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Cambodian Journal of Natural History Volume 2014 Issue 1

  • Re:wild and Fauna & Flora International
August 2014 Vol. 2014 No. 1
Aquatic Special Issue:
Dragon ies and damsel ies
New crabs discovered as by-catch
Seagrasses of Koh Rong Archipelago
Koh Sdach Archipelago coral reef survey
Zoning Cambodia’s rst Marine Fisheries
Management Area
Cambodian Journal
of Natural History
Cambodian Journal of Natural History
ISSN 2226–969X
Dr Jenny C. Daltry, Senior Conservation Biologist, Fauna & Flora International.
Dr Neil M. Furey, Research Associate, Fauna & Flora International: Cambodia Programme.
• Hang Chanthon, Former Vice-Rector, Royal University of Phnom Penh.
Dr Nicholas J. Souter, Project Manager, University Capacity Building Project, Fauna & Flora International: Cambodia
Dr Stephen J. Browne, Fauna & Flora International,
Dr Martin Fisher, Editor of Oryx—The International
Journal of Conservation, Cambridge, United Kingdom.
Dr L. Lee Grismer, La Sierra University, California,
Dr Knud E. Heller, Nykøbing Falster Zoo, Denmark.
Dr Sovanmoly Hul, Muséum National d’Histoire
Naturelle, Paris, France.
Dr Andy L. Maxwell, World Wide Fund for Nature,
Dr Jörg Menzel, University of Bonn, Germany.
Dr Brad Pe i , Murdoch University, Australia.
Dr Campbell O. Webb, Harvard University Herbaria,
Other peer reviewers for this volume
The Cambodian Journal of Natural History (ISSN 2226–969X) is an Open Access journal published by the Centre
for Biodiversity Conservation, Royal University of Phnom Penh. The Centre for Biodiversity Conservation is
a non-pro t making unit dedicated to training Cambodian biologists and to the study and conservation of
Cambodia’s biodiversity.
Dr Shane T. Ahyong, Australian Museum Research
Institute, Sydney, Australia.
Dr Alexander E. Balakirev, Severtsov’s Institute of
Ecology and Evolution of RAS, Moscow, Russia.
Jan-Willem van Bochove, UNEP World Conservation
Monitoring Centre, Cambridge, UK.
Dr Stephen Bortone, Osprey Aquatic Sciences, Inc.,
Tampa, Florida, USA.
Dr Jackson Freche e, Fauna & Flora International,
Phnom Penh, Cambodia.
Dr Hedley Grantham, Conservation International,
Arlington, Virginia, USA.
Dr Peter Houk, University of Guam Marine
Laboratory, Mangilao, Guam, USA.
Dr Kathe Jensen, Zoological Museum, Copenhagen,
Dr Luke Leung, School of Agriculture and Food
Sciences, University of Queensland, Australia.
Prof. Colin L. McLay, Canterbury University,
Christchurch, New Zealand.
• Neang Thy, GDANCP, Ministry of Environment,
Phnom Penh, Cambodia.
Dr Niphon Phongsuwan, Phuket Marine Biological
Center, Phuket, Thailand.
Dr Carly Starr, Northern Gulf Resource Management
Group, Mareeba, Queensland, Australia.
Keith D.P. Wilson, Brighton, UK.
Cover photo: A sea whip (order Alcyonacea) near Koh Bong in the Song Saa Marine Reserve (© Jelena Vukosav-
ljevic, Song Saa Private Island). The reserve is within the proposed Koh Rong Archipelago Marine Fisheries
Management Area, the focus of papers in this issue led by Cambodian scholars Leng Phalla and Boon Pei Ya.
International Editorial Board
© Centre for Biodiversity Conservation, Phnom PenhCambodian Journal of Natural History 2014 (1) 13
Guest Editorial—Lots of information collected about marine living
resources, but where is it? And can it be trusted?
Kathe R. JENSEN¹ and ING Try²
1 Zoological Museum (Natural History Museum of Denmark), Universitetsparken 15, DK-2100 Copenhagen, Denmark.
2 Fisheries Administration, Ministry of Agriculture, Forestry and Fisheries, Preah Norodom Boulevard 186, P.O. Box
582, Phnom Penh, Cambodia. Email: or
“Out of sight—out of mind”, is an excuse that has
often been used to explain why so li le is known
about biodiversity in the sea. With the exception of
what washes up on the beach and what can be seen in
the sh market, marine biodiversity remains largely
hidden from human eyes. This means we have to
remind ourselves—or be reminded—that marine
biodiversity needs our a ention, care and protection,
just like that of forests, mountains and wetlands.
Cambodia has one of the world’s largest inland
sheries and naturally the focus of international
organizations, as well as the national government,
has been largely on the conservation and sustain-
able use of freshwater sheries resources. The unique
hydrological features associated with the seasonal
reversal of ow to and from the Tonle Sap Lake have
intrigued scientists from all over the world, and there
is great concern about the potential damage caused
by constructing hydro-electric power plants across
the Lower Mekong water course (see e.g. Hogan et
al., 2004; Campbell et al., 2006; Baran & Myschowoda,
Much less a ention has been given to the marine
biodiversity of Cambodia. Marine capture sheries,
though increasing, remain less than 20% the size of
the inland sheries, and hence fewer resources have
been allocated to this eld. Over the years, numerous
international projects have operated in the Cambo-
dian coastal zone, and much information has been
gathered about vulnerable ecosystems, the socio-
economics of coastal residents, and the impacts of
legislation relating to environmental degradation and
use of sheries resources.
One of the largest projects, usually called the
UNEP South China Sea Project for short, involved
seven countries surrounding the South China Sea,
over 16 million dollars in direct funding, and about
the same amount in local “in-kind” contributions, and
lasted from 2002 through 2008 (Vo & Perne a, 2010).
The results from this project have entered the inter-
national scienti c literature (e.g. Vo et al., 2013), and
provided government agencies with national reports,
trained sta members, established demonstration
sites for the general public (e.g. UNEP, 2007) and
published posters and booklets on various ecosystems
(e.g. UNEP, 2004).
For Cambodia, this project funded assessments of
coral reef, seagrass and mangrove habitats. We now
have a fair knowledge of the size of these vulnerable
ecosystems and, equally importantly, where they are
located. This information constituted the baseline
information for developing a National Action Plan
for Coral Reef and Seagrass Management in Cambodia
2006–2015 (FiA, 2006). The project also funded the
publication of a bilingual eld guide to the marine
living resources in Cambodia, which unfortunately
was printed in far too few copies (Ing et al., 2006).
Smaller projects have taken over and carry out Reef
Check and Seagrass Watch procedures to monitor the
state of selected reefs and seagrass beds. Some of these
projects are carried out by local Cambodian govern-
ment employees, but most use foreign volunteers,
often marine biology students. Few Cambodians
have diving skills or can a ord the equipment and
other associated costs. However, dive shops exist for
tourists, and the sta and boats from these shops have
been involved in several projects.
Every NGO has its own “mission and vision”,
but this may seem like semantics when applying for
foreign grants: you put your pen where the money
CITATION: Jensen, K.R. & Ing T. (2014) Lots of information collected about marine living resources, but where is it? And can
it be trusted? Cambodian Journal of Natural History, 2014, 1–3.
© Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2014 (1) 13
is. If the donor wants you to include gender issues,
you include them in your application; if they focus
on sustainability, you make this a priority goal; and
so on. One project may be speci cally dedicated to
seahorses—small threatened sh used in traditional
medicine. Another project may focus on the economic
costs and bene ts of tourism in coral reef areas.
Other projects may focus on socioeconomics and the
development of small-scale sheries. In spite of this
diversity in aims and goals, there is a lot of overlap
in the reports that result from these projects. Many
socioeconomic questionnaires containing basically
the same questions are presented to basically the same
people by di erent projects. Similarly, the same coral
reef may be surveyed by transects, manta-tows and
random quadrat photography. Unfortunately there
are sometimes con icting results among reports.
The information gathered from this multitude
of projects is valuable and useful, but often di cult
to locate. In many cases, project teams write status
reports for their funding bodies only. Most nish their
projects with impressive reports on glossy paper, but
rarely with an ISBN identi cation number, and rarely
have these reports been subject to peer-review. Some
information is available in Environmental Impact
Assessment reports in connection with major devel-
opment projects, usually in the private sector, but EIA
reports are often inaccessible to the public, including
scientists, because they are considered the private
property of the company. Other information is avail-
able only in Khmer-language internal government
reports, such as most sheries statistics. There is a
great need for a comprehensive review of the avail-
able data, and to improve the quality and reliability of
the data from di erent studies.
The sta of government agencies, both national
and local, try their best to keep up with all the ndings
from the various projects, but it often looks as if legis-
lation is drafted, implemented and enforced at the
same time as the eld data are collected. One could
say that Cambodia implements the holistic, integrated
approach from need rather than choice, and “citizen
science” was in use here years before it became
fashionable in western countries (after the economic
crisis drastically cut funding for science).
Government sta depend on scienti c information
to draft meaningful and e ective regulations. Cambo-
dian sheries o cers, however, usually have to
collect eld data at the same time they draft manage-
ment measures, such as the National Plan of Action
for Sharks (Ing et al., 2004). Fortunately international
organizations provide guidelines, but data still have
to be collected. Although an increasing number of
Cambodian ministerial o cers have received upper
level academic degrees (MSc and PhD), they rarely
get the opportunity to be involved in basic research
or participate in research publications. There is a need
for collaboration between universities and govern-
ment agencies that goes beyond supervising graduate
thesis projects.
Cambodia is a signatory party to several interna-
tional conventions related to marine biodiversity (Nao
et al., 2013). This means that Cambodia is expected to
participate in meetings, submit national reports, and
most importantly, comply with international regula-
tions. This has occasionally led to awkward situations.
At one time, Cambodia was urged to take measures
against illegal
shing vessels, which were shing
tuna in the Atlantic Ocean. Obviously no Cambo-
dian shermen own vessels capable of shing tuna
in the Atlantic, but somehow these vessels had been
registered in Cambodia and were ying the ag of
Cambodia, but without a Cambodian license to sh.
Recently, the European Union imposed an embargo
on sheries goods from Cambodia because “despite
several warnings” the country had not taken su -
cient measures to prevent illegal, unreported and
unregulated (IUU) shing. Cambodia is working on
a National Action Plan to prevent, deter and eliminate
illegal, unreported and unregulated shing (NPOA-
IUU shing), but this takes time and requires funding.
Cambodia has four nationally protected areas in
the coastal zone: Three National Parks (Bokor, Kep
and Ream), and one Wildlife Sanctuary (Peam Krasop,
part of which—Koh Kapik—is also a Ramsar Wetland
of International Importance). These are all land-based
but have a marine component. In addition, the UNEP
South China Sea Project introduced a new concept
for conserving critical habitats of certain species—
Fisheries Refugia—and the South-East Asian Fisheries
Development Center (SEAFDEC) assisted Cambodia
to implement marine refugia (Ing et al., 2010). A
real Marine Protected Area, however, has yet to be
formally established in Cambodia. The underwater
area surrounding the islands of the Koh Rong Archi-
pelago has been mentioned several times, and many
reef areas have been surveyed again and again. In spite
of all the information available, however, con icting
interests have so far stalled the formal establishment
of a Marine Fisheries Management Area.
One of the latest government targets of the
Strategic Planning Framework for Fisheries 2010–2019, is
to increase aquaculture production by 15% annually.
One can only guess at the impacts to vulnerable
© Centre for Biodiversity Conservation, Phnom PenhCambodian Journal of Natural History 2014 (1) 13
coastal environments of such an immense enterprise.
Once again, it appears that research and policy devel-
opment are carried out simultaneously with cu ing
mangroves and digging ponds. Moreover, this is
happening at a time when most western countries are
trying to ban imports of cultured sh and shrimps
from countries that use wild-caught trash- sh for
In Cambodia, conservation issues, which are often
supported by small NGOs, have to compete with
development projects, which are often supported
by wealthy private companies. Unfortunately, the
economy usually wins over ecology. The coastal
marine waters of Cambodia house a diverse and
possibly unique marine life, and it is important
that this resource be protected for future genera-
tions (as well as to a ract tourists). Having multiple
NGO-supported projects without much coordination
of their e orts has created a lot of information, but has
had li le or no lasting e ect on the conservation or
management of marine resources. The action plans
remain plans without action.
It is time for someone in Cambodia to review
the existing information and extract data from avail-
able reports that might be used for knowledge-based
selection of sites to be protected. Also, NGO and inter-
national projects should be required to make their
nal results publicly available at a common website
repository in Cambodia. The existing websites that
contain some of this information should be urged to
collaborate. The conservation of Cambodia’s marine
biodiversity is too important to end up as a pile of
glossy paper reports on bookshelves in government
o ces.
Baran, E. & Myschowoda, C. (2009) Dams and sheries in
the Mekong Basin. Aquatic Ecosystem Health & Manage-
ment, 12, 227–234.
Campbell, I.C., Poole, C., Giesen, W. & Valbo-Jørgensen, J.
(2006) Species diversity and ecology of Tonle Sap Great
Lake. Aquatic Sciences, 68, 355–373.
FiA—Fisheries Administration (2006) National Action Plan for
Coral Reef and Seagrass Management in Cambodia 2006–2015.
Ministry of Agriculture, Forestry and Fisheries, Fisheries
Administration, Phnom Penh, Cambodia.
Hogan, Z.S., Moyle, P.B., May, B., Vander Zanden, M.J. &
Baird, I.G. (2004) The imperiled giants of the Mekong.
American Scientist, 92, 228–237.
Ing T., Jensen, K.R., Pich S. & Va L. (2004) Shark and ray
sheries in Cambodia: a review of national management
activities. Fish for the People, 2, 24–31.
Ing T., Jensen, K.R. & Va L. (2006) Field Guide to the Marine
Living Resources of Cambodia. Fisheries Administration,
Phnom Penh, Cambodia.
Ing T., Etoh, S. & Sornkliang, J. (2010) The role of shers’
group in the establishment and management of a refugia
system: experience of Cambodia. Fish for the People, 8, 32.
Nao T., Ing T. & Jensen, K.R. (2013) Relevant international
and regional instruments for sustainable development of
small-scale marine sheries: signi cance to Cambodia.
Fish for the People, 11, 24–31.
UNEP—United Nations Environment Programme (2004)
Coral Reefs in the South China Sea. UNEP/GEF/SCS
Technical Publication No. 2., United Nations Environment
Programme, Bangkok, Thailand.
UNEP—United Nations Environment Programme (2007)
Seagrass Demonstration Sites in the South China Sea. UNEP/
GEF/SCS Technical Publication No. 6., United Nations
Environment Programme, Bangkok, Thailand.
Vo, S.T. & Perne a, J.C. (2010) The UNEP/GEF South China
Sea Project: lessons learnt in regional cooperation. Ocean &
Coastal Management, 53, 589–596.
Vo, S.T., Perne a, J.C. & Paterson, C.J. (2013) Status and
trends in coastal habitats of the South China Sea. Ocean &
Coastal Management, 85, 153–163.
© Centre for Biodiversity Conservation, Phnom Penh
F. Chassagne & S. Hul
Cambodian Journal of Natural History 2014 (1) 47
Short Communication
A range extension for the new plant species
Solanum sakhanii
Hul, and its medicinal uses in a Bunong community in Mondulkiri
François CHASSAGNE1,* and Sovanmoly HUL2
1 Nomad RSI Organization, Doh Kromom Village, Sokhadom Commune, Senmonorom District, Mondulkiri Province,
2 Muséum national d’Histoire naturelle (MNHN), Département de Systématique et Évolution, UMR 7205 (ISYEB),
Herbier national (P), CP 39, 57 rue Cuvier, 75231 Paris Cedex 5, France.
*Corresponding author. Email
Paper submitted 4 June 2014, revised manuscript accepted 17 July 2014.
CITATION: Chassagne, F. & Hul S. (2014) A range extension for the new plant species Solanum sakhanii Hul, and its medicinal
uses in a Bunong community in Mondulkiri Province. Cambodian Journal of Natural History, 2014, 4–7.
Thanks to its wide range of ecosystems (Tan, 1952),
including evergreen forests, deciduous dry forests,
grasslands, mangroves, coral reefs, swamp forests,
seagrass meadows, and wetlands, Cambodia harbours
a rich oristic diversity. The country is home to about
2,308 known vascular plant species belonging to 852
genera in 164 families, including 214 endemic species
(Dy Phon, 1982; Ministry of Environment, 2010). In the
1990s, Ashwell (1997) estimated that with a complete
and systematic study, the ora would exceed 3,000
species. The total number of vascular plants remains
unknown, but we now estimate that Cambodia has
as many as 5,000 vascular plant species (Hul, unpub-
lished data).
Cambodia’s native ora has fascinated scien-
tists through the ages (e.g. Loureiro, 1790; Pierre,
1879–1899; Lecomte, 1907–1951; Aubréville, 1960
onwards; Vidal et al., 1969; Martin, 1971; Matras &
Martin, 1972), and botanists, ethnobotanists and
biologists continue to take a keen interest in it to
this day. With research over the past ve years, the
number of plant species recorded in Cambodia has
increased, and 20 new vascular plants species have
been described: Acampe hulae Telepova (Telepova,
2009), Aeschynanthus cambodiensis D.J. Middleton
(Middleton, 2009), Alocasia jiewhoei V.D. Nguyen
(Nguyen et al., 2010), Amorphophallus hemicrypticus
He erscheid (Maxwell, 2009), Areca riparia Heatubun
(Heatubun, 2011), Ceratopteris oblongiloba Masuyama
& Watano (Masuyama & Watano, 2010), Curculigo
fabrei Hul (Leti et al., 2013), Dasymaschalon acuminatum
Wang & Saunders (Wang et al., 2009), Doritis boulbetii
Telepova (Telepova, 2013), Doritis pulcherrima f. cinna-
barina Telepova (Telepova, 2013), Exacum darae Hul
(Hul, 2010), Lagerstroemia kratiensis W.J. de Wilde &
Duy es (de Wilde & Duy es, 2013), Lespedeza cambodi-
anum V.D. Nguyen (de Kok & Nguyen, 2013), Miliusa
cambodgensis Chaowasku & Kessler (Chaowasku &
Keßler, 2014), Nepenthes bokorensis Mey (Mey, 2009),
Nepenthes holdenii Mey (Mey et al., 2010), Peliosanthes
cambodiana Aver. & N. Tanaka (Averyanov et al., 2013),
Peliosanthes triandra Aver. & N. Tanaka (Averyanov et
al., 2014), Solanum sakhanii Hul (Leti et al., 2013) and
Syzygium bokorense W.K. Soh & J. Parn. (Soh & Parnell,
These ndings have contributed to a be er under-
standing of the country’s ora, but suggest that many
species remain to be discovered.
In Northeast Cambodia, near the Vietnamese
border, lies the province of Mondulkiri. This province
has been studied primarily by zoologists, whose
research has proved that the area’s wide range of
habitats supports an exceptionally high diversity of
fauna (Walston et al., 2001; Rawson et al., 2009; Phan
et al., 2010). Although its rich diversity of habitats also
© Centre for Biodiversity Conservation, Phnom Penh
Range extension of a new medicinal plant
Cambodian Journal of Natural History 2014 (1) 47
makes Mondulkiri an extremely interesting subject
for botanists, only a handful of surveys have been
conducted thus far (Jung et al., 2014).
The province’s largest ethnic group is the indig-
enous Bunong, one of the highland minorities who
are considered to be among the rst inhabitants
of Cambodia. Over time, the Bunong people have
developed a strong relationship with their natural
environment (Bourdier, 2006). They are known for
their extensive knowledge of plants, which they use
for various purposes, such as medicine, food, building
and hunting (Laval et al., 2011). Their local knowledge
has already been advantageous for ecological assess-
ments of mammal species (Starr et al., 2011) and we
predicted this could also be used as a basis for botan-
ical eld studies, potentially helping to uncover new
plant species.
The results of an ethnobotanical survey carried
out in 2013 and 2014 with the Bunong people have
enhanced our understanding of Mondulkiri’s ora
(Chassagne, in prep.). It is thanks to the knowledge of
the Bunong that we discovered the newly described
plant species Solanum sakhanii Hul during our investi-
gation. The villagers of Dak Dam Commune (O’Reang
District) have long known and used this plant as a
traditional treatment for malaria and fevers. The plant
grows on dry rice elds during the rainy season. The
Bunong people collect the whole plant and cut it into
small pieces, which are boiled for 15 minutes. The
patient drinks this remedy four to six times per day
until his condition has improved. It is also possible to
make a steam bath with the boiling water.
This is the rst paper to describe the plant Solanum
sakhanii Hul as having medicinal uses, and also
con rms the rst record in Mondulkiri, and only the
second record of this species in Cambodia and South-
east Asia.
Solanum sakhanii Hul is a small herb that was rst
discovered in 2008 and published in 2013 by Leti et
al. (Chéng, Hul & David CL 929; holotype, P00836398;
isotypes, P00836399, P00836400) near Kbal Chhay
in Preah Sihanouk Province (10°39’23”N, 103°36’
57”E, elevation 60 m) (Figs 1 and 2). It is a creeping,
stoloniferous plant, which grows in open, humid
environments or in the understory of dense forests.
Other characteristics are listed as follows: Leaves are
solitary or unequally paired; lamina ovate-elliptic,
4–7 × 2.5–4.0 cm, margin 3–5-sinuate lobed, pubescent
with stellate hairs adaxially and abaxially, prickly
along main veins on both surfaces. In orescences
extra-axillary, scorpioid-racemose, 1–4- owered.
Flowers 5-merous, 10–12 mm long; pedicel pubescent
with stellate hairs, prickly. Calyx of 5 sepals, pubes-
cent with stellate hairs, prickly abaxially. Corolla of 5
petals, pale blue or purplish, pubescent with stellate
hairs abaxially. Stamens 5, laments 1–1.5 mm long;
anthers narrow. Fruits: berries globose or subglo-
bose, 1–1.5 cm in diameter, glabrous, ± included in the
accrescent calyx, greenish-white to yellow.
We collected Solanum sakhanii Hul (Chassagne
138, P00723707) on 27 October 2013. This species was
found to be very common in an upland rice eld on
red clay soil, in a disturbed area near an evergreen
forest, in Dak Dam Commune, O’Reang District,
Mondulkiri Province (12°24’37”N, 107°19’25”E, eleva-
tion 785 m) (Fig. 1). Specimens have been deposited
in the National Herbarium of Cambodia (RUPP) in
the Department of Biology, Faculty of Science, Royal
University of Phnom Penh, and in the Paris Herbarium
in France.
This record in Mondulkiri Province, in a di erent
habitat (with red clay soil rather than sandy soil) and
at a much higher altitude (785 m) than the type locality
in Preah Sihanouk Province, indicates that the species
Solanum sakhanii could be widely distributed. Further
investigations in neighbouring countries, especially
Vietnam and Laos, could be helpful to assess its
presence in these countries. Moreover, the discovery
that this plant is considered by the Bunong as having
medicinal uses could be used for future phytochem-
ical and pharmaceutical investigations.
Fig. 1 Cambodia, showing the locations of the two
records of Solanum sakhanii Hul.
© Centre for Biodiversity Conservation, Phnom Penh
F. Chassagne & S. Hul
Cambodian Journal of Natural History 2014 (1) 47
The authors would like to thank the Bunong villagers
in Dak Dam Commune for sharing their knowledge
with us, and Moeub Chamroeun for helping trans-
late this information. We are grateful to the sta of
Nomad RSI Cambodia for their participation, and
Bruno David (Pierre Fabre Institute) for his photo-
graphs. We sincerely thank Edwinstaël Ramanantsoa
and Élodie Lerat (Global Plants Initiative, MNHN).
Finally, we would like to thank Svjetlana Bosanac for
proofreading this article.
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Fig. 2 Solanum sakhanii Hul, in its type locality in Preah Sihanouk Province (© B. David).
© Centre for Biodiversity Conservation, Phnom Penh
Range extension of a new medicinal plant
Cambodian Journal of Natural History 2014 (1) 47
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About the authors
FRANÇOIS CHASSAGNE is a pharmacist and
holds a Master’s degree in Biodiversity, Ecology and
Evolution. He has acquired special skills in ethnop-
harmacology and conducted ethnobotanical surveys
in Southeast Asia. He organised and led this study in
Mondulkiri Province.
SOVANMOLY HUL is a professional botanist at the
National Museum of Natural History in Paris. She is
an associate professor, editor-in-chief of the Flore du
Cambodge, du Laos et du Viêtnam and curator for Genti-
anales and Solanales at the Paris Herbarium. She is
a co-author of the Flore Photographique du Cambodge,
published by Editions Privat in 2013.
© Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2014 (1) 8–11
O.E. Kosterin & G. Chartier
Short Communication
Two more Odonata species recorded for Cambodia
Oleg E. KOSTERIN1,2,* and Gerard CHARTIER3
1 Institute of Cytology & Genetics SB RAS, Acad. Lavrentyev ave. 10, Novosibirsk, 630090, Russia.
2 Novosibirsk State University, Pirogova str. 2, Novosibirsk, 630090, Russia.
3 Rainbow Lodge, Tatai, Koh Kong, Cambodia.
*Corresponding author. Email
Paper submitted 9 November 2013, revised manuscript accepted 30 June 2014.
CITATION: Kosterin, O.E. & Chartier, G. (2014) Two more Odonata species recorded for Cambodia. Cambodian Journal of
Natural History, 2014, 8–11.
A checklist of Odonata species recorded in Cambodia
was published in the December 2012 issue of the
Cambodian Journal of Natural History (Kosterin et al.,
2012a), but was estimated to comprise only about
a half of the country’s actual species. The paper
included dragon y and damsel y species registered
in the Cardamom Mountains foothills—speci cally in
Tatai Commune—by the resident second author over
a number of years. Since that publication, the second
author has recorded two more species of Odonata that
are new records for Cambodia. They are published
herein, along with additional new data from the same
Gerard Chartier has compiled photographic
records of Odonata using a Canon EOS 500D camera
with 18–55 mm and 55–250 mm lenses during his
regular excursions in Tatai Commune, Koh Kong
Province, within the area encompassed by the
following points: Rainbow Lodge on the Stoeng Kep
River (“Left Tatai River”), 1.6 km north of Phum Doung
Bridge in Tatai Village (11.580˚N, 103.127˚E), the “lake
area” of the Stoeng Sala Munthun River (“Right Tatai
River”), 3.9 km north-northwest of Phum Doung
Bridge (11.599–11.601˚N, 103.120–121˚E), and the Tatai
Waterfall at the same river, 4 km northwest of Phum
Doung Bridge (11.586˚N, 103.097˚E). This area is
covered by evergreen forest and bamboo thickets over
a low ridge. For more details of the area see Kosterin et
al. (2012a). Two odonate individuals were found dead
in a house at Rainbow Lodge and preserved dry in a
paper envelope.
Heliaeschna simplicia (Karsch, 1891)
A male was found dead inside a house at Rainbow
Lodge (11.580˚N, 103.127˚E) on 12 March 2013 (Fig. 1),
seemingly after being a racted by light.
There used to be taxonomical and nomenclatural
confusion around this species, but this was resolved
by Lieftinck (1940). The identi cation of our male was
unmistakable because of the shape of its cerci (Fig. 2),
which are unique for H. simplicia (Karsch, 1891) (see
also Fig. 161 for “H. Weelei” in Martin, 1909; Figs 22–23
for “H. Weelei” in Ris, 1927; and plate 14h in Orr, 2003).
Other characters matched as well.
Until now, H. simplicia was known only from
Borneo, Sumatra (Lieftinck, 1940; 1954), the Philip-
pines (Hämäläinen & Müller, 1997; Tsuda, 2000) and
Peninsular Malaysia (Orr, 2008). Ours is the rst
published record of this species in Indochina and in
continental (not peninsular or insular) Asia.
Epophthalmia vi igera (Rambur, 1842) ssp. bellicosa
Lieftinck, 1948.
A female was photographed by Gerard Chartier at
Rainbow Lodge (11.580 N, 103.127 E) on 14 June
2013 (Fig. 3a) and a male on 18 June 2013 (Fig. 3b).
Another female (Fig. 4) was found dead inside a house
at Rainbow Lodge on 23 June 2013. More males were
observed at the same place every day until at least 30
June 2013.
© Centre for Biodiversity Conservation, Phnom Penh
Cambodian Journal of Natural History 2014 (1) 8–11
New records of dragon ies
Identi cation of this species was reliably based on
the extensive dark patches at the wing bases in both
sexes, the strong subapical inward angulation of the
male cerci and the epiproct of the same length, and
vertex tubercles without yellow markings (Fraser,
1936; Lieftinck, 1948).
Fig. 1 Heliaeschna simplicia (Karsch, 1891), a male found
dead in Rainbow Lodge, Tatai Commune, Koh Kong
Province, 12 March 2013 (© G. Chartier).
Fig. 2 Heliaeschna simplicia (Karsch, 1891), anal append-
ages of the same male (© O. Kosterin).
Fig. 3 Epophthalmia vi igera bellicosa Lieftinck, 1948, a
female (a) and male (b) photographed at Rainbow Lodge,
Tatai Commune, Koh Kong Province on 14 and 18 June
2013 respectively (© G. Chartier).
Fig. 4 Epophthalmia vi igera bellicosa Lieftinck, 1948, details
of a female found at Rainbow Lodge, Tatai Commune,
Koh Kong Province, 23 June 2013 (© G. Chartier).
© Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2014 (1) 8–11
O.E. Kosterin & G. Chartier
As to the subspeci c diagnostic characters, the
basal brown patches extended to antenodal 5 and 6 on
the forewing and hindwing, respectively, in the female,
and were vestigal on the forewing and extended to
antenodal 1 in the male, which were within the range
of variation indicated for the subspecies bellicosa Lieft-
inck, 1948 by Fraser (1936) and Lieftinck (1948). This
subspecies is known to range across India, Myanmar,
Thailand, Laos and Vietnam (Tsuda, 2000). The yellow
rings on abdominal segments 2–7 were not as broad
as indicated in the original description (Lieftinck,
1948) and interrupted in segments 2–6, but were
within the variation indicated by Fraser (1936); their
fore margins being indistinct. In the female specimen,
the postclypeus missed the yellow transversal stripe
mentioned by Fraser (1936) and Lieftinck (1948)
and was “uniformly brownish” as characterised by
Asahina (1987). Generally, the male and two females
gave the impression of being aged and darkened.
Epophthalmia vi igera bellicosa was expected to
occur in Cambodia, having been found in Chaya-
phum and Saraburi Provinces of Central Thailand
(Hämäläinen & Pinratana, 1999).
Macromia chaiyaphumensis Hämäläinen 1985.
Two males photographed on 11 October 2011 and 19
September 2012 in the study area were not immediately
identi ed to species, being referred to as “Macromia
sp.” and illustrated by Kosterin et al. (2012a) in Fig.
12 (showing the male from 19 September 2012 but
erroneously captioned “April 2012”). They showed a
yellow mark between the vertex pyramidal processes.
Of the known Thai and Indochinese species, this trait
is found only in M. chaiyaphumensis. The original
description, based on a single male, states: “Posterio-
medial slopes of pyramidal processes in frons yellow”
(Hämäläinen 1985: 105), although its author did not
indicate this character as diagnostic. Photographs of
a male and female identi ed as M. chaiyaphumensis
were uploaded by Reinthong Ruanrong to www.
asia-dragon on 18 September 2013 (use the
‘Global query’ to nd these records of this species).
Ruanrong’s male and female were photographed on
24 May 2013 and 20 April 2013, respectively, at the
Borwee Waterfall, Sounphung, Ratchaburi Province,
Thailand. They showed the aforementioned yellow
mark in the frontal crevice very clearly and were
obviously conspeci c.
Our males from Cambodia corresponded to the
male photographed by R. Ruanrong in most details,
but with the following di erences: yellow ring at
abdominal segment 2 interrupted at auricles, anterio-
lateral yellow spots on S3 large (vs very small in the
Ratchaburi male), dorsal and lateral yellow marks
on abdominal segment 8 separated, yellow lateral
spots and lateroventral dots on abdominal segment
9 absent, one less antenodal on each wing (12 on
forewing and nine on hindwing vs 13 on forewing and
10 on hindwing in the Ratchaburi male). The pinkish-
grey eyes and brownish-black thoracic ground colour
in the Tatai males may be ascribed to their subteneral
The males from the Tatai Commune and Ratch-
aburi Province show some common di erences
from the original description of M. chaiyaphumensis
(Hämäläinen 1985: 105), as follows:
• Abdominal segment 3 with lateroventral spots
(small in the Ratchaburi male and large in the Tatai
males), not mentioned in the original description;
Yellow marking on segments 3–7 seems to be larger
(not depicted in loc. cit.), spots on segment 6 not
“very small” but occupy half of segment height,
mark on segment 7, however, occupies apical
one- fth rather than one-third of its segment;
Fewer antenodals on forewing (vs 14 on forewing
and nine on hindwing in the holotype);
• Two crossveins in hindwing hypertrigones (vs
three in the holotype).
In spite of these not-so-large di erences, which may
be a ma er of geographical variation, we provision-
ally identi ed the Tatai males as M. chaiyaphumensis. A
female of this species was earlier reported from Ratan-
akiri Province, Cambodia, by Kosterin (2014).
Macromia ?cincta (Rambur, 1842)
A female was photographed on 1 July 2013. Its external
characters correspond to M. cincta, a male of which was
collected by the rst author on 23 May 2013, also in Koh
Kong Province. The la er record of this species, which
was rather unexpected in Cambodia, was published
by Kosterin (2014) without supporting arguments.
These will be specially presented elsewhere in a paper
devoted to Cambodian Macromia.
The following three species, already known from
Cambodia, were added to the already quite long list
of 72 species recorded in the study area by Kosterin
et al. (2012a):
Camacinia gigantea (Brauer, 1867). A female photo-
graphed on 29 June 2013 at Rainbow Lodge.
© Centre for Biodiversity Conservation, Phnom Penh
Cambodian Journal of Natural History 2014 (1) 8–11
New records of dragon ies
Brachydiplax farinosa Krüger, 1902. A male photo-
graphed on 12 June 2013 at a pond on the western
bank of the Stoeng Kep River (11.582˚N, 103.130˚E).
Brachythemis contaminata (Fabricius, 1793). A male
photographed at the same place and date as above.
The following recent ndings of rare species in the
study area, although already recorded in Cambodia
by Kosterin et al. (2012a), are also worth mentioning:
Heliaeschna crassa Krüger, 1899. A male was photo-
graphed near Rainbow Lodge on 4 March 2013. This
species was previously reported from the same area
from a photograph of a female (Day, 2011).
Heliaeschna uninervulata Martin, 1909. A female was
found alive inside a house at Rainbow Lodge and
released on 2 May 2013.
Burmagomphus asahinai Kosterin, Makbun &
Dawwrueng, 2012. A teneral female of this recently
described species was found and photographed on
15 July 2013 at Tatai Waterfall (11.586˚N, 103.097˚E).
Together with the data from Kosterin et al. (2012b),
these records suggest a prolonged emergence period
for imagines of this species from at least late March to
Amphithemis curvistyla Selys, 1891. An immature
male was photographed at Rainbow Lodge on 1
October 2013. Earlier, a female of this very rare species
was reported from the same site (Kosterin et al., 2012a).
The new records of two species for Cambodia
published herein and the recent paper by Kosterin
(2014), added to the checklist published by Kosterin et
al. (2012b), has raised the number of named Odonata
species known to occur in Cambodia to 154.
The authors are grateful to Martin Schorr for valuable
taxonomical information and great help with litera-
ture. Two anonymous referees greatly improved the
language and style of the paper.
Asahina, S. (1987) A list of the Odonata from Thailand. Part
XVIII. Corduliidae 2. Kontû, 55, 699–720.
Day, L. (2011) Odonata seen in Tatai, Koh Kong Province,
Cambodia. International Dragon y Fund Report, 42, 1–6.
Fraser, F.C. (1936) The Fauna of British India, including Ceylon
and Burma: Odonata, Volume III. Taylor and Francis,
Hämäläinen, M. (1985) Macromia chaiyaphumensis sp. n.
(Odonata, Corduliidae) from Northeast Thailand. Annales
Entomologici Fennici, 51, 105–107.
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Philippine Odonata, with lists of species recorded from
forty islands. Odonatologica, 26, 249–315.
Hämäläinen, M. & Pinratana, A. (1999) Atlas of the Dragon ies
of Thailand: Distribution Maps by Provinces. Brothers of St.
Gabriel in Thailand, Bangkok, Thailand.
Kosterin, O.E. (2014) Odonata of the south-west and north-
east of Cambodia as studied in early rainy season of 2013.
International Dragon y Fund Report, 67, 1–94.
Kosterin, O.E., Chartier, G., Holden, J. & Mey F.S. (2012a)
New records of Odonata from Cambodia, based mostly on
photographs. Cambodian Journal of Natural History, 2012,
Kosterin, O.E., Makbun, N. & Dawwrueng, P. (2012b)
Burmagomphus asahinai sp. nov., a new species from
Cambodia and Thailand, with a description of the male of
B. gratiosus Chao, 1954. International Journal of Odonatology,
15, 275–292.
Lieftinck, M.A. (1940) Descriptions and records of South-
East Asiatic Odonata (II). Treubia, 17, 337–390.
Lieftinck, M.A. (1948) Entomological results from the
Swedish Expedition 1934 to Burma and British India.
Arkiv för zoologi, Uppsala, 41A, 19–21.
Lieftinck, M.A. (1954) Handlist of Malaysian Odonata:
a catalogue of the dragon ies of the Malay Peninsula,
Sumatra, Java and Borneo, including the adjacent small
islands. Treubia (Supplement), 22, 1–201.
Martin, M. (1909) Collections Zoologiques du Baron Edm.
de Selys Longchamps. Catalogue Systématic et Descriptif.
Aeschnines. XX. Hayez, Brussels, Belgium.
Orr, A. (2003) A Guide to the Dragon ies of Borneo, their Identi-
cation and Biology. Natural History Publications (Borneo),
Kota Kinabalu, Sabah, East Malaysia.
Orr, A. (2008) Eight days in the Peninsular Malaysia—and
the bene ts of hindsights. Agrion, 31, 24–27.
Ris, F. (1927) Odonaten von Sumatra, gesammelt von
Edward Jacobson. Zoologische Mededeelingen, 10, 1–49.
Tsuda S. (2000) A Distributional List of World Odonata. Osaka,
© Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2014 (1) 12–13
T. Hartmann et al.
Short Communication
First record of the rare parachute gecko
Ptychozoon trinotaterra
Brown, 1999 from Cambodia
Timo HARTMANN1,*, Anna Bella BETTS2, Stéphane DE GREEF2 and Flora IHLOW2
1 Zoological Research Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany.
2 Wat Polanka, Slorkram Village, Siem Reap, Cambodia.
3 Center for Khmer Studies, PO Box 9380, Wat Damnak, Siem Reap, Cambodia.
*Corresponding author. Email
Paper submitted 5 December 2013, revised manuscript accepted 8 June 2014.
CITATION: Hartmann, T., Be s, A.B., De Greef, S. & Ihlow, F. (2014) First record of the rare parachute gecko Ptychozoon trino-
taterra Brown, 1999 from Cambodia. Cambodian Journal of Natural History, 2014, 12–13.
The rare parachute gecko Ptychozoon trinotaterra
Brown, 1999 was rst described based on a type series
consisting of a single female from Nakhon Ratch-
asima Province, Thailand, and three males from Gia
Lai and Dak Lak provinces, Vietnam (Brown, 1999).
In addition to the type series, populations of P. trino-
taterra have been reported from Ubon Ratchathani
Province in eastern Thailand (Kunya et al., 2011) and
Dong Nai Province in southern Vietnam (Nguyen
et al., 2009) (see Fig. 1). Stuart & Emme (2006) and
Nguyen et al. (2009) expected P. trinotaterra to occur
in northern Cambodia and southern Laos. Here, we
con rm their hypothesis and present the rst record
of P. trinotaterra from north-western Cambodia based
on photographs taken in Siem Reap Province.
In October 2013, a single unsexed specimen was
found resting on a tree trunk, approximately 1.5 m
above ground, during voluntary entomological eld-
work by Stéphane De Greef and Anna Bella Be s in
old secondary semi-evergreen forests south of the
Preah Khan Temple in the Angkor temples complex,
Nokor Thom Commune, Siem Reap Province,
Cambodia (13.458°N 103.869°E WGS84) (Fig. 2). The
specimen was found shortly after midday and a series
of detailed photographs was taken (Fig. 3).
Through the combination of the following charac-
ters the photo-vouchered individual can be assigned
to the species P. trinotaterra: three transverse M-shaped
dark bands in the axilla-groin region (vs four in
P. lionotum Annandale, 1905), M-shaped chevrons
without two posterior projections (vs distinct poste-
rior projections of chevrons in P. kaengkrachanense
Sumontha, Pauwels, Kunya, Limlikhitaksorn, Ruksue,
Fig. 1 Map showing the distribution of Ptychozoon trino-
taterra. THAILAND: (1) Nakhon Ratchasima Province
(Type Locality) (Brown, 1999); (2) Ubon Ratchathani
Province (Kunya et al., 2011); CAMBODIA: (3) Siem
Reap Province; VIETNAM: (4) Gia Lai Province (Brown,
1999); (5) Dak Lak Province (Brown, 1999); (6) Dong Nai
Province (Nguyen et al., 2009).
© Centre for Biodiversity Conservation, Phnom PenhCambodian Journal of Natural History 2014 (1) 12–13
First country record of parachute gecko
Fig. 3 Dorsal views of the live adult Ptychozoon trino-
taterra from Siem Reap Province, Cambodia (tail tip
regenerated) (© Anna Bella Be s).
Fig. 2 Habitat of Ptychozoon trinotaterra in Siem Reap
Province, Cambodia (© Stéphane De Greef).
Taokratok, Ansemet & Chanhome, 2012), and contin-
uation of caudal tubercles distally on to the dorsal
surface of tail terminus (vs no caudal tubercles on
terminal tail in P. lionotum) (Brown, 1999; Sumontha et
al., 2012) (see Fig 3.).
While Brown (1999) also stated the presence of a
mid-vertebral tubercle row as a diagnostic character
of P. trinotaterra, our specimen shows no such tuber-
cles. This supports the views of Kunya et al. (2011),
who postulate this character to be variable and hence
not diagnostic.
Besides records of P. lionotum from Phnom Sruoch,
Kirirom National Park, Kampong Speu Province,
south-western Cambodia by Stuart & Emme (2006)
our record of P. trinotaterra from north-western
Cambodia represents the second documented species
of the parachute gecko genus Ptychozoon from this
Brown, R.M. (1999) New species of parachute gecko
(Squamata: Gekkonidae: genus Ptychozoon) from north-
eastern Thailand and central Vietnam. Copeia, 1999,
Kunya, K., Pauwels, O.S.G., Sairum, P. & Taokratok, A. (2011)
Squamata, Gekkonidae, Ptychozoon trinotaterra Brown,
1999: rediscovery in Thailand. Check List, 7, 820–822.
Nguyen V.S., Ho T.C. & Nguyen Q.T. (2009) Herpetofauna of
Vietnam. Edition Chimaira, Frankfurt am Main, Germany.
Stuart, B.L. & Emme , D.A. (2006) A collection of amphib-
ians and reptiles from the Cardamom Mountains,
southwestern Cambodia. Fieldiana (Zoology), 109, 1–27.
Sumontha, M., Pauwels, O.S.G., Kunya, K., Limlikhitaksorn,
C., Ruksue, S., Taokratok, A., Ansermet, M. & Chanhome,
L. (2012) A new species of parachute gecko (Squamata:
Gekkonidae: genus Ptychozoon) from Kaeng Krachan
National Park, western Thailand. Zootaxa, 3513, 68–78.
© Centre for Biodiversity Conservation, Phnom Penh
S. Ravon et al.
Cambodian Journal of Natural History 2014 (1) 1418
Short Communication
A rapid assessment of ying fox (
spp.) colonies in
Sébastien RAVON1, Neil M. FUREY2,*, HUL Vibol3 and Julien CAPPELLE1,4
1 Institut Pasteur du Cambodge, Epidemiology and Public Health Unit, BP983, Phnom Penh, Cambodia.
2 Fauna & Flora International (Cambodia Programme), PO Box 1380, No. 19, Street 360, Boeng Keng Kong 1, Phnom
Penh, 12000, Cambodia.
3 Institut Pasteur du Cambodge, Virology Unit, BP983, Phnom Penh, Cambodia.
4 Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD), UR Animal et
Gestion Intégrée des Risques (AGIRs), F-34398, Montpellier, France.
*Corresponding author. Email neil.furey@fauna-
Paper submitted 11 May 2014, revised manuscript accepted 10 July 2014.
CITATION: Ravon, S., Furey, N.M., Hul V. & Cappelle, J. (2014) A rapid assessment of ying fox (Pteropus spp.) colonies in
Cambodia. Cambodian Journal of Natural History, 2014, 14–18.
Flying foxes (Chiroptera: Pteropodidae: Pteropus) are
among the few wide-ranging frugivores still found
in many parts of Southeast Asia and play important
seed-dispersion and pollination roles in their ecosys-
tems (Cox et al., 1991; Fujita & Tu le, 1991; Struebig et
al., 2007). Three ying fox species were depicted for
Cambodia in the range maps of Francis (2008): large
ying fox Pteropus vampyrus, Lyle’s ying fox P. lylei
and island ying fox P. hypomelanus. These species are
currently considered to be globally Near-threatened,
Vulnerable and Least Concern by the IUCN (Bates et
al., 2008; Bumrungsri et al., 2008; Francis et al., 2008)
respectively, and as nationally common (P. vampyrus
and P. lylei) or nationally rare (P. hypomelanus) in
Cambodian legislation (MAFF, 2007). All three species
are included in Appendix II of CITES, but almost
nothing is known about their conservation status in
Cambodia. Although likely present, the occurrence
of P. vampyrus remains uncon rmed, having yet to
be validated by the unequivocal documentation of a
live animal or museum specimen to our knowledge.
As colony surveys are central to determining conser-
vation priorities for ying foxes (Mickleburgh et al.,
1992), we provide here the ndings of a rapid assess-
ment of pteropodid colonies in Cambodia.
Using an unpublished list of roost sites provided
by the Wildlife Conservation Society, supplemented
by additional sites reported by local eld workers,
we conducted eld surveys between June 2013 and
February 2014 to assess all of the known or suspected
Pteropus colonies in Cambodia. At every site, the
location and basic se ing of the roost environment
was recorded and standardized estimates of roost
populations made using direct census methods, and,
where possible, nightly dispersal counts (Kunz et al.,
1996). E orts were also made to identify the species
present at each site, but because these necessarily
relied upon impressions of relative size (Francis,
2008) using binoculars, species identi cations were
uncertain and so no a empts were made to estimate
total population sizes for each species. As P. hypome-
lanus primarily occurs in coastal areas and on marine
islands, however (Francis, 2008), we assumed that the
species present at all inland sites surveyed were P.
lylei and/or P. vampyrus (Fig. 1).
Direct censuses were undertaken by visually
counting the bats during the day at each site with the
aid of 8 x 42 binoculars and manual hand-counters,
whereas nightly dispersal counts began at dusk when
the bats emerged to forage until all had left the roost
site (typically from 1830 h to 1910 h). The la er was
© Centre for Biodiversity Conservation, Phnom PenhCambodian Journal of Natural History 2014 (1) 1418
Flying fox colonies
site-based conservationists around the country, it
appears likely these comprise most of the ying fox
colonies in Cambodia. (The possibility that other
colonies might be discovered in the future cannot be
excluded, however, particularly in poorly surveyed
coastal regions and areas surrounding the Tonle Sap
Lake). At least one interview was completed at each
site. According to local informants, all of the colonies
assessed were present year-round. O spring were
largely reported by respondents as appearing in April
each year, and this is supported by monthly observa-
tions at the Wat Pi Chey Sa Kor (Kandal Province) and
Wat Bay Dam Ram (Ba ambang Province) colonies
where mating takes place in November and parturi-
tion primarily occurs in April (Hul, 2013; J. Cappelle,
unpublished data).
Most of the roost sites were situated inside the
grounds or within the vicinity of a religious or govern-
con rmed by checking roost trees with a spotlight
after the dispersal count at each site. Due to the
density of bats and brevity of the evening dispersal,
in some instances bats were counted in groups of 10
as they dispersed. The higher count from the two
methods was rounded down to the nearest hundred
and accepted as the estimated population size for a
given site. Interviews were also undertaken by the
rst author with local authorities and residents at
roost sites to determine: (i) the status of the colony
(permanent or seasonal); (ii) annual breeding periods
(de ned as birth periods); (iii) whether the colony
receives any protection; (iv) conservation threats at
each site; and (v) local perceptions concerning the
ying fox colony.
Over the course of the rapid eld survey, 12
roost sites were located and assessed (Table 1, Fig.
2). Based on our experience and discussions with
Fig. 1 Flying foxes on Koh Trong Island on the Mekong River (© Gordon Congdon, left) and Koh Bong Island o the
coast of Cambodia (© Jeremy Holden, right). The species on the left is thought to be P. lylei and the species on the right
P. hypomelanus.
© Centre for Biodiversity Conservation, Phnom Penh
S. Ravon et al.
Cambodian Journal of Natural History 2014 (1) 1418
Fig. 2 Locations and
relative sizes of 12 ying
fox colonies in Cambodia.
Table 1 Summary characteristics of 12 ying fox colonies in Cambodia. Key: * Site where hunting of bats was reported
or observed; 1 DC = Direct Census, ND = Night Dispersal, n/a = not applicable; 2 Value given is the highest gure from
the count methods, rounded down to the nearest hundred.
# Site Name (Province) Latitude,
Longitude Census Date Roost Environment Census
1 Ang Trapeang Thmor
(Banteay Meanchey)
13.804 N,
103.261 E.
14 Aug 2013 One roost tree on small
island in reservoir.
DC 200
2 Wat Bay Dam Ram
(Ba ambang)
12.993 N,
103.161 E.
23 Jun 2013 Pagoda. Three roost
trees in site vicinity.
DC / ND 1,400
3 Royal Gardens *
(Siem Reap)
13.363 N,
103.859 E.
10 Aug 2013 O cial site. 14 roost
trees in urban park.
DC / ND 5,000
4 Kampong Thom *
(Kampong Thom)
12.714 N,
104.883 E.
8 Aug 2013 O cial site. Three roost
trees along roadside.
DC / ND 6,000
5 Koh Trong Island
12.507 N,
105.993 E.
26 Aug 2013 Pagoda. Two roost trees
on site perimeter.
DC 200
6 Koh Chreng Island *
12.361 N,
106.044 E.
27 Aug 2013 Pagoda. 17 roost trees
on site perimeter.
DC 900
7 Wat Srey Santaor *
(Kampong Cham)
11.915 N,
105.183 E.
8 Aug 2013 Pagoda. Small forest on
site perimeter.
n/a Extirpated
8 Council for Development
of Cambodia
(Phnom Penh)
11.577 N,
104.924 E.
18 Oct 2013 O cial site. One roost
tree on site perimeter.
DC / ND 1,800
9 Wat Prek Chey Lech *
11.465 N,
105.235 E.
1 Aug 2013 Pagoda. Five roost trees
on site perimeter.
DC / ND 500
10 Wat Pi Chey Sa Kor *
11.200 N,
105.058 E.
15 Jan 2014 Pagoda. 21 roost trees
on site perimeter.
DC / ND 4,000
11 Wat Veal Lbang
(Prey Veng)
11.173 N,
105.310 E.
17 Jan 2014 Pagoda. 12 roost trees in
site vicinity.
DC 700
12 Koh Bong Island
10.759 N,
103.265 E.
1 Feb 2014 Three roost trees in
forest on small private
DC 200
© Centre for Biodiversity Conservation, Phnom PenhCambodian Journal of Natural History 2014 (1) 1418
Flying fox colonies
colony areas to eliminate misconceptions regarding
their medicinal values and to generate local support
for their protection. To this end, a website (www. has been developed to
gain public information about ying fox colonies
in Cambodia, which we hope will help to stimulate
greater conservation interest in these charismatic and
inherently vulnerable animals.
The authors are grateful to Mr Prak Bali for his assis-
tance in nding ying fox colonies, Benjamin Hayes
for his advice and to the SE Asian Bat Conservation
and Research Unit for their support. We also thank the
two anonymous reviewers who kindly commented on
the text. The study was supported by the SouthEast
Asia Encephalitis project which is funded by Aviesan
Sud and Fondation Total.
Bates, P., Francis, C., Gumal, M., Bumrungsri, S., Walston,
J., Heaney, L. & Mildenstein, T. (2008) Pteropus vampyrus.
In IUCN Red List of Threatened Species. H p://www.> [accessed 2 July 2014].
Bumrungsri, S., Suyanto, A. & Francis, C. (2008) Pteropus
lylei. In IUCN Red List of Threatened Species. H p://www.> [accessed 2 July 2014].
Cox, P.A., Elmqvist, T., Pierson, E.D. & Rainey, W.A. (1991)
Flying foxes as strong interactors in South Paci c island
ecosystems: a conservation hypothesis. Conservation
Biology, 5, 448–454.
Francis, C.M. (2008) A Guide to the Mammals of Southeast Asia.
Princeton University Press, Princeton, New Jersey, USA.
Francis, C., Rosell-Ambal, G., Bonaccorso, F.A., Heaney,
L., Molur, S. & Srinivasulu, C. (2008) Pteropus hypome-
lanus. In IUCN Red List of Threatened Species. H p://www.> [accessed 2 July 2014].
Fujita, M.S. & Tu le, M.D. (1991) Flying foxes (Chiroptera:
Pteropodidae): threatened animals of key ecological and
economic importance. Conservation Biology, 5, 455–463.
Hul V. (2013) Ecology of ying fox (Pteropus species) and assess-
ment of the risk of emergence of Nipah virus in Ba ambang and
Kandal Provinces, Cambodia. MSc thesis, Royal University
of Phnom Penh, Cambodia.
Kunz, T.H., Thomas, D.W., Richards, G.C., Tidemann,
C.R., Pierson, E.D. & Racey, P.A. (1996) Observational
techniques for bats. In Measuring and Monitoring Biological
Diversity: Standard Methods for Mammals (eds D.E. Wilson,
F.R. Cole, J.D. Nichols, R. Rudran & M.S. Foster), pp.
105–114, Smithsonian Institution Press, Washington D.C.,
ment building, which apparently a orded them some
protection from hunting (seven roosts were near a
pagoda and three were near a government building).
Only two roosts were protected by a natural barrier:
the very small uninhabited islands of Ang Trapaeng
Thmor and Koh Bong in the country’s Northwest and
Southwest respectively (Table 1, Fig. 2). This situa-
tion is similar to Thailand—another predominantly
Buddhist country—where many ying fox colonies
are near pagodas (Wacharapluesadee et al., 2010),
but di ers from the Philippines—a largely Christian
and Muslim country—where most colonies are in
forest areas (Mildenstein, 2012). Religion may there-
fore play an important role in ying fox conservation
in Cambodia, and a be er understanding of local
perceptions of bats could aid the design of more e ec-
tive conservation initiatives.
Despite the potential in uence of religious views
however, hunting of bats for bushmeat, trade and/or
medicinal use was reported at half of the 12 sites in
Table 1. In fact, one colony in Srey Santaor (Kampong
Cham Province) was recently extirpated by large-scale
hunting, despite protests from local monks (Prak Bali,
pers. comm.), and colonies at other sites could well
be declining. Even though most of the remnant ying
fox colonies in Cambodia now appear to be con ned
to sites which a ord some measure of protection,
the bats are still actively hunted while foraging or
occur in non-protected areas (Timmins, 2008; present
study). Thus, while there is a clear need for further
surveys—preferably entailing synchronized counts
employing standard methods at all known colonies to
establish seasonal variation and population trends—
our data nonetheless suggest that ying fox colonies
in mainland Cambodia are heavily threatened and by
no means nationally common.
This poses an obvious concern, not least because
ying foxes can cease to be e ective seed dispersers
long before they become rare (McConkey & Drake,
2006). Besides their ecological services to humans,
ying foxes may also play a role as reservoir of
pathogens of public health importance in Cambodia.
Evidence of Nipah virus circulation was reported
in national ying fox populations some years ago
(Reynes et al., 2005), but very li le information is
available on the risk of transmission to domestic
animals and humans in the country. Further research
on the status and ecology of Cambodian Pteropus is
therefore central to overcoming current challenges to
reliable eld identi cation and designing conserva-
tion plans and public health risk mitigation strategies.
Campaigns to raise awareness are also required in
© Centre for Biodiversity Conservation, Phnom Penh
S. Ravon et al.
Cambodian Journal of Natural History 2014 (1) 1418
MAFF—Ministry of Agriculture, Forestry & Fisheries (2007)
Prakas on Classi cation and List of Wildlife Species. Ministry
of Agriculture, Forestry and Fisheries, No. 020 PR. Ministry
of Agriculture, Forestry & Fisheries, Phnom Penh,
McConkey, K.R. & Drake, D.R. (2006) Flying foxes cease to
function as seed dispersers long before they become rare.
Ecology, 87, 271–276.
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World Fruit Bats: an Action Plan for their Conservation.
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Mildenstein, T.L. (2012) Conservation of endangered ying foxes
in the Philippines: e ects of anthropogenic disturbance and
methods for monitoring. PhD thesis, University of Montana,
Reynes, J.M., Counor, D., Ong, S. & Faure, C. (2005) Nipah
virus in Lyle’s ying foxes, Cambodia. Emerging Infectious
Diseases, 11, 1042–1047.
Struebig, M.J., Harrison, M.E., Cheyne, S.M., Limin, S.H.
(2007) Intensive hunting of large ying foxes Pteropus
vampyrus natunae in Central Kalimantan, Indonesian
Borneo. Oryx, 41, 390–393.
Timmins, R. (2008) Large mammals. In Biological Surveys of
the Mekong River between Kratie and Stung Treng Towns,
Northeast Cambodia, 2006–2007 (eds M.R. Bezuijen, R.
Timmins & Seng T.), pp. 82–89. WWF Cambodia Country
Programme, Cambodian Fisheries Administration &
Cambodian Forestry Administration, Phnom Penh,
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asetyuth, N., Supavonwong, P., Saengsen, D., Gongal,
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Nipah virus in Pteropus lylei bats in Thailand: evidence for
seasonal preference in disease transmission. Vector-borne
and Zoonotic Diseases, 10, 183–190.
© Centre for Biodiversity Conservation, Phnom Penh
Small mammal trapping in Bokor
Cambodian Journal of Natural History 2014 (1) 1923
Short Communication
A small terrestrial mammal survey and analysis of bait
consumption at Bokor National Park, Cambodia
Marissa PARROTT1,*, CHHIN Sophea2, HUN Seiha2, CHHEANG Sarak2 and Neil FUREY2,3
1 Wildlife Conservation & Science, Zoos Victoria, Victoria 3052, Australia.
2 Centre for Biodiversity Conservation, Room 415, Department of Biology, Faculty of Science, Royal University of
Phnom Penh, Confederation of Russia Boulevard, Phnom Penh, Cambodia.
3 Fauna & Flora International (Cambodia), #19, Street 360, Boeung Keng Kong 1, Phnom Penh, Cambodia.
*Corresponding author. Email
Paper submitted 30 April 2014, revised manuscript accepted 15 July 2014.
CITATION: Parro , M., Chhin S., Hun S., Chheang S. & Furey, N. (2014) A small terrestrial mammal survey and analysis of bait
consumption at Bokor National Park, Cambodia. Cambodian Journal of Natural History, 2014, 19–23.
Cambodia boasts a rich array of species and the
Greater Cardamom Mountains in the country’s South-
west are recognised as part of a global biodiversity
hotspot (Myers et al., 2000) and one of the planet’s 200
Global Ecoregions (Olson et al., 2000). Survey e ort in
this region has recently intensi ed and uncovered new
species of plants (Mey, 2009), amphibians and reptiles
(e.g. Daltry & Wuster, 2002; Grismer et al., 2008; Neang
et al., 2012). Medium-sized to large mammals have
also been a research focus in Cambodia, with several
papers describing their distribution, populations and
threats (e.g. Emme & Olssen, 2005; Royan, 2010;
Edwards et al., 2012; Gray et al., 2012). In contrast,
li le work has been conducted on the small terrestrial
mammals of Cambodia. Bats (order Chiroptera) have
been the most intensively researched small mammals
in Cambodia, with 70 species presently known
(Chheang et al., 2013), including taxa new to science
(Csorba, 2011; Csorba et al., 2011). However, a recent
study identi ed a new species of shrew (Jenkins et al.,
Within the Cardamom Mountains, only two
systematic surveys of small terrestrial mammals have
been reported. The rst used aluminium box traps
and pitfall traps and caught mostly rodents (Muridae)
except for a single capture of the insectivore Hylomys
suillus (Swan & Kry, 2000). The second study employed
box, pitfall and mesh traps and encountered two
species of shrew (Crocidura spp.), two species of tree
shrews and eight rats (Emme & Olssen, 2005).
Capture rates for small terrestrial mammals in
forested ecosystems in Cambodia tend to be low. For
instance, Swan & Kry (2000) recorded a mean capture
rate of 1.3% and 0.9% for small terrestrial mammals in
the Phnom Samkos Wildlife Sanctuary, while Emme
& Olssen (2005) achieved a capture rate of 2.6% for
small and medium-sized mammals using traps in
the Central Cardamoms Protected Forest. These are
similar to the capture rates of 0.04–2.6% in northern
Cambodia (Edwards et al., 2012), but lower than a
study of rodents across four habitat types in eastern
and southern Cambodia, including se lements and
agricultural areas (mean 13.5%: Ivanova et al., 2012).
The type of bait used to a ract mammals can
signi cantly a ect the species and number of individ-
uals trapped (Patric, 1970; Kok et al., 2013). Most
studies in Cambodia have used vegetable ma er,
including sticky rice, peanuts, bananas and/or cassava
as bait (Kemper & Bell, 1985; Swan & Kry, 2000;
Emme & Olssen, 2005; Blasdell et al., 2011; Edwards
et al., 2012; Starr et al., 2012), but two studies used
dried sh or shrimp (Swan & Kry, 2000; Conservation
International, 2007), one used dried beef (Conserva-
tion International, 2007) and one used chicken or
egg (Starr et al., 2012). Shrews may be particularly
a racted to meat lures, but li le is known of these
© Centre for Biodiversity Conservation, Phnom Penh
M. Parrott et al.
Cambodian Journal of Natural History 2014 (1) 1923
species in Cambodia (Emme & Olssen, 2005; Jenkins
et al., 2010). As species composition is likely to vary
markedly across Cambodia and di erent bait types
may a ect the numbers of species a racted to traps,
this paper reports the results of a pilot investigation of
small mammal assemblages and their bait consump-
tion in two sites in Bokor National Park.
Bokor National Park (Fig. 1) covers an area of
>1,400 km2 and is in the Elephant Mountains of
Cambodia (Seng et al., 2003), a southern o shoot of
the Cardamom Mountains. The small mammal study
was undertaken at two sites in the southern portion of
the park from 11–24 February 2013 as part of a broader
biodiversity study investigating amphibians, reptiles,
birds, bats, large mammals and gibbons from October
2012 to May 2013.
In Site 1 (Fig. 1), Ellio small live mammal traps
(9 cm × 10 cm × 33 cm) were set along a transect at
approximately 15–20 m intervals for a total of 60
trap-nights. In Site 2 (Fig. 1), Ellio traps were set
for a total of 50 trap-nights along four transects with
approximately 15–20 m intervals between traps. The
four transects were along a dry creek bed (15 trap-
nights), o a steep path (21 trap-nights), on a sloped
path (six trap-nights) and along a stream (eight trap-
nights). Where necessary, traps were moved within
their chosen locations to protect trapped animals from
ants and to reduce the amount of bait consumed by
non-target species.
Traps were placed in areas likely to be used by
mammals including alongside fallen logs, under
boulders, and near runways, holes and observed
scats. All traps were checked at dawn (0630 h),
midday (1300 h) and dusk (1730 h). Every trap was
covered with thick plastic to protect animals from
wind and rain and was lled with co on wadding and
tissue paper to provide nesting material. As animal
preferences for bait were unknown, every trap was
baited simultaneously with 5 g of locally produced
Fig. 1 Location of study sites in Bokor National Park, Cambodia. Site 1 was at 10°43.826’N, 103°55.601’E, at an eleva-
tion of 408 metres above sea level (asl), and consisted of wet dipterocarp forest with bananas and sugar palms. Canopy
cover was estimated at >80% and there was extensive ground cover with many fallen logs. Site 2 was at 10°45.542’N,
103°55.154’E, at an elevation of 416 m asl, and consisted of drier dipterocarp forest with none of large-leafed plants that
occurred in Site 1. Canopy cover was estimated at >60% and there was less ground cover than in Site 1.
© Centre for Biodiversity Conservation, Phnom Penh
Small mammal trapping in Bokor
Cambodian Journal of Natural History 2014 (1) 1923
lap cheong (fa y pork salami), uncooked rice, dried
papaya, dried cranberries, liver dog treats and a mixed
ball of peanut bu er, rolled oats and honey. Following
every capture, the remaining bait was collected and
the amount consumed was assigned to the following
classes: 100%, 75–99%, 50–74%, 25–49% and <25%.
Upon capture, every animal was weighed using a
Pesola™ 60 g or 600 g spring balance and measured
using vernier callipers. Reproductive status and age
class were also recorded (Aplin et al., 2003). On their
rst capture, animals were individually marked with
a unique set of ear clips and the removed tissue was
stored in 90% ethanol for genetic analyses. Animals
were generally handled in cloth bags, but the Maxomys
sp. was measured brie y in a large clear plastic bag
because the animals appeared calmer and handling
of their delicate tail skin was avoided (Francis, 2008).
Genetic samples were deposited and stored at -20°C
at the Royal University of Phnom Penh Centre for
Biodiversity Conservation (they will be analysed at
the Museum Victoria, Australia). Following measure-
ment, all animals were released at their point of
All captures occurred overnight with no animals
trapped during daylight hours. There were 29
successful captures from 110 trap-nights giving an
overall capture rate of 26%.
From the total of 60 trap-nights in Site 1, four speci-
mens were trapped, giving a capture rate of 7%. They
included a 160 g female rat with silver upperparts,
white underparts and feet, and a mo led grey tail
(Fig. 2), which was identi ed as Berylmys berdmorei,
albeit smaller than published measurements for this
species (Aplin et al., 2003). Two female shrews were
trapped, which were dark grey with a tail shorter than
their head-body length and had sparse hairs near the
base of the tail (Fig. 3). They weighed 7.3 and 7.5 g
(mean 7.4 g) and had a distinctive musky odour. These
animals were likely in the genus Crocidura, but their
measurements did not match any species presently
known in Cambodia or Southeast Asia. A palm civet
Paradoxurus hermaphrodites was also trapped, but after
consuming the bait it broke out of the trap and was
subsequently identi ed from its hair and scats.
A total of 50 trap-nights were conducted in Site 2
with 15 rats (seven females, eight males) trapped on 25
occasions (including 10 recaptures), giving a capture
rate of 50%. The rats’ upperparts were a distinctive
reddish-orange colour with a darker midline and
numerous sti dark spines throughout the fur (Fig.
4). Underparts were white with small sti spines and
Fig. 2 Rat resembling Berylmys berdmorei trapped in Site
Fig. 3 Crocidura sp. trapped in Site 1.
Fig. 4 Maxomys sp. trapped in Site 2 (being brie y held
in plastic bag).
© Centre for Biodiversity Conservation, Phnom Penh
M. Parrott et al.
Cambodian Journal of Natural History 2014 (1) 1923
were sharply demarcated from upperparts. The tail
was bicoloured, being dark grey above and white
underneath, with a white tip comprising one-third of
the tail length. Feet were white and ears were dark
grey. Mean mass was 170.27 ± 22.38 g (males 173.6 ±
21.6 g; females 166.4 ± 24.3 g) and ranged from 130–209
g. This species appeared most similar to Maxomys
surifer (Aplin et al., 2003), but had a number of di er-
ences, including consistently shorter ears. The rats
were all adults in breeding condition with enlarged
testes in the males and signs of early-mid pregnancy
in three of the seven females.
Rates of bait consumption by the rodents were:
lap cheong 100%, dog treats 50–74%, bait ball 50–74%,
dried papaya 25–49%, dried cranberries 25–49% and
rice <25%. The shrews consumed lap cheong only. The
Maxomys rats trapped on consecutive nights gained
between 2–13 g (mean 5.14 ± 1.64 g), the equivalent
of up to 8% of their body mass, indicating that large
amounts of bait had been consumed.
This is the rst study of small terrestrial mammals
at Bokor National Park, and insectivores, carnivores
and rodents were trapped. To the best of our knowl-
edge, the trap success rate achieved at Site 2 (50%) is
the highest recorded for small terrestrial mammals
in Cambodia to date. This may be a ributed to the
rats occurring at high population densities, becoming
“trap happy” (whereby individuals that are trapped
have a positive experience and are more inclined to
enter traps again), and/or the use of more a ractive
The pork-based lap cheong was entirely consumed
by all species caught, and shrews consumed this bait
alone. This is likely due to their small size (7.4 g) and
inability to eat any further items during their brief
period in the traps. Rodents also consumed greater
amounts of liver dog treats and bait ball mixture
than dried fruits. Li le to no rice was consumed. This
mirrors studies in South Africa where peanut bu er
and oat mixes were the most successful bait, achieving
more captures than fruit or meat lures (Kok et al., 2013).
Similarly, peanut bu er mixes are the most e ective
lure in the USA, although meat lures are e ective at
a racting carnivores (Patric, 1970). However, a study
of arboreal species in Cambodia found banana to
be a more e ective lure than chicken for small and
medium-sized mammals, including carnivores (Starr
et al., 2012). Small mammal surveys in Cambodia have
commonly used fruit or sticky rice as bait (Kemper
& Bell, 1985; Swan & Kry, 2000; Blasdell et al., 2011),
although two studies also used dried sh, shrimp and
beef (Swan & Kry, 2000; Conservation International,
2007). We recommend the use of multiple food types,
including meat lures and peanut bu er mixes, to
increase capture success in Cambodia.
This short study may have uncovered new species
of rat and shrew, but further genetic and morpholog-
ical work is required and will be conducted shortly.
Further studies on shrews are likely to uncover new
records for Cambodia and potentially species new to
science. In 2004, Emme & Olssen (2005) collected two
unidenti ed species of shrew. Two further unidenti-
ed species were found in Virachey National Park in
2007 (Conservation International, 2007), and a new
species of Crocidura was recently described from
Northeast Cambodia (Jenkins et al., 2010). Among the
rodents, the genus Maxomys in particular has highly
divergent populations within and between island
regions (Achmadi et al., 2013) and new Maxomys
species continue to be described (e.g. Achmadi et
al., 2012). Emme & Olssen (2005) also discovered
Maxomys and Niviventer species in the Cardamom
Mountains which appear to di er externally from
previously recorded species. With so li le research on
small terrestrial mammals in Cambodia, new records
and possibly new species likely await discovery.
Further research and allied conservation initiatives
are vital to protect biodiversity in Bokor National
Park. Several areas within the park are likely to
support high numbers of small mammal species and
need protection from inappropriate land-use. More
intensive small mammal surveys, using a variety of
traps and baits over a longer time period in di erent
regions of Bokor National Park, and Cambodia as a
whole, are required to elucidate species diversity and
conservation requirements for future management.
This research was conducted under a Conservation
Leadership Programme grant to Chhin Sophea and a
Zoos Victoria Fellowship to Marissa Parro . We thank
Nick Souter, Ouk Dane, Chris Banks and Michael
Magrath for their assistance with the organisation of
this project.
Achmadi, A.S., Maryanto, I. & Maharadatunkamsi (2012)
Systematic and descriptions of new species within genus
Maxomys from East Kalimantan, Borneo Island. Treubia,
39, 1–28.
© Centre for Biodiversity Conservation, Phnom Penh
Small mammal trapping in Bokor
Cambodian Journal of Natural History 2014 (1) 1923
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Abdullah, M.T. (2013) Phylogeny, diversity, and biogeog-
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G.R. (2003) Field Methods for Rodent Studies in Asia and
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Blasdell, K., Cosson, J.K., Chaval, Y., Herbreteau, V., Douang-
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T. Naruse et al.
Cambodian Journal of Natural History 2014 (1) 2436
Notes on a collection of stomatopod and decapod crustaceans
from Cambodia
Tohru NARUSE1,*, Darren C.J. YEO2 and Masayuki OSAWA3
1 Tropical Biosphere Research Center, Iriomote Station, 870 Uehara, Taketomi, Okinawa 907–1541, Japan.
2 Department of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore 117543,
Republic of Singapore.
3 Research Center for Coastal Lagoon Environments, Shimane University, 1060 Nishikawatsu-cho, Matsue, Shimane
690–8504, Japan.
*Corresponding author. Email
Paper submitted 23 June 2014, revised manuscript accepted 17 July 2014.
CITATION: Naruse, T., Yeo, D.C.J. & Osawa, M. (2014) Notes on a collection of stomatopod and decapod crustaceans from
Cambodia. Cambodian Journal of Natural History, 2014, 24–36.
The crustacean fauna of Cambodia is poorly known. The present study records 21 species of Malacostraca (three
stomatopods, one scyllarid lobster and 17 brachyuran crabs) from the by-catch of shermen in Kampot City,
Kampot Province, Cambodia. Twelve species are new records for this country. Photographs and illustrations as
well as taxonomic and ecological notes are provided for every species.
Marine biodiversity, crustaceans, faunal survey, by-catch, Gulf of Thailand.
Cambodia has a coastline of 435 km in length,
covering an area estimated at between 17,791 km² and
18,477 km², including mangroves (728 km²), seagrass
beds (301–325 km²), and coral reefs (approximately
10 times smaller than area covered by seagrass)
(Spalding et al., 2010; Rizvi & Singer, 2011). So far,
only about 50 species of crustaceans have been listed
for Cambodia (UNEP, 2007). Considering that more
than 600 crustacean species have been recorded from
the Gulf of Thailand ports in neighbouring Thailand
(Naiyanetr, 2007), the marine and coastal biodiversity
of Cambodia is clearly understudied.
In December 2010, the rst and second authors
had an opportunity to make a brief visit to a local
shing port in Kampot City, Kampot Province, and
found many crustacean species among the by-catch of
© Centre for Biodiversity Conservation, Phnom Penh
Crustaceans found in by-catch
Cambodian Journal of Natural History 2014 (1) 2436
(CL) × carapace width (CW). All measurements are in
Species accounts
Order STOMATOPODA Latreille, 1817
Family SQUILLIDAE Latreille, 1802
Carinosquilla multicarinata (White, 1849) (Fig. 1a)
Material examined: RUMF-ZC-1539, two males, 17.0 /
73.6 mm, 18.2 / 78.3 mm, two females, 20.2 / 86.7 mm,
17.3 / 72.4 mm.
Distribution: Southern India to Indonesia, Thailand
(Andaman Sea and Gulf of Thailand), Cambodia,
Vietnam, the Philippines, China, Taiwan and Japan
(Naiyanetr, 2007; Ahyong et al., 2008; present study).
Remarks: The present specimens agree well with recent
descriptions of C. multicarinata (e.g. Ahyong & Moosa
2004; Ahyong et al., 2008). As discussed by Ahyong
(2001) and Ahyong et al. (2008), C. multicarinata closely
trash sh that were being sold to a shmeal factory.
Such by-catch can be a very useful source of material
for faunal studies, including rare or unusual species
(Jensen et al., 2011). The present paper reports some of
the crustaceans sampled (21 species in total). Photo-
graphs as well as taxonomic and ecological notes are
provided for the species.
The material examined was deposited in the Ryukyu
University Museum, Fujukan (RUMF), University of
the Ryukyus, Okinawa, Japan; the Zoological Refer-
ence Collection (ZRC) of the Lee Kong Chian Natural
History Museum (formerly known as Ra es Museum
for Biodiversity Research) of the National Univer-
sity of Singapore; and the National Museum of the
Philippines in Manila (NMCR). All specimens from
Cambodia were collected from the Kampong Kandal
Fishery Port (10°34’7.01”N, 104°14’33.21”E), Kampot
City, Kampot Province, on 27 December 2010 by T.
Naruse, D.C.J. Yeo, L. Sarorn and E.C.K. Khoo.
In the following accounts, the body sizes of
stomatopods and scyllarid lobsters are presented as
carapace length (CL) / total length (TL), while those
of the brachyurans are presented as carapace length
Fig. 1 Stomatopod and scyllarid species collected from Kampong Kandal Fishery Port: a, Carinosquilla multicarinata
(White, 1849) (RUMF-ZC-1539, male, 17.0 / 73.6 mm); b, Erugosquilla woodmasoni (Kemp, 1911) (RUMF-ZC-1541, male,
31.1 / 138.3 mm); c, d, Odontodactylus cultrifer (White, 1851) (RUMF-ZC-1540, male, 23.3 / 93.4 mm, in live colouration);
e, Biarctus sordidus (Stimpson, 1860) (RUMF-ZC-1328, male, 16.5 / 42.5 mm, in live colouration). a, b, c, e, habitus, dorsal
view; d, telson and uropods, lateral view.
© Centre for Biodiversity Conservation, Phnom Penh
T. Naruse et al.
Cambodian Journal of Natural History 2014 (1) 2436
resembles C. carita Ahyong, 2001, known only from
Australia, but di ers from the la er in the presence
of the mandibular palp and the prelateral lobe of the
telson with a spinular instead of blunt apex.
Erugosquilla woodmasoni (Kemp, 1911) (Fig. 1b)
Material examined: RUMF-ZC-1541, two males, 31.1 /
138.3 mm, 21.0 / 81.6 mm.
Distribution: Western Indian Ocean to Indonesia;
Thailand (Gulf of Thailand), Cambodia, Vietnam, the
Philippines, Taiwan, Japan and Australia (Naiyanetr,
2007; Ahyong et al., 2008; present study).
Remarks: The present material agrees well with recent
descriptions of E. woodmasoni (e.g. Ahyong, 2001;
Ahyong et al., 2008). This species is readily distin-
guished from its congeners by the merus of the
raptorial claw possessing an outer inferodistal spine
(versus no spine in the other species).
Family ODONTODACTYLIDAE Manning, 1980
Odontodactylus cultrifer (White, 1851) (Figs 1c, d)
Material examined: RUMF-ZC-1540, one male, 23.3 /
93.4 mm.
Distribution: Thailand (Andaman Sea and Gulf of
Thailand), Cambodia, South China Sea, Indonesia,
Taiwan, Australia and New Caledonia (Naiyanetr,
2007; Ahyong et al., 2008; present study).
Remarks: The present specimen agrees well with recent
descriptions of O. cultrifer (e.g. Ahyong, 2001; Ahyong
et al., 2008). This species is immediately distinguished
from its congeners by the high, thin median crest on
the telson (Fig. 1d) (versus no such crest in the others).
Order DECAPODA Latreille, 1802
Infraorder ACHELATA Scholtz & Richter, 1995
Family SCYLLARIDAE Latreille, 1825
Biarctus sordidus (Stimpson, 1860) (Fig. 1e)
Material examined: RUMF-ZC-01328, one male, 16.5 /
42.5 mm.
Distribution: Persian Gulf to Australia, the Gulf of
Thailand, Cambodia, southern China (Holthuis, 2002;
Naiyanetr, 2007; present study).
Remarks: The present specimen agrees well with the
redescription of Biarctus sordidus by Holthuis (2002).
Yang et al. (2012) performed phylogenetic analyses of
species of the family Scyllaridae and pointed out the
need for a generic re-arrangement of Scyllarinae.
Infraorder BRACHYURA Latreille, 1802
Family DROMIIDAE De Haan, 1833
Conchoecetes arti ciosus (Fabricius, 1798) (Fig. 2a)
Material examined: RUMF-ZC-1907, one male, 23.1 ×
24.7 mm.
Distribution: Western Indian Ocean to Australia;
Thailand (Gulf of Thailand); China; Taiwan; Japan
(Naiyanetr, 2007; Ahyong et al., 2009) and now
Cambodia (present study).
Remarks: Dromiid crabs usually carry a piece of
sponge, an ascidian, or a bivalve shell to cover the
dorsal surface of the cephalothorax using the last two
pairs of pereopods (except for Conchoecetes species that
use the fourth pereopods only), which are subchelate
(McLay, 1993). Species of Conchoecetes, Hypoconcha and
Desmodromia have been known to carry bivalve shells
as camou age (Wicksten, 1986; Guinot et al., 1995;
Guinot & Tavares, 2000; McLay & Hosie, 2012). Conch-
oecetes arti ciosus is unique within the genus because
of the presence of two distinct anterolateral teeth on
the carapace (McLay, 1993). It may carry bivalve shells
for camou age (Ahyong et al., 2009).
Dromidiopsis indica (Gray, 1831) (Fig. 2b)
Material examined: RUMF-ZC-1903, one male, 43.6 ×
43.9 mm, one female, 53.9 × 56.5 mm.
Distribution: Thailand (Phuket, Andaman Sea), Singa-
pore (McLay et al., 2001; Naiyanetr, 2007), and now
Cambodia (present study).
Remarks: Dromidiopsis indica is diagnosed by the
presence of three anterolateral teeth, which are close
to each other (present study).
Family DORIPPIDAE MacLeay, 1838
Dorippe quadridens (Fabricius, 1793) (Fig. 2c)
Material examined: RUMF-ZC-2318, one male, 31.0 ×
30.1 mm, one ovigerous female, 43.0 × 42.8 mm.
Distribution: Widely distributed in the Indo-West
Paci c from the Suez Canal, the Red Sea and south-
eastern Africa to Hong Kong, the Philippines,
Indonesia, Australia, Thailand (Andaman Sea and
Gulf of Thailand) (Holthuis & Manning, 1990;
Naiyanetr, 2007), and now Cambodia (present study).
© Centre for Biodiversity Conservation, Phnom Penh
Crustaceans found in by-catch
Cambodian Journal of Natural History 2014 (1) 2436
Fig. 2 Brachyuran species collected from Kampong Kandal Fishery Port: a, Conchoecetes arti ciosus (Fabricius, 1798)
(RUMF-ZC-1907, male, 23.1 × 24.7 mm); b, Dromidiopsis indica (Gray, 1831) (RUMF-ZC-1903, male, 43.6 × 43.9 mm);
c, Dorippe quadridens (Fabricius, 1793) (RUMF-ZC-2318, male, 31.0 × 30.1 mm); d, Dorippoides facchino (Herbst, 1785)
(RUMF-ZC-2319, male, 19.6 × 23.7 mm); e, Doclea armata De Haan, 1839 (RUMF-ZC-2320, male, 59.4 × 52.1 mm); f,
Halimede ochtodes (Herbst, 1783) (RUMF-ZC-1917, male, 28.2 × 37.9 mm, in live colouration).
23.7 mm, one ovigerous female, 19.9 × 25.1 mm.
Distribution: This species is known from Sri Lanka,
southern India, Thailand (Gulf of Thailand), southern
China, Vietnam, Thailand, Malaysia, Indonesia
(Holthuis & Manning, 1990), and now Cambodia
Remarks: Dorippe quadridens has been recorded to carry
various types of material for camou age (e.g. sponge,
pieces of shell or other debris): see Guinot et al. (1995).
Dorippoides facchino (Herbst, 1785) (Fig. 2d)
Material examined: RUMF-ZC-2319, one male, 19.6 ×
© Centre for Biodiversity Conservation, Phnom Penh
T. Naruse et al.
Cambodian Journal of Natural History 2014 (1) 2436
(present study).
Remarks: Dorippoides facchino often carries a shell with
a sea anemone a ached (see Guinot et al., 1995).
Family EPIALTIDAE MacLeay, 1838
Doclea armata De Haan, 1839 (Fig. 2e)
Material examined: RUMF-ZC-2320, one male, 59.4 ×
52.1 mm, one female, 57.6 × 49.3 mm.
Comparative material: Doclea armata: four males, 45.0 ×
ca. 34.4–53.4 × 42.2 mm, three females, 30.1 × 23.3 to
46.8 × 43.0 mm, three ovigerous females, 45.3 × 38.4
to ca. 49.3 × 47.2 mm, ZRC 1998.1054, Angsila shing
port, Chonburi, Thailand (Gulf of Thailand), collected
by P.K.L. Ng, 29 September 1998; six males, 46.5 × 41.2
to 56.1 × 54.2 mm, one ovigerous female, 49.5 × 46.3,
ZRC 2002.0071, Pichai shing port, Phuket, Thailand
(Andaman Sea), collected by J.C.Y. Lai, 22–25 August
Distribution: This species is known from Sri Lanka,
India, Myanmar, Singapore, Malaysia, Thailand (Gulf
of Thailand and Andaman Sea), Vietnam, Hong Kong,
the Philippines (Wagner, 1986; Naiyanetr, 2007), and
now Cambodia (present study).
Remarks: This species can be distinguished from
congeners by the absence of a spine on the supraor-
bital margin, the comparatively long rostrum, and
the presence of four longitudinal rows of dense and
rather long setae on the ambulatory legs (Wagner,
1986; Chen & Ng, 2004).
Family GALENIDAE Alcock, 1898
Halimede ochtodes (Herbst, 1783) (Fig. 2f)
Material examined: RUMF-ZC-1917, one male, 28.2 ×
37.9 mm.
Comparative material: Halimede ochtodes (Herbst, 1783):
two males, 33.2 × 46.6, 35.5 × 50.5 mm, ZRC 1998.1053,
Fig. 3 Brachyuran species collected from Kampong Kandal Fishery Port. a, Arcania novemspinosa (Lichtenstein, 1816)
(RUMF-ZC-1910, male, 25.9 × 22.9 mm); b, Leucosia formosensis Sakai, 1937 (RUMF-ZC-1911, male, 23.6 × 20.9 mm);
c, Myra a nis Bell, 1855 (RUMF-ZC-1909, male, 39.0 × 31.0 mm); d, Ixa cylindrus (Fabricius, 1777) (RUMF-ZC-1908,
male, 19.5 × 56.1 mm); e, Cryptopodia fornicata (Fabricius, 1787) (RUMF-ZC-1574, male, 35.8 × 59.8 mm); f, Rhinolambrus
longispinus (Miers, 1879) (RUMF-ZC-1912, male, 34.3 × 36.8 mm).
© Centre for Biodiversity Conservation, Phnom Penh
Crustaceans found in by-catch
Cambodian Journal of Natural History 2014 (1) 2436
Angsila shing port, Chonburi, Thailand (Gulf of
Thailand), collected by P.K.L. Ng, 29 September
1998; one male, 23.9 × 31.8 mm, 27.2 × 36.7 mm, ZRC
2012.0148, Changi Coast, Singapore, low tide, sand at
and sea grasses, collected by S.Y. Chan & L. Nguang,
7 May 2012.
Distribution: Northern half of Australia, Pakistan,
India, Thailand (Gulf of Thailand), Singapore, China
(Guangxi, Hainan, Hong Kong), the Philippines,
Indonesia (Dai & Yang, 1991; Davie, 2002; Naiyanetr,
2007), and now Cambodia (present study).
Remarks: Halimede currently includes four species (Ng
et al., 2008). Halimede ochtodes can be distinguished
from its congeners by the ill-de ned regions of the
carapace (versus well-de ned) (see Miers, 1884; Galil,
Family LEUCOSIIDAE Samouelle, 1819
Arcania novemspinosa (Lichtenstein, 1816) (Figs 3a,
4a, b)
Material examined: RUMF-ZC-1910, two males, 23.6 ×
21.0, 25.9 × 22.9 mm.
Comparative material: Arcania novemspinosa (Lichten-
stein, 1816): One male, 18.5 × 17.0 mm, RUMF-ZC-2122,
Oura Bay, Okinawa Island, Japan, -15 m, collected by
T. Fujii, 20 August 2011; three juveniles, 6.7 × 5.8 to 8.5
× 7.9 mm, RUMF-ZC-2123, Oura Bay, Okinawa Island,
-19.6 m, collected by T. Fujii, T. Naruse, D. Uyeno &
S. Nishihira, 13 November 2012; six males, 13.5 × 12.3
to 22.1 × 21.5 mm, 13 females, 13.9 × 13.2 to 27.2 × 25.2
mm, ZRC 2008.1321, Ang Sila shing port, Chonburi,
Thailand (Gulf of Thailand), collected by Z. Jaafar et al.
12 August 2003; one male, 21.3 × 18.2 mm, one female,
22.2 × 20.0 mm, ZRC 2005.0065, Aer Prang, Lembeh
Straits, Bitung, North Sulawesi, Indonesia, collected
by Tan Heok Hui, 17 July 2003.
Distribution: Andaman Sea, Malaysia, Singapore,
Thailand (Gulf of Thailand), Taiwan Strait, Okinawa
Island, Indonesia, the Philippines, Australia (Galil,
2001a; Naiyanetr, 2007; Fujii & Naruse, 2013), and
now Cambodia (present study).
Remarks: Arcania novemspinosa (Lichtenstein, 1816)
has been misidenti ed as, or regarded as possibly
being conspeci c with, A. undecimspinosa De Haan,
1841 (Alcock, 1896; Hill, 1982; Tan, 1996), but they are
clearly two di erent species. Arcania novemspinosa can
be distinguished from A. undecimspinosa by the very
long posterolateral spines of the carapace (posterolat-
eral spines distinctly longer than lateral spines versus
posterolateral spines as long as lateral spines) (Fujii
& Naruse, 2013: Fig. 3). Galil (2001a) identi ed what
Campbell & Stephenson (1970) recorded as “A. novem-
spinosa” from Moreton Bay as A. undecimspinosa, but
Naruse (2014) raised doubts as to whether it is either
species. Careful reexamination of previous records
is necessary to draw conclusions regarding the true
distribution of A. novemspinosa.
Leucosia formosensis Sakai, 1937 (Figs 3b, 4c–e)
Material examined: RUMF-ZC-1911, one male, 23.6 ×
20.9 mm,
Comparative material: Leucosia formosensis: ZRC
1984.6327–6329, one male, 22.5 × 19.0 mm, one female,
22.0 × 19.4 mm, about 30 miles from Horsburgh Light-
house, South China Sea, collected by Hee Huat, 10
September 1983.
Distribution: Singapore, Gulf of Tongking, Taiwan,
China (Guangdong and Fujian), the Philippines (Tan,
1996; Sakai, 2004; present study), and now Cambodia
(present study).
Remarks: This species is diagnosed by the G1
possessing a screw-like, longitudinally coiled shaft
with setose and swollen distal end bearing a long
slender process. Dai et al. (1986), Dai & Yan (1991),
and Tan (1996) recorded L. formosensis from the Philip-
pines. Tan (1996) noted that her record was based
on specimens deposited in the National Museum of
Natural History (Smithsonian Institution), but did not
provide detailed information.
Myra a nis Bell, 1855 (Figs 3c, 4f, g)
Material examined: RUMF-ZC-1909, one male, 39.0 ×
31.0 mm, one female, 30.9 × 26.3 mm; ZRC 2014.0338,
one male, 37.9 × 29.8 mm, one female, 28.6 × 24.3 mm.
Distribution: Sri Lanka, Indonesia, Thailand (Gulf of
Thailand), the Philippines (Galil, 2001b; Naiyanetr,
2007), and now Cambodia (present study).
Remarks: Myra a nis is characterised by a promi-
nently granulate carapace, acuminate posteromedian
carapace denticle, presence of a horizontal granulate
band on the male third thoracic sternite, cheliped
dactylus about three-quarters as long as the upper
margin of the palm, and male cheliped merus being
shorter than the carapace length (Galil, 2001b).
Ixa cylindrus (Fabricius, 1777) (Figs 3d, 4h, i)
Material examined: RUMF-ZC-1908, two males, 12.7 ×
32.8, 19.5 × 56.1 mm, one female, 19.3 × 52.7 mm; ZRC
2014.0339, one male, 18.7 × 52.5 mm, one female, ca. 19
× ca. 60 mm.
© Centre for Biodiversity Conservation, Phnom Penh
T. Naruse et al.
Cambodian Journal of Natural History 2014 (1) 2436
Distribution: Widely distributed. Known from East
Africa, Mauritius, India, Sri Lanka, Singapore,
Thailand (Gulf of Thailand), Cambodia, Malaysia
(Borneo), China, the Philippines, Taiwan, Australia
(Komatsu et al., 2004; Naiyanetr, 2007; Takeda &
Nagai, 2009; UNEP, 2007).
Remarks: Among the nine known species of Ixa Leach,
1816 (Ng et al., 2008; also see Komatsu et al., 2004,
Takeda & Nagai 2009), I. cylindrus can be easily distin-
guished by the presence of a pair of sharp-edged
narrow and deep grooves that run longitudinally
along the sides of the gastric and cardiac regions to the
intestinal regions. Komatsu et al. (2004) indicated that
there is considerable variation in the condition of the
lateral process of the carapace. Takeda & Nagai (2009)
noted that the geographical ranges of I. cylindrus and I.
edwardsii Lucas, 1858, reach Mersin Bay, southeastern
Turkey, in the Mediterranean Sea. This information
appears to be cited from Holthuis & Go lieb (1956).
However, the only Ixa species recorded from the
Mediterranean Sea by Holthuis & Go lieb (1956) was
their new species, I. monodi. Since Takeda & Nagai
(2009) regarded I. monodi as a valid species, it is likely
they cited Holthuis & Go lieb (1956) in error.
Fig. 4 Male rst and second gonopods of leucosiid species collected from Kampong Kandal Fishery Port: a, b, Arcania
novemspinosa (Lichtenstein, 1816) (RUMF-ZC-1910, 25.9 × 22.9 mm) (a, left G1, ventral view; b, left G2, ventral view); c–e,
Leucosia formosensis Sakai, 1937 (RUMF-ZC-1911, 23.6 × 20.9 mm) (c, left G1, ventral view; d, distal part of left G1, mesial
view; e, right G2, ventral view); f, g, Myra a nis Bell, 1855 (RUMF-ZC-1909, 39.0 × 31.0 mm) (f, left G1, ventral view; g,
left G2, ventral view); h, i, Ixa cylindrus (Fabricius, 1777) (RUMF-ZC-1908, 19.5 × 56.1 mm) (h, left G1, ventral view; i, left
G2, ventral view). Scales = 1 mm.
© Centre for Biodiversity Conservation, Phnom Penh
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Cambodian Journal of Natural History 2014 (1) 2436
Family PARTHENOPIDAE MacLeay, 1838
Cryptopodia fornicata (Fabricius, 1787) (Figs 3e, 5a,
Material examined: RUMF-ZC-1574, one male, 35.8 ×
59.8 mm.
Distribution: Central Indian Ocean, Thailand (Gulf of
Thailand), Cambodia, South China Sea, Japan (Chiong
& Ng, 1998; Naiyanetr, 2007; UNEP, 2007).
Remarks: Cryptopodia fornicata can be distinguished
from all congeners by the following combination of
morphological characters: entire posterior to postero-
lateral margins of the carapace; rostrum being broader
than long, with convex lateral margins; and smooth
pterygostomial region and outer surface of third
maxilliped (Chiong & Ng, 1998).
Rhinolambrus longispinus (Miers, 1879) (Figs 3f, 5c,
Material examined: RUMF-ZC-1912, one male, 34.3 ×
36.8 mm.
Comparative material: Rhinolambrus longispinus: one
male, 38.3 × 38.8 mm, ZRC 1984.6380, about 30
Fig. 5 Male rst and second gonopods of parthenopid species collected from Kampong Kandal Fishery Port: a, b,
Cryptopodia fornicata (Fabricius, 1787) (RUMF-ZC-1574, 35.8 × 59.8 mm) (a, left G1, ventral view; b, left G2, ventral view);
c, d, Rhinolambrus longispinus (Miers, 1879) (RUMF-ZC-1912, 34.3 × 36.8 mm) (c, left G1, ventral view; d, left G2, ventral
view). Scale = 1 mm.
© Centre for Biodiversity Conservation, Phnom Penh
T. Naruse et al.
Cambodian Journal of Natural History 2014 (1) 2436
miles from Horsburgh Lighthouse, South China Sea,
collected by Hee Huat, 10 September 1983; one male,
39.5 × 40.7, ZRC 1984.6068, Singapore. Rhinolambrus
contrarius (Herbst, 1804): one male, 45.3 × 42.6 mm,
ZRC 1999.0058, Kii Peninsula, Japan, collected by
NSMT, 27 Feb. 1999.
Distribution: Amirante Islands, Seychelles,
Arrakanküste, Maldives, Sri Lanka, Cambodia, Singa-
pore, Thailand (Gulf of Thailand), China (Shanghai),
Sabira, Kepulauan Seribu, Sulawesi, and Torres Strait,
Australia (Flipse, 1930; Naiyanetr, 2007; UNEP, 2007).
Remarks: The present specimen agrees well with
the original description of the species (Miers, 1879).
Shen et al. (1982) recorded R. longispinus [as Parthe-
nope (Rhinolambrus) longispinus] from Hainan Island,
China. However, their photographed specimen (Shen
et al., 1982: pl. 1–4) had a relatively long and sharp
rostrum, and long cheliped and carapace spines.
These Chinese specimens most probably belong to a
di erent species.
Family PORTUNIDAE Ra nesque, 1815
Podophthalmus vigil (Fabricius, 1798) (Fig. 6a)
Material examined: RUMF-ZC-1914, one male, 26.4 × ca.
57.3 mm.
Distribution: Western Indian Ocean including the Red
Sea to Malaysia, Cambodia, China (Guangxi, Hainan),
Taiwan, Japan, Australia, Hawaii, Samoa and Tahiti
(Dai & Yang, 1991; Naiyanetr, 2007; UNEP, 2007).
Remarks: The genus Podophthalmus Lamarck, 1801,
contains three species. Podophthalmus vigil di ers from
its two congeners in the following combination of
characters: eyestalk entire (without pterygoid expan-
sion) and carapace broad (width ca. 2.5 times carapace
length) (Stephenson, 1972).
Charybdis (Charybdis) anisodon (De Haan, 1850)
(Fig. 6b)
Material examined: RUMF-ZC-1915, one male, 31.2 ×
53.7 mm.
Distribution: Madagascar, Red Sea, Thailand
(Andaman Sea and Gulf of Thailand), Cambodia,
Malaysia, Singapore, Indonesia (Sulawesi), China,
Taiwan, Japan, the Philippines, Australia and New
Caledonia (Wee & Ng, 1995; Naiyanetr 2007; UNEP,
Remarks: Charybdis anisodon is characterized by the
olive-coloured dorsal surface of the carapace. Identi-
cation keys to species of Charybdis were provided by
Stephenson (1972), Wee & Ng (1995) and Apel & Spiri-