Sukhirin & Waeng districts).
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.
Additional specimens.— Poopath-B82, 16
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).
Distribution.— Peninsular Malaysia. (Fig. 7).
Ecology.— Rare, frequently gregarious on
high ridges or hilltops in hill evergreen dipterocarp
forest; alt. 800–900 m. Flowering: April–July.
Fruiting: September–October.
Additional specimens.— Poopath-A107, 7
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,
Narathiwat (BKF).
Note.— A newly recorded species for
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,
Malayan Forest Rec. 16: 222, t. 108, f. 2. 1943;
Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 369.
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.
Thailand.— PENINSULAR: Narathiwat
(Sukhirin).
Distribution.— Peninsular Malaysia. (Fig. 7).
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:
223, t. 109, f. 1. 1943; Ashton, Fl. Mal. Ser. I,
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);
Poopath-B120, 12 July 2005, Sukhirin, Narathiwat
(BKF).
Note.— A newly recorded species for
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.
Thailand.— PENINSULAR: Narathiwat (Sukhirin
& Waeng districts).
THAI FOREST BULLETIN (BOTANY) 40
Distribution.— Peninsular Malaysia, Borneo
(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.
Additional specimens.— Poopath-B89, 4
April 2005, Waeng, Narathiwat (BKF); Poopath-
B101, 14 May 2005, Waeng, Narathiwat (BKF);
Poopath-B109, 9 July 2005, Waeng, Narathiwat
(BKF).
Note.— A newly recorded species for
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).

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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:
349. 1982; Pooma & Newman, Thai For. Bull.
(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).
Ecology.— Common, scattered on undulating
to ridged terrains in lowland evergreen dipterocarp
forest; alt. to ca. 300 m. Flowering: March.
Fruiting: March–October.
Additional specimens.— Poopath-B69, 11
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..
THAI FOREST BULLETIN (BOTANY) 40
— 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.
Thailand.— PENINSULAR: Yala (Bannang
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,
Narathiwat (BKF).
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;
Smitinand, Thai For. Bull. (Bot.) 12: 57. 1980;
Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 383.
1982; Pooma & Newman, Thai For. Bull. (Bot.)
29: 154, f. 6. 2001.— Shorea stellata (Kurz) Dyer
in Hook.f., Fl. Brit. Ind. 1: 304. 1874.— S. cinerea

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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,
Symington, Malayan Forest Rec. 16: 102, t. 48, f.
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,
Peninsular Malaysia. (Fig. 8).
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);
Poopath-B130, 24 Oct. 2003, Sukhirin, Narathiwat
(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:
388. 1982; Pooma & Newman, Thai For. Bull.
THAI FOREST BULLETIN (BOTANY) 40
KEY TO THE SPECIES
(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).
Distribution.— Peninsular Malaysia. (Fig. 9).
Ecology.— Rare, sporadic on ridges or hill
slopes in lowland and hill evergreen dipterocarp
forests; alt. 300–600 m. Flowering: July. Fruiting:
Janualy–April.
Additional specimens.— Poopath-B81, 15
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

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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.
Thailand.— PENINSULAR: Narathiwat
(Sukhirin).
Distribution.— Peninsular Malaysia, Borneo,
Sumatra. (Fig. 9).
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.
Thailand.— PENINSULAR: Yala (Bannang
Sata & Than To districts).
Distribution.— Peninsular Malaysia, Sumatra,
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.
Additional specimens.— Poopath-A73, 15
July 2004, Than To, Yala (BKF); Poopath-A97, 14
July 2005, Bannang Sata, Yala (BKF); Poopath-
A104, 15 July 2005, Than To, Yala (BKF);
Poopath-A131, 2 Oct. 2005, Bannang Sata, Yala
(BKF).
Note.— A newly recorded species for Thailand.
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.
(Bot.) 12: 49. 1980; Ashton, Fl. Mal. Ser. I,
Spermatophyta 9: 404. 1982; Pooma & Newman,
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).
Distribution.— Peninsular Malaysia, Borneo
(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.
Fruiting: July–August.
Additional specimens.— Poopath-A71, 21
THAI FOREST BULLETIN (BOTANY) 40
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.
Thailand.— PENINSULAR: Yala (Bannang
Sata & Betong districts).
Distribution.— Peninsular Malaysia, Borneo
(Sabah), Sumatra. (Fig. 10).
Ecology.— Very rare, gregarious on ridges in
hill and upper evergreen dipterocarp forests; alt.
750–1,000 m. Fruiting: August.
Additional specimens.— Poopath-A61, 20
Oct. 2004, Bannang Sata, Yala (BKF); Poopath-
A95, 14 July 2005, Bannang Sata, Yala (BKF).
Note.— A newly recorded species for
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;
Symington, Malayan Forest Rec. 16: 137, t. 64, f.
2. 1943; Smitinand, Thai For. Bull. (Bot.) 12: 50.
1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9:
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,

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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,
Peninsular Malaysia. (Fig. 10).
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;
Smitinand, Thai For. Bull. (Bot.) 12: 51. 1980;
Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 408.
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.
Fruiting: July.
THAI FOREST BULLETIN (BOTANY) 40
Additional specimens.— Poopath-A15, 11
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);
Poopath-B117, 11 July 2005, Sukhirin, Narathiwat
(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
(Bannang Sata, Betong, Than To districts),
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.
Additional specimens.— Poopath-A9, 16
July 2004, Than To, Yala (BKF); Poopath-A89, 25
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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).
Distribution.— Peninsular Malaysia. (Fig. 11).
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.
Additional specimens.— Poopath-A21, 16
July 2004, Than To, Yala (BKF); Poopath-A22, 14
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.
Thailand.— PENINSULAR: Narathiwat (Sukhirin
& Waeng districts).
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.
THAI FOREST BULLETIN (BOTANY) 40
KEY TO THE SPECIES
Additional specimens.— Poopath-B39, 22
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;
Smitinand, Thai For. Bull. (Bot.) 12: 57. 1980;
Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 436.
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

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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.
1943; Smitinand, Thai For. Bull. (Bot.) 12: 71.
1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9:
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).
Distribution.— Peninsular Malaysia, Sumatra.
(Fig. 11).
Ecology.— Very common, scattered or gregarious
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.
Additional specimens.— Poopath-A47, 16
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,
Narathiwat (BKF).
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).
Distribution.— Peninsular Malaysia, Singapore,
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.
Additional specimens.— Poopath-B12, 3 April
2005, Waeng, Narathiwat (BKF); Poopath-B67, 10
Sept. 2004, Waeng, Narathiwat (BKF); Poopath-
B99, 14 May 2005, Waeng, Narathiwat (BKF).
81

82
THAI FOREST BULLETIN (BOTANY) 40
Figure 11. Distribution maps of Hopea sublanceolata, Hopea sp., Shorea assamica and S. bracteolata in Peninsular Thailand-
Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors).

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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.
1925; Symington, Malayan Forest Rec. 16: 67, t.
33, f. 1. 1943; Ashton, Gard. Bull. Singapore 31(1):
48. 1978; Smitinand, Thai For. Bull. (Bot.) 12: 76.
1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9:
541. 1982; Pooma & Newman, Thai For. Bull.
(Bot.) 29: 158. 2001. Fig. 19.
Thailand.— PENINSULAR: Satun, Yala (Bannang
Sata, Than To, Betong districts), Narathiwat
(Sukhirin, Sungai Padi, Waeng districts).
Distribution.— Peninsular Malaysia, Singapore,
Borneo (Brunei, Sarawak). (Fig. 12).
Ecology.— Very common, locally frequent
on ridges or hilltops in stands in lowland and hill
evergreen dipterocarp forests; alt. 250–1,000 m.
Flowering: May–July. Fruiting: August–September.
Additional specimens.— Poopath-A17, 16
July 2004, Than To, Yala (BKF); Poopath-B26, 13
May 2005, Waeng, Narathiwat (BKF); Poopath-
B27, 8 June 2004, Waeng, Narathiwat (BKF);
Poopath-B58, 8 Sept. 2004, Sukhirin, Narathiwat
(BKF); Poopath-B112, 11 July 2005, Waeng,
Narathiwat (BKF).
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.
Thailand.— PENINSULAR: Yala (Bannang
Sata, Betong, Than To districts), Narathiwat
(Sukhirin & Waeng districts).
Distribution.— Peninsular Malaysia, Borneo
(Sabah, Sarawak), Sumatra. (Fig. 12).
Ecology.— Common, scattered on undulating
areas or ridges in lowland and hill evergreen
dipterocarp forests; altitude up to 600 m. Flowering:
April–May. Fruiting: July–August.
Additional specimens.— Poopath-A103, 15
July 2005, Than To, Yala (BKF); Poopath-B97, 13
May 2005, Waeng, Narathiwat (BKF); Poopath-
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).
Ecology.— Very common, scattered or gregarious
on ridges or hilltops in lowland to upper
evergreen dipterocarp forests; alt. 200–1,200 m.
Fruiting: October–November.
Additional specimens.— Poopath-B15, 24
Oct. 2003, Sukhirin, Narathiwat (BKF); Poopath-
B62, 9 Sept. 2004, Sukhirin, Narathiwat (BKF);
Poopath-B76, 15 Oct. 2004, Sukhirin, Narathiwat
(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.
1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9:
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
THAI FOREST BULLETIN (BOTANY) 40
Figure 12. Distribution maps of Shorea curtisii, S. faguetiana, S. gratissima and S. guiso in Peninsular Thailand-Malaysia (♦ Pooma
(2003), ▲ Symington (1943) and * Authors).

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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
(Bannang Sata, Betong, Than To districts),
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.
Additional specimens.— Poopath-A29, 14
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;
Symington, Malayan Forest Rec. 16: 75, t. 36, f. 2.
1943; Smitinand, Thai For. Bull. (Bot.) 12: 78.
1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9:
540. 1982; Pooma & Newman, Thai For. Bull.
(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).
Distribution.— Peninsular Malaysia, Singapore,
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.
Additional specimens.— Poopath-B37, 22
July 2004, Waeng, Narathiwat (BKF); Poopath-
B63, 9 Sept. 2004, Sukhirin, Narathiwat (BKF);
Poopath-B77, 15 Oct. 2004, Sukhirin, Narathiwat
(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:
56, t. 26, f. 1. 1943; Ashton, Fl. Mal. Ser. I,
Spermatophyta 9: 481. 1982. Figs. 19, 26C.
Thailand.— PENINSULAR: Narathiwat (Sukhirin
& Waeng districts).
Distribution.— Peninsular Malaysia, Borneo
(Brunei, Sarawak), Sumatra. (Fig. 13).
Ecology.— Very rare, infrequently on hill
slopes in lowland evergreen dipterocarp forest; alt.
ca. 300 m. Fruiting: July–October.
Additional specimens.— Poopath-B41, 22
July 2004, Waeng, Narathiwat (BKF); Poopath-
B45, 23 July 2004, Sukhirin, Narathiwat (BKF);
Poopath-B60, 8 Oct. 2004, Sukhirin, Narathiwat
(BKF); Poopath-B127, 27 Oct. 2005, Sukhirin,
Narathiwat (BKF).
9. Shorea macroptera Dyer in Hook. f., Fl. Brit.
Ind. 1(2): 308. 1874; Ridl., Fl. Mal. Pen. 1: 225.
1922; Symington, Malayan Forest Rec. 16: 78, t.
38, f. 1. 1943; Smitinand, Thai For. Bull. (Bot.) 12:
79. 1980; Ashton, Fl. Mal. Ser. I, Spermatophyta 9:
532. 1982; Pooma & Newman, Thai For. Bull.
(Bot.) 29: 167. 2001.— S. auriculata Scort. ex
Foxw., Malayan Forest Rec. 10: 195. 1932. Fig. 19.
Thailand.— PENINSULAR: Yala (Betong).
Distribution.— Peninsular Malaysia, Singapore,
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.
Additional specimens.— Poopath-A92, 11
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
THAI FOREST BULLETIN (BOTANY) 40
Figure 13. Distribution maps of Shorea leprosula, S. longisperma, S. macroptera and S. ochrophloia in Peninsular Thailand-
Malaysia (♦ Pooma (2003), ▲ Symington (1943) and * Authors).

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
Thailand.— PENINSULAR: Narathiwat (Waeng).
Distribution.— Peninsular Malaysia, Sumatra.
(Fig. 13).
Ecology.— Very rare, infrequently on undulating
terrain in lowland evergreen dipterocarp
fores; alt. ca. 150 m. Flowering: April. Fruiting: July.
Additional specimens.— Poopath-B44, 22
July 2004, Waeng, Narathiwat (BKF); Poopath-
B96, 5 April 2005, Waeng, Narathiwat (BKF).
Note.— A newly recorded species for
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.
Thailand.— PENINSULAR: Yala (Betong &
KEY TO THE SUBSPECIES
Than To district), Narathiwat (Sukhirin).
Distribution.— Peninsular Malaysia, Sumatra,
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.
Additional specimens.— Poopath-A20, 15
July 2004, Than To, Yala (BKF); Poopath-A59, 14
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).
Note.— A newly recorded species for Thailand.
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;
Symington, Malayan Forest Rec. 16: 85, t. 40, f. 2.
1943; Ashton, Gard. Bull. Singapore 20: 278:
1963; Man. Dipt. Brun.: 206, f. 16. 1964;
Smitinand, Thai For. Bull. (Bot.) 12: 80. 1980;
Ashton, Fl. Mal. Ser. I, Spermatophyta 9: 546.
1982; Pooma & Newman, Thai For. Bull. (Bot.)
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
(Bannang Sata, Betong, Than To districts),
87

88
Narathiwat (Cho Ai Rong, Sungai Padi, Sukhirin,
Waeng districts).
Distribution.— Peninsular Malaysia, Singapore,
Borneo (Brunei, Kalimantan, Sabah, Sarawak),
Sumatra. (Fig. 14).
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.
Additional specimens.— Poopath-A28, 13
June 2004, Than To, Yala (BKF); Poopath-B46, 23
July 2004, Sukhirin, Narathiwat (BKF); Poopath-
B64, 9 Sept. 2004, Sukhirin, Narathiwat (BKF);
Poopath-B70, 14 Oct. 2004, Sukhirin, Narathiwat
(BKF); Poopath-B78, 15 Oct. 2004, Sukhirin,
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).
Distribution.— Peninsular Malaysia, Borneo
(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.
Additional specimens.— Poopath-A41, 21
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,
THAI FOREST BULLETIN (BOTANY) 40
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.
Thailand.— PENINSULAR: Yala (Betong &
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.
Fruiting: July.
Additional specimens.— Poopath-A10, 20
June 2005, Than To, Yala (BKF); Poopath-A70, 10
April 2005, Than To, Yala (BKF); Poopath-A72,
17 May 2005, Than To, Yala (BKF); Poopath-A99,
15 July 2005, Than To, Yala (BKF).
Note.— A newly recorded species for
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:
536. 1982; Pooma & Newman, Thai For. Bull.
(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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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
Thailand.— PENINSULAR: Narathiwat (Sukhirin
& Waeng districts).
Distribution.— Peninsular Malaysia, Sumatra.
(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.
Additional specimens.— Poopath-B21, 4
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).
THAI FOREST BULLETIN (BOTANY) 40
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
Figure 16. Species of Anisoptera and Dipterocarpus in the Hala-Bala forest complex: leaves and fruits.
91

92
THAI FOREST BULLETIN (BOTANY) 40
Figure 17. Species of Dipterocarpus and Vatica in the Hala-Bala forest complex: leaves and fruits.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
Figure 18. Species of Parashorea, Neobalanocarpus and Hopea in the Hala-Bala forest complex: leaves and fruits.
sp.
93

94
THAI FOREST BULLETIN (BOTANY) 40
Figure 19. Species of Shorea in the Hala-Bala forest complex: leaves and fruits.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
Figure 20. Species of Shorea in the Hala-Bala forest complex: leaves and fruits.
95

96
THAI FOREST BULLETIN (BOTANY) 40
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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
THAI FOREST BULLETIN (BOTANY) 40
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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
THAI FOREST BULLETIN (BOTANY) 40
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.

THE DIPTEROCARPACEAE OF HALA-BALA FOREST COMPLEX, NARATHIWAT AND YALA PROVINCES, PENINSULAR THAILAND
(M. POOPATH, D. SOOKCHALOEM & T. SANTISUK)
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

THAI FOR. BULL. (BOT.) 40: 102–104. 2012.
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.
ACKNOWLEDGEMENTS
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.
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.
Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall,
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
THAI FOREST BULLETIN (BOTANY) 40
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

THAI FOR. BULL. (BOT.) 40: 105–107. 2012.
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
THAI FOREST BULLETIN (BOTANY) 40
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)].
Distribution.— Peninsular Malaysia, Sumatra,
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.
ACKNOWLEDGEMENTS
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.
IUCN. (2001). IUCN Red List Categories and
Criteria: Version 3.1. IUCN Species Survival
Commission. IUCN, Gland, Switzerland and
Cambridge, UK.
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.

THAI FOR. BULL. (BOT.) 40: 108–113. 2012.
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).
Myanmar.— Kachin State: Mountains west
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
THAI FOREST BULLETIN (BOTANY) 40
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.

NEW RECORDS IN THE ORCHID FLORA OF MYANMAR (H. KURZWEIL & S. LWIN) 111
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).
Myanmar.— Kachin State: Mountains west
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
THAI FOREST BULLETIN (BOTANY) 40
Figure 2. A. Cleisostoma linearilobatum (Seidenf. & Smitinand) Garay, fl ower. Kurzweil & Lwin 2772; B–C. Panisea tricallosa
Rolfe. Kurzweil & Lwin 2573.

NEW RECORDS IN THE ORCHID FLORA OF MYANMAR (H. KURZWEIL & S. LWIN) 113
ACKNOWLEDGEMENTS
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.

THAI FOR. BULL. (BOT.) 40: 114–117. 2012.
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.
ACKNOWLEDGEMENTS
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
THAI FOREST BULLETIN (BOTANY) 40
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.

THAI FOR. BULL. (BOT.) 40: 118–120. 2012.
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.
ACKNOWLEDGEMENTS
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
THAI FOREST BULLETIN (BOTANY) 40
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.

THAI FOR. BULL. (BOT.) 40: 121–124. 2012.
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
THAI FOREST BULLETIN (BOTANY) 40
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].
IUCN. (2001). IUCN Red List Categories and
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.
THAI FOREST BULLETIN (BOTANY) 40
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.

THAI FOR. BULL. (BOT.) 40: 125–129. 2012.
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
Thailand.
Ecology.— Epiphyte on trees and lithophyte
on moss-covered sandstone rock platforms in montane
forest, 1100–2100 m altitude. Flowering:
August–September.
THAI FOREST BULLETIN (BOTANY) 40
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).
ACKNOWLEDGEMENTS
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
THAI FOREST BULLETIN (BOTANY) 40
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.

THAI FOR. BULL. (BOT.) 40: 130–133. 2012.
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.
ACKNOWLEDGEMENTS
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
THAI FOREST BULLETIN (BOTANY) 40
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

THAI FOR. BULL. (BOT.) 40: 134–140. 2012.
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,
Thailand.
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
THAI FOREST BULLETIN (BOTANY) 40
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.
ACKNOWLEDGEMENTS
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,

TWO NEW RECORDS OF XYRIS L. (XYRIDACEAE) FOR THAILAND (P. PHONSENA, P. CHANTARANOTHAI & A. MEESAWAT) 137
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
Commission. IUCN, Gland, Switzerland and
Cambridge, UK.
Van Royen, P. (1954). Some new Australasian species
of Xyris. Blumea 7: 477. 1954.

138
A
1 cm
B
THAI FOREST BULLETIN (BOTANY) 40
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
TWO NEW RECORDS OF XYRIS L. (XYRIDACEAE) FOR THAILAND (P. PHONSENA, P. CHANTARANOTHAI & A. MEESAWAT) 139
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

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THAI FOREST BULLETIN (BOTANY) 40
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).

THAI FOR. BULL. (BOT.) 40: 141–143. 2012.
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.

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THAI FOREST BULLETIN (BOTANY) 40
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).
ACKNOWLEDGEMENTS
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.