FORSCHUNGS

museum

KOENIG

Bonn

zoological

Bulletin

formerly: Bonner zoologische Beitrage

Volume 63

Issue 1

2014

H. eni

H. kyaboboen sis

tus

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Bonn zoological Bulletin

Editor-in-Chief

Fabian Herder, Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK), Ichthyology Section, Adenauerallee 160,

53113 Bonn, Germany, tel. +49 228-9122-255,

fax: +49 228-9122-212; f.herder@zfmk.de

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Dirk Ahrens, Insects: Coleoptera, ZFMK,

tel. +49 228-9122-286; d.ahrens@zfmk.de

Wolfgang Boh me, Amphibians and Reptiles, ZFMK,

tel. +49 228-9122-250; w.boehme@zfmk.de

Jan Decher, Mammals, ZFMK, tel. +49 228-9122-262;

j.decher@zfmk.de

Netta Borchin, Insects: Diptera, Department of Zoology, Tel

Aviv University, Tel Aviv 69978, Israel,

tel. +972-3-6409808; ndorchin@post.tau.ac.il

Bernhard Huber, Invertebrates except Insects, ZFMK,

tel. +49 228-9122-294; b.huber@zfmk.de

Rainer Hutterer, Mammals, ZFMK, tel. +49 228-9122-261;

r.hutterer@zfmk.de

Ximo Mengual, Insects: Diptera, ZFMK,

tel. +49 228-9122-292; x.mengual@zfmk.de

Gustav Peters, Mammals, ZFMK, tel. +49 228-9122-279;

g.peters@zfmk.de

Ralph Peters, Insects: Hymenoptera, ZFMK,

tel +49 228-9122-290; r.peters@zfmk.de

Bradley Sinclair, Canadian National Collection of Insects,

Ottawa Plant Laboratory - Entomology, CFIA, K.W. Neat-

by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada

K1A0C6, tel. + 1 613-759-1787;

bradley.sinclair@inspection.gc.ca

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ZFMK, tel. +49 228-9122-220;

d.stuening.zfmk@uni-bonn.de

Till Topfer, Birds, ZFMK, tel. +49 228-9122-246;

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Philipp Wagner, Amphibians and Reptiles, ZFMK,

tel. +49 228 9122-254; philipp.wagner.zfmk@uni-bonn.de

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niz-Institut fur Evolutions- und Biodiversitatsforschung an

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10115 Berlin, Germany, tel. +49 30-2093-8504/ 8400,

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ment of Entomology, Cromwell Road, London, SW7 5BD,

U.K.; j.holloway@nhm.ac.uk

Tan Heok Hui, Raffles Museum of Biodiversity Research,

National University of Singapore, Department of Biological

Sciences, 6 Science Drive 2, #03-01, Singapore 117546,

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PO. Box 290, Ljubljana, Slovenia;

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Rosenstein 1, 70191 Stuttgart, Germany,

tel. +49 711-8936-221, fax: +49 711-8936-100;

schawaller.smns@naturkundemuseum-bw.de

Ulrich K. Schliewen, Department of Ichthyology, Bavarian

State Collection of Zoology, Miinchhausenstr. 21, 81247

Miinchen, Germany, tel. + 49 89-8107-1 10;

sch 1 iewen@zsm . mwn . de

Michael Schmitt, Ernst-Moritz-Amdt-Universitat, Allge-

meine & Systematische Zoologie, Anklamer Str. 20, 17489

Greifswald, Germany, tel. +49 3834-86-4242, fax: +49

3834-86-4098; michael.schmitt@uni-greifswald.de

W. David Sissoni, Dept, of Life, Earth and Environmental

Sciences, W. Texas A. & M. University, WTAMU Box

60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578,

fax: +1 806-651-2928; dsissom@mail.wtamu.edu

Miguel Vences, Technical University of Braunschweig, Zoo-

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Germany, tel. +49 531-391-3237, fax: +49 3 1-391-8198;

m.vences@tu-bs.de

Erich Weber, Eberhard-Karls-Universitat, Zoologische

Schausammlung, Sigwartstr. 3, 72076 Tubingen,

tel. +49 7071-2972616, fax +49 7071-295170;

erich.weber@uni-tuebingen.de

Bonn zoological Bulletin 63 (1): 1-14

June 2014

Description of four new West African forest geckos

of the Hemidactylus fasciatus Gray, 1842 complex,

revealed by coalescent species delimitation

Philipp Wagner' , Adam D. Leache2 & Matthew K. Fujita’

' Department of Biology, Villanova University, 800 Lancaster Avenue, ViUanova, Pennsylvania 19085, USA and Zoologisches

Forschungsmuseum A. Koenig, Adenauerallee 160, D53113 Bonn, Germany.

■ -Department ofBiolog\> & Burke Museum of Natural History and Culture, University of Washington, Seattle, WA 98195, USA.

Department of Biology, University of Texas at Arlington, 501 S. Nedderman Drive, Box 19498, Arlington, TX 76019-0498, USA

"Corresponding Author

Abstract. The gecko Hemidactylus fasciatus is widespread in rainforest regions of equatorial Africa, from Guinea to

Cameroon. Recently, this taxon was identified as a cryptic complex of at least five species, using multilocus genetic da-

ta and coalescent models for species delimitation. Here, we provide the formal descriptions of four new species from

tropical West and Central Africa. As typical for cryptic species, the new species are genetically distinct, but difficult to

distinguish using external morphology. However, coloration, shape of the body crossbands, and body size, are important

distinguishing characters for this complex. We provide a new taxonomy for this complex that includes the following for-

est gecko species: H. fasciatus is now restricted to West Africa occurring eastwards to the Dahomey Gap, H. kyaboboen-

sis sp. n. is known only from within the Dahomey Gap, H. eniangii sp. n. is distributed from the Dahomey Gap to west-

ern Cameroon, H. coalescens sp. n. occurs from central Cameroon to southern Gabon, H. biokoensis sp. n. is restricted

to Bioko Island, and H. ituriensis , herein recognized as full species, is known from several localities in eastern Africa.

Key words. Africa, rainforest, Sauria, Gekkonidae, Hemidactylus fasciatus complex, Hemidactylus ituriensis.

INTRODUCTION

An accurate inventory of species diversity forms the foun-

dations for scientific inquiries in biogeography, phylo-

geography, ecology, conservation, and comparative biol-

ogy. However, a fundamental problem is that a large pro-

portion of species remain scientifically undocumented, es-

pecially those that are not easily distinguishable based on

casual examinations of external morphology or coloration.

Bickford et al. (2006) defined cryptic species as two or

more species that have been recognized as a single nom-

inal species, because they are at least superficially indis-

tinguishable based on external morphology. The uncov-

ering of cryptic species is important for accurate species

inventory, and for providing new opportunities to study

mechanisms of speciation, biogeography, mate recogni-

tion, and conservation management (Bickford et al. 2006).

Molecular data and coalescent theory provide useful tools

for species delimitation in cryptic species assemblages

(Fujita et al. 2012).

The speciation history of Hemidactylus fasciatus Gray,

1842 was recently investigated by Leache and Fujita

(2010) using five nuclear loci, and by Leache et al. (2014)

using single nucleotide polymorphism (SNP) data from

over 1,000 loci. Both studies used coalescent-based

species delimitation methods to statistically test alterna-

tive species delimitation models. The species diversity in

Received: 17.09.2013

Accepted: 21.01.2014

H. fasciatus was found to be underestimated in both stud-

ies, and revealed four new genetically distinct species. De-

tails on the methods are presented by Leache & Fujita

(2010) and Leache et al. (2014). Bauer et al (201 1 ) iden-

tified insufficiencies in the diagnoses according to the In-

ternational Code of Zoological Nomenclature (ICZN

1999, subsequently named 'the code’), and argued that the

names proposed by Leache and Fujita (2010) were nomen

nuda. Here, we follow the recommendations of Bauer et

al. (201 1) and Fujita & Leache (201 1), and provide for-

mal descriptions for these new species of African forest

geckos that meet the criteria of the code (ICZN 1999).

The distributions of the proposed species match the con-

temporary major blocks of rainforests in equatorial

Africa (Wagner et al. 2008; Wagner et al. 2009), suggest-

ing that allopatric divergence, as a result of habitat frag-

mentation, can explain the formation of these lineages

(Leache & Fujita 20 1 0; Wagner et al. 2009; Wagner 20 1 0;

Leache et al. 2014).

SPECIES ACCOUNT

The Hemidactylus fasciatus species group is distinct by

its characteristic coloration to all other African Hemidacty-

Corresponding editor: F. Herder

2

Philipp Wagner et al.

Fig. 1. [A] Living specimen of Hemidactylus fasciatus (not collected) from Liberia; [B] Living specimen of Hemidactylus kyaboboen-

sis sp. n. from the type locality.

lus species, and is therefore not compared to them in the

present study.

Specimens, including type specimens, from the follow-

ing institutions were examined for the present study: Acad-

emy of Natural Sciences at Drexel University (ANSP),

Philadelphia, PA, USA. American Museum of Natural His-

Bonn zoological Bulletin 63 (1): 1-14

tory (AMNH), New York, NY, USA. Natural History Mu-

seum (BMNH), England, UK. California Academy of Sci-

ences (CAS), San Francisco, CA, USA. Museum of Ver-

tebrate Zoology (MVZ), Berkeley, CA, USA. Zoologi-

sches Forschungsmuseum Alexander Koenig (ZFMK),

Bonn, Germany.

©ZFMK

Description of New African Forest Geckos

3

Hemidactylus fasciatus Gray, 1842

1842 Hemidactylus fasciatus Gray, Description of some

new species of Reptiles, chiefly from the British Muse-

um collection. The Zoological Miscellany: 57-59.

1845 Leiurus oniatus Gray (fide Loveridge 1947), Cata-

logue of the specimens of lizards in the collection of the

British Museum. Trustees of die British Museum/Edward

Newman, London: xxvii + 289 pp. [Holotype in BMNH,

from “W. Africa”].

1857 Hemidactylus formosus Hallowed, ( fide Loveridge

1 947) Notes on the reptiles in the collection of the muse-

um of the Academy of Natural Sciences. Proc. Acad. Nat.

Sci. Philadelphia, 8 (4): 146-153 [Syntypes: ANSP 745 1 -

53; Liberia].

Holotype. BMNH XXI.24.a, from an unknown locality

(fide Gray 1842).

Description. A fairly large species of Hemidactylus , with

a maximum SVL of 95 mm, and a maximum total length

of 172 mm. Like in the other species of the group, the body

and limbs are moderate; the snout is slightly longer than

the distance between the eye and the ear opening, the ear-

opening is suboval and oblique; the upper surface of the

body is covered with small granules, which are intermixed

with small, round, and convex tubercle scales, forming

more than 20 less regular longitudinal rows; digits are dis-

tinctly webbed at the base, and have six to eight lamellae

under the inner digits, and nine to ten under the median

ones. Like other species of the group, excluding H. iturien-

sis , it is morphologically characterized by a long row of

femoral pores in males, which are interrupted in the mid-

dle, resulting in a high number of femoral pores ( 16-20,

vs. 8 in H. ituriensis ) on each side. In coloration it is dis-

tinct from its close relative H. kyaboboensis sp. n., by lack-

ing the various pale tubercle spots and the broad pale fram-

ing of the dark body crossbands, and by having the dark

body crossbands more distinct to the pale interspaces (see

Fig. 1). It is distinct from other species of the group by

having a broad dark band between the eyes and the neck

(vs. a narrow stripe in H. coalescens sp. n. and vs. lack-

ing in H. ituriensis), and by having a pale upper lip (vs.

dark in H. eniangii sp. n.).

Distribution. We herein recognize the distribution of H.

fasciatus as restricted to the Guinean rainforest system

west of the Dahomey Gap. Therefore, it is known from

localities in Guinea, Liberia, Cote d’Ivoire, and Ghana.

Relationships. Genetic data was presented by Leache &

Fujita (2010), who showed that this species includes all

populations that cluster with those from the western

Guinean rainforest with strong support in the Bayesian

species delimitation model. This species is closely relat-

ed to the other West African species described herein (H.

kyaboboensis sp. n.), which is so far only known from

forests within the Dahomey Gap.

Hemidactylus kyaboboensis sp. n.

2010 Hemidactylus kyaboboensis Leache & Fujita (nom.

nud. fide Bauer et al. 2010), Bayesian species delimita-

tion in West African forest geckos (Hemidactylus fascia-

tus). Proc Roy Soc B 277: 3071-3077.

Holotype. MVZ 245291 (Fig. 2), adult male from Ghana,

Volta Region, Togo Hills, Kyabobo National Park, Water-

fall, 08.3301 9° N, 00.5941 1° E, 5 1 5 m, collected by Adam

D. Leache, Raul Diaz and Matthew K. Fujita on 16 June

2004.

Paratypes. MVZ 245292-245299, same data as the holo-

type. ZFMK 93689, subadult from Togo, Missahohe

[=Missahoe or Missahohe], near Kpalime. ZFMK 19922,

adult female from Faille de Bafilo [=fault proper near Bafi-

lo (9.282399, 1.216621)], Togo.

Diagnosis. A large species of Hemidactylus, with a max-

imum SVL of 80 mm, and a maximum total length of 160

mm, a broad head (vs. slender in H. coalescens). and three

enlarged intemasal scales (vs. two in H. eniangii ). Char-

acteristic is the dorsal scalation of irregular arranged

smooth and convex tubercle scales, each circumdated by

a ring of small granular scales. In coloration, it is unique

by having indistinct dark body crossbands, and more

prominent whitish stripes and dots. This new species is

distinct in coloration in detail to H. fasciatus by having a

band on the side of the head as broad as the crossband on

the neck, and reaching the lower tip of the ear hole (vs.

not reaching the lower tip). It is distinct from H. coa-

lescens by having the broad band on the side of the head

(extension of the crossband on the neck) as broad as the

band on the neck (vs. a crossband on the neck becoming

a narrow stripe on the side of the head); by having the last

indistinct body crossband in contact with the hindlimbs

(vs. a distinct crossband not in contact); by having the first

indistinct body crossband reaching the head skull (vs. a

distinct first body crossband restricted to the neck). It is

distinct from H. eniangii by having a band on the side of

the head as broad as the crossband on the neck (vs. a cross-

band becoming narrower on the side of the head).

Description of the holotype. Habitus stout, head elongat-

ed. SVL length 79.2 mm, tail length 77.4 mm (posterior

60 % regenerated). Snout elongate, distance from anteri-

or margin of the tympanum to hind margin of the eye 4.6

mm, from anterior margin of the eye to tip of snout 9.3

mm; eye diameter 4.5 mm. Rostral broad, with a notch on

Bonn zoological Bulletin 63 (1): 114

©ZFMK

4

Philipp Wagner et al.

Fig. 2. The holotype of (MVZ 245291 ) Hemidactylus kyaboboensis sp. n.

Bonn zoological Bulletin 63 (1): 1-14

©ZFMK

Description of New African Forest Geckos

5

its anterior midline, which continues as a vertical suture,

but does not separate the rostral scale. Nostril bordered

by the rostral, the 1st labial and two to three postnasal

scales, three enlarged internasal scales present. Granular

scales on the snout small, smooth and convex, larger as

the scales on top of the head of the frontal and parietal

region. Head scales become intermixed with tubercle

scales at the posterior part of the neck. Temple covered

with convex scales, with only few intermixed enlarged tu-

bercle scales. Mouthline straight, but uplifted at its hind-

part. Supralabials (left/right) 10/12, sublabials 10/10. Men-

tal scale triangular, followed by a single pair of postmen-

tal scales. Gulars small, smooth, passing posteriorly grad-

ually into the rhombic shaped, imbricate and smooth chest

and belly scales. The convex and smooth dorsal tubercles

are arranged irregularly, and are circumdated by a ring of

granular scales. There is no or only one row of granular

scales between these rings. Limbs covered with coarse

granular scales, and intermixed with flat and smooth larg-

er tubercle scales. Palmar scales smooth. All fingers and

toes with distinct claws and with divided adhesive lamel-

lae beneath, except the terminal subdigital scales, which

are undivided like the basal ones.

Coloration in preservation. Upper side uniform grey, with

darker crossbands from neck to tail. One band on the neck,

three between the limbs with the middle band broken in-

to two smaller bands, one on the tail base and two on the

unregenerated portion of the tail. Tubercle scale pale

whitish possessing a spotted pattern. Underside uniform

dirty white.

Distribution. This species is distributed in the Togo Hills

in Ghana, and in the forests of Missahohe in Togo.

Habitat. At Togo Hills, specimens were collected in moist

semi-deciduous rainforest, while the specimen at Mis-

sahohe was found in an old stone building within moist

semi-deciduous rainforest.

Relationships. This species includes all populations that

cluster with those presented by Leache & Fujita (2010)

from the Togo Hills with strong support in the Bayesian

species delimitation model. It is the sister taxon of H.fas-

ciatus from the Guinean rainforest system in West Africa

(Leache & Fujita 2010; Leache et al., 2014).

Etymology’. This species is named after its type locality,

the Kyabobo National Park, Togo Hills, Volta Region,

Ghana.

Hemidactylus eniangii sp. n.

2010 Hemidactylus eniangii Leache & Fujita (nom. nud.

fide Bauer et al. 2010), Bayesian species delimitation in

West African forest geckos (Hemidactylus fasciatus). Proc

Roy Soc B 277: 3071-3077.

Holotype. MVZ 253215 (Fig. 3), adult male from Nige-

ria, Cross River State, Cross River National Park, Oban

Hills Sector, Southern Portion, Erokut Park entry gate,

05.3639° N, 08.43341° E, 143 m, collected by Adam D.

Leache, Anne M. Leache, and Edem A. Eniang on 6 April

2006.

Paratypes. MVZ 253213, 253214, same data as the holo-

type.

Diagnosis. A medium sized species of Hemidactylus , with

a maximum SVL of 60 mm, a maximum total length of

about 140 mm, and a broad head (vs. slender in H. coa-

lescens sp. n.). Characteristic is the dorsal scalation of ir-

regular arranged smooth and convex tubercle scales, each

encircled by a ring of small granular scales, and having

two enlarged intemasal scales (vs. three in all other

species). In coloration, it is distinct from H. fasciatus by

having a dark upper lip (vs. a pale one), and in having the

band on the side of the head narrower than the crossband

on the neck (vs. as broad as the crossband); from H. co-

alesces sp. n. by having the last body crossband in con-

tact with the hindlimbs (vs. not in contact), having the first

body crossband starting on the hind part of the head skull

(vs. restricted to the neck), and by having a narrow band

(vs. a stripe as extension of the neck crossband on the side

of the head); and from H. k\>aboboensis sp. n. by having

distinct body crossbands (vs. indistinct bands) and stripes,

and by having the extension band on the side of the head

narrower than the crossband on the neck (vs. as broad as

the crossband).

Description of the holotype. Habitus stout, head elongat-

ed. SVL length 61.5 mm, tail length 71 mm (posterior 20

% regenerated). Snout elongate, distance from anterior

margin of the tympanum to hindmost margin of the eye

4.8 mm, from anterior margin of the eye to tip of snout

6.7 mm; eye diameter 3.7 mm. Rostral broad, with a notch

on its anterior midline which continues as a vertical su-

ture but does not separate the rostral scale. Nostril bor-

dered by the rostral, the 1st labial and two to three post-

nasal scales, two enlarged internasal scales present. Gran-

ular scales on the snout small, smooth and convex, larg-

er as the scales on top of the head of the frontal and pari-

etal region. Head scales become intermixed with tuber-

Bonn zoological Bulletin 63 (1): 1-14

©ZFMK

6

Philipp Wagner et al.

Fig. 3. The holotype (MVZ 253215) of Hemidactylus eniangii sp. n.

Bonn zoological Bulletin 63 (1): 1-14

©ZFMK

Description of New African Forest Geckos

7

cle scales at the posterior part of the neck. Temple cov-

ered with convex scales, with only few intermixed en-

larged tubercle scales. Mouthline straight, but uplifted at

its hindpart. Supralabials (left/right) 12/12, sublabials

10/10. Mental scale triangular, followed by a single pair

of postmental scales. Gulars small, smooth, passing pos-

teriorly gradually into the rhombic shaped, imbricate and

smooth chest and belly scales. The convex and smooth

dorsal tubercles are arranged irregularly, and are encircled

by a ring of granular scales. There is no or only one row

of granular scales between these rings. Limbs covered with

coarse granular scales and intermixed with flat and smooth

larger tubercle scales. Palmar scales smooth. All fingers

and toes with distinct claws and with divided adhesive

lamellae beneath, except the terminal subdigital scales

which are undivided like the basal ones.

Coloration in preservation. Upper side uniform grey, with

darker crossbands from neck to tail. One band on the neck,

three between the limbs, one on the tail base and three on

the unregenerated portion of the tail. Tubercle scale pale

whitish possessing a spotted pattern. Underside uniform

dirty white.

Distribution. This species is distributed in the northern

portion of the Congolian rainforest north of the Sanaga

River, including northern Cameroon, and Nigeria.

Habitat. This is a nocturnal species. Mertens ( 1 938) men-

tioned the gecko as very common in rainforests of lower

elevations (400-550m) at Mount Cameroon, but also

found one specimen in a small hut outside the forest.

Relationships. This species includes all populations that

cluster with those presented by Leache & Fujita (2010)

from the northern portion of the Congolian rainforest (east-

ern Nigeria and northern Cameroon) with strong support

in the Bayesian species delimitation model. The SNP loci

used by Leache et al. (2014) provide strong support plac-

ing this taxon sister to H. coalescens sp. n. and H. biokoen-

sis sp. n. This topology contradicts the Leache and Fuji-

ta (2010) study, which suggests a close relationship be-

tween H. eniangii sp. n. and H. biokoensis sp. n.

Etymology. This species is named in honor of the Niger-

ian conservation biologist and herpetologist Dr. Edem A.

Eniang for his studies on the Nigerian herpetofauna.

Hemidactylus coalescens sp. n.

2010 Hemidactylus coalescens Leache & Fujita (nom.

nud. fide Bauer et al. 2010), Bayesian species delimita-

tion in West African forest geckos (Hemidactylus fascia-

tus). Proc Roy Soc B 277: 3071-3077.

Holotype. ZFMK 87679 (Fig. 4), adult male from

Cameroon, Campo Region, Nkoelon, 2.3972° N,

1 0.045 1 5° E, 85 m, collected by Michael F. Barej and Ju-

lia Wurstner on 27 October 2007.

Paratvpes. ZFMK 87680-82, same data as the holotype.

Diagnosis. A medium sized species of Hemidactylus , with

a max. SVL of 78 mm, a maximum total length of 174

mm, a more elongated head than the other species of the

H. fasciatus group, and three enlarged internasal scales

present in most specimens. It is unique in coloration by

possessing body crossbands which are more narrow than

the pale interspaces; the interspaces contain indistinct dark

bands which are not connected at the vertebrate; by hav-

ing the first body crossband restricted to the neck (vs.

reaching the cranium as in the other species); by having

the last crossband on the body between the legs not in con-

tact with the hindlimbs (vs. in contact); and by having a

distinctly fine narrow dark stripe on the side of the head

between eye and ear, as the extension of the crossband on

the neck (vs. a band slightly narrower or as broad as the

crossband in the other species).

Description of the holotype. Habitus stout, head elongat-

ed. SVL length 59.2 mm, tail length 76.2 mm. Snout elon-

gate, distance from anterior margin of the tympanum to

hindmargin of the eye 4.6 mm, from anterior margin of

the eye to tip of snout 5.4 mm; eye diameter 3.3 mm. Ros-

tral broad, with a notch on its anterior midline which con-

tinues as a vertical suture but does not separate the ros-

tral scale. Nostril bordered by the rostral, the 1st labial and

two to three postnasal scales, three enlarged intemasal

scales present. Granular scales on the snout small,

smooth and convex, larger as the scales on top of the head

of the frontal and parietal region. Head scales become in-

termixed with tubercle scales at the posterior part of the

neck. Temple covered with convex scales, with only few

intermixed enlarged tubercle scales. Mouthline straight,

but uplifted at its hindpart. Supralabials (left/right) 12/12,

sublabials 10/10. Mental scale triangular, followed by a

single pair of postmental scales. Gulars small, smooth,

passing posteriorly gradually into the rhombic shaped, im-

bricate and smooth chest and belly scales. The convex and

smooth dorsal tubercles are arranged irregularly and are

encircled by a ring of granular scales. There is no or on-

ly one row of granular scales between these rings. Limbs

covered with coarse granular scales and intermixed with

flat and smooth larger tubercle scales. Palmar scales

smooth. All fingers and toes with distinct claws and with

divided adhesive lamellae beneath, except the terminal

subdigital scales which are undivided like the basal ones.

Coloration in preservation. Upper side uniform light

brownish, with darker crossbands from neck to tail tip.

Bonn zoological Bulletin 63 (1): 1-14

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Philipp Wagner et al.

Fig. 4. The holotype (ZFMK 87679) of Hemidactylus coalescens sp. n.

Bonn zoological Bulletin 63 ( 1 ): 114

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Description of New African Forest Geckos

9

One band on the neck, three between the limbs, one on

the tail base and six on the tail. Tubercle scale pale whitish

possessing a spotted pattern. Underside uniform dirty

white.

Distribution. This species is distributed in the southern

portion of the Congolian rainforest south of the Sanaga

River, including southern Cameroon, Gabon and Congo.

Relationships. This species includes all populations that

cluster with those from the southern portion of the Con-

golian rainforest (southern Cameroon, Gabon and Congo)

presented by Leache & Fujita (2010), with strong support

in the Bayesian species delimitation model. The SNP loci

used by Leache et al. (20 1 4) provide strong support plac-

ing this taxon in a clade with H. biokoensis sp. n.

Etymology. This species is named after the coalescent

process used to delimit the species.

Hemidactylus biokoensis sp. n.

Holotype. ZFMK 9342 (Fig. 5), adult female from Ureca

on Bioko Island, Equatorial Guinea, collected by Martin

Eisentraut between 17.-30. January 1963.

Parah’pes. CAS 207624, Equatorial Guinea, Bioko Island,

near Malabo (03° 42’ 39.8” N, 08° 39’ 59.9” E ), collect-

ed by L.G. Henwood and J.V. Vindum on 18 October

1998; CAS 207663-4, Equatorial Guinea, Bioko Island,

road S of Luba (03° 28’ 06. 1 ” N, 08° 29’ 34.2” E), col-

lected by L.G. Henwood and J.V. Vindum on 14 October

1998; CAS 207777-8, Equatorial Guinea, Bioko island,

ca 3.6 km N (by road) of Luba (03° 28’ 58.9” N, 08° 34’

55.2” E); ZFMK 9339-49, same data as the holotype.

Diagnosis. A medium sized, broad headed species of

Hemidactylus , with a maximum SVL of 80.6 mm and a

maximum total length of 166.7 mm. It is overall similar

to H. coalescens sp. n. in its morphological characters.

Characteristic is the dorsal scalation of pale, smooth and

convex tubercle scales irregularly arranged in longitudi-

nal rows, and distinct to the dark body crossbands in col-

oration and the dark bands on the lower side of the tail.

In coloration, it is distinct from H.fasciatus by having pale

dots on the dark body crossbands (vs. uniform dark cross-

bands), and by lacking fine, dark stripes within the inter-

spaces between the crossbands, and by having the inter-

spaces slightly broader than the crossbands (vs. crossbands

slightly broader than the interspaces); it is distinct from

H. kyaboboensis sp. n. by having distinct body crossbands

(vs. indistinct bands and stripes), and by having the ex-

tension band on the dorsolateral side of the head narrow-

er than the crossband on the neck (vs. as broad as the

crossband); it is distinct from H. eniangii sp. n. by hav-

ing the upper lip dirty white (vs. brownish), and by hav-

ing the first body crossband broader at the median of the

neck than on the lateral parts of the neck (vs. broader on

the lateral sides); it is distinct from H. coalescens sp. n.

by having the band between eye and neck as broad as the

crossband on the neck and half the size of the eye height

(vs. having a narrow stripe a quarter the size of the eye

height), by having the interspaces slightly broader than the

crossbands (vs. distinctly broader than the crossbands), by

lacking the dark stripe in the center of each interspace be-

tween the limps which is present in H. coalescens sp. n.,

and by having the first body crossband starting on the hind

part of the head skull (vs. restricted to the neck); and it is

distinct to H. ituriensis by having a dark band as exten-

sion of the neck crossband between the eye and the neck

(vs. lacking this band), by having interspaces which are

slightly broader than the body crossbands (vs. body cross-

bands which are distinctly broader than the interspaces),

and by having body crossbands with a median notching

(vs. without it).

Description of the holotype. Habitus stout, head elongat-

ed. SVL length 66.4 mm, tail length 79.9 mm. Snout elon-

gate, distance from anterior margin of the tympanum to

hind margin of the eye 5.5 mm, from anterior margin of

the eye to tip of snout 7.6 mm; eye diameter 4.5 mm. Ros-

tral broad, with a notch on its anterior midline which con-

tinues as a vertical suture but does not separate the ros-

tral scale. Nostril bordered by the rostral, one small and

one elongated postnasal scale, in contact with the inter-

nasal at its upper tip and with the 1st labial scale on its

lower tip, three enlarged internasal scales present. Gran-

ular scales on the snout small, smooth and strongly con-

vex, larger as the scales on top of the head of the frontal

and parietal region. Head scales become intermixed with

tubercle scales just behind the eyes. Temple covered with

convex scales, with only few intermixed enlarged tuber-

cle scales. Mouthline straight, but uplifted at its hindpart.

Supralabials (left/right) 9/10, sublabials 9/9. Mental

scale triangular, followed by a single pair of postmental

scales. Gulars small, smooth, passing posteriorly gradu-

ally into the rhombic shaped, imbricate and smooth chest

and belly scales. Dorsal tubercle scales numerous, con-

vex and smooth, arranged in irregular longitudinal rows

and circumdated by a ring of granular scales. No or only

one row of granular scales between these rings on the up-

per side of the body. Limbs covered with coarse granular

scales, and intermixed with flat and smooth larger tuber-

cle scales. Palmar scales smooth. All fingers and toes with

distinct claws and with divided adhesive lamellae beneath,

except the terminal subdigital scales which are undivid-

ed.

Bonn zoological Bulletin 63 ( 1 ): 114

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10

Philipp Wagner et al.

Fig. 5. The holotype (ZFMK 9342) of Hemidactylus biokoensis sp. n.

Bonn zoological Bulletin 63 (1): 1-14

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Description of New African Forest Geckos

1 1

Fig. 6. [A] Living specimen of Hemidactylus ituriensis (AMNH 10273) from Akenge (obtained from Schmidt 1919, plate XVI);

[B] living specimen of Hemidactylus coalescens sp. n. (ZFMK 87679, holotype) from the type locality; and [C] uncollected liv-

ing specimen of Hemidactylus biokoensis from Bioko Island (Reserva cientifica de la Caldera de San Carlos, 3°14’2.39"N,

8°37’38.60”E.).

Coloration in preservation. Upper lip white. Upper side

of the body uniform grey, with darker, white framed, cross-

bands from neck to tail. One band on the neck, three be-

tween the limbs, one on the tail base and seven on the tail.

Tubercle scale pale whitish possessing a spotted pattern.

Underside uniform dirty white, except the tail where the

crossbands are also obvious on the lower side.

Bonn zoological Bulletin 63 (1): 1 14

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12

Philipp Wagner et al.

Fig. 7. Species tree for the Hemidactylus fasciatus species group based on a coalescent-based Bayesian analysis of 1,087 single

nucleotide polymorphisms (Leache et al. 2014). Posterior probabilities are shown on branches. Museum specimen records were

downloaded from the HcrpNET database, and geographic distribution was predicted using Maxent. Populations with uncertain tax-

onomic placement are indicated with “?”. Type localities are: [A]= H. kyaboboensis sp. n.; [B]= H. eniangii sp. n.; [C]= H. biokoen-

sis sp. n.; and [D]= H. coalescens sp. n.

Distribution. Endemic to Bioko Island (former Fernando

Poo, Equatorial Guinea) and only known from some lo-

calities on the island like e.g., Luba, Malabo, San Carlos,

and Ureca.

Habitat. Only known from coastal areas on Bioko Island

where it does not avoid cultural areas (Mertens 1964).

Within the Reserva cientifica de la Caldera de San Car-

los is was found in a forest almost immediately adjacent

to the beach.

Relationships. This species includes all populations that

cluster with those from the Bioko Island with strong sup-

port in the Bayesian species delimitation model (Leache

and Fujita, 2010). It is the sister species to H. coalescens ,

which together form the sister group to H. eniangii sp. n.

(Leache et al. 2014).

Etymology ’. This species is named after Bioko Island,

Equatorial Guinea, where it is endemic.

Hemidactylus ituriensis Schmidt, 1919

1919 Hemidactylus fasciatus ituriensis Schmidt, Contri-

butions to the Herpetology of the Belgian Congo based

on the Collection of the American Congo Expedition,

1909-1915. Part I: turtles, crocodiles, lizards, and

chamaeleons. Bull Amer Mus Nat Hist 39 (2): 385-624

Holoty’pe. AMNH 10272, from “Avakubi,” Democratic

Republic of the Congo.

Description. A fairly large species of Hemidactylus with

a maximum SVL of 89mm and a maximum total length

of 162mm. The species is morphologically characterized

by a high number of subdigital lamellae (9-1 1 on 4"’ fin-

ger; 11-13 on 4th toe); by having the enlarged subcaudal

scales about one-third of the tail width; and by having a

low number of femoral pores in males (8 vs. 16-20 in the

West African species). In coloration the species is distinct

from all other taxa of the H. fasciatus group by lacking

the sharply defined band between neck and eye on the side

of the head and by having the body crossbands, which are

wider than the interspaces, without a median notching (see

Fig. 6).

Distribution. The species is recently only known from

northeastern parts of the DR Congo. Wagner (2010) sug-

gests a wider distribution along the Albertine Rift and

probably eastwards to Uganda and probably Kenya, sim-

ilar to the distribution pattern of forest species in this area

(Wagner et al. 2008, 2009).

Bonn zoological Bulletin 63 (1): 1-14

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Description of New African Forest Geckos

13

Relationships. So far genetic information of this taxon are

lacking, but we expect, due to morphological characters

and the distribution pattern, a close, probably sister tax-

on, relationship to the clade including H. eniangii sp. n.,

H. coalescens sp. n., and H. hiokoensis sp. n.

DISCUSSION

Speciation in the Hemidactylus fasciatus species group (in-

cluding//. biokoensis sp. n., H. coalescens sp. n., H. eni-

angii sp. n., H. fasciatus, and H. ky’aboboensis sp. n.) il-

lustrates some of the common patterns of geographic di-

vergence observed across the reptile and amphibian for-

est fauna of West and Central Africa. The current levels

of West African rainforest fragmentation are due to an ex-

pansion of dry forest and savannah into the rainforest

blocks during the last glacial maximum of the Pleistocene

(Hamilton & Taylor 1991). Within these different frag-

ments, many species diverged in spatial separation (Wag-

ner et al. 2008; Wagner et al. 2009; Wagner 2010). The

Dahomey Gap, a stretch of dry savannah extending from

central Ghana through western Nigeria (Salzmann &

Hoelzmann 2005), separates many West and Central

African forest species (e.g., Amietophrynis superciliaris ,

Lepidothyris fernandi). The formation of this arid barri-

er, which was frequently opened and closed over time

(Salzmann & Hoelzmann 2005), corresponds with the ini-

tial divergence in the H. fasciatus species group (Fig. 7).

The Togo Hills as well as nearby forests, like the Mis-

sahohe, are situated within the Dahomey Gap. These

mountainous areas contain moist semi-deciduous rain-

forests, which are isolated from the more expansive rain-

forest blocks to the west and east (Leache et al. 2006, Wag-

ner 2010). Therefore, these rainforest ‘islands’ harbor

many specialists, including H. ky’aboboensis sp. n.,

which were isolated over time from other populations.

Likewise geographically isolated, H. biokoensis sp. n. is

endemic to Bioko Island in the Gulf of Guinea, separat-

ed from the mainland by about 40 km and sea depths of

less than 60 m. Bioko was probably connected to the main-

land during the last glaciation (Lee et al. 1994). Finally,

the Sanaga River in Cameroon appears the likely barrier,

separating H. eniangii sp. n. from H. coalescens sp. n.

However, fine-scale population sampling is necessary for

investigating the geographic limits of these new species.

The taxonomy within the H. fasciatus species complex

is hampered by the lack of a specific type locality in the

original description of H. fasciatus by Gray (1842); the

collection locality of the name bearing type (BMNH

xxi.24.a) is also not given in the BMNH catalogue. The

first available synonym of the species also has an unspe-

cific type locality ( Leiurus ornatus Gray, 1845 from “W.

Africa”), but the second synonym is more specific ( Hemi -

dactylus formosus Hallowed, 1857 from “Liberia”). Fol-

lowing article 76 of the Code, the type locality is the col-

lecting locality of the name bearing type, but despite rec-

ommendations set forth in 76A. 1 of the Code, it is not pos-

sible to specify the type locality any further. However, fol-

lowing recommendation 76. A. 1.4 of the Code, localities

within the known range of the taxon can be used as the

type locality, but without prejudice to other clarifications.

Given that one of the synonyms of the species has a rel-

atively specific type locality (Liberia), we restrict the dis-

tribution of H. fasciatus to the western Guinean rainfor-

est system from Sierra Leone to the Dahomey Gap in

Ghana, excluding the Togo Hills and other forests with-

in the Dahomey Gap e.g. the nearby Missahohe, which is

not in conflict with the herein described new species. Fol-

lowing Leache and Fujita (2010), H. fasciatus includes

populations from the western and central Guinean rain-

forest (all west of the Dahomey Gap) and whose evolu-

tionary history has been independent from other lineag-

es.

Two other large-bodied, forest-dwelling taxa of Hetni-

dactylus occur in equatorial Central Africa: H. ituriensis

Schmidt, 1919 and H. makolowodei Bauer, LeBreton,

Chirio, Ineieh & Talla Kouete, 2006. However, the latter

is clearly distinct from the H. fasciatus complex (Bauer

et al. 2006), and although the phylogenetic relationships

among these species are currently unknown, we predict

that these large-bodied, forest-dwelling species of Hemi-

dactylus are closely related to the Hemidactylus fasciatus

complex. Schmidt (1919) considered the non-overlapping

morphological characters (e.g., larger head, higher num-

ber of subdigital lamellae [10-10 vs. 12-12], enlarged sub-

caudals about one-third the width of the tail [vs. more than

half of the width], lower number of femoral pores [8 vs.

16-20 on each side]) of H. fasciatus and H. ituriensis suf-

ficient for species differentiation. Loveridge ( 1936) also

regarded H. ituriensis as distinct from H. fasciatus at the

species level, but later recognized the taxon ituriensis as

a subspecies of H. fasciatus (Loveridge 1947). The char-

acters reported by Schmidt (1919) are sufficient for rec-

ognizing H. ituriensis as a full species. Currently, the char-

acters diagnosing the H. fasciatus group refer to color pat-

tern, which was already used by Schmidt (1919) to dis-

tinguish H. ituriensis. An evaluation of other possible mor-

phological characters including body proportions and scale

counts is needed.

Acknowledgements. The authors are thankful to the

Ghanaian Wildlife Division, the Nigerian Biodiversity Re-

search Center, Conservation International, the Nature Con-

servation Research Center and IUCN for assistance with

fieldwork. For the use of their images we are grateful to

Jessica Weinberg (Bioko Biodiversity Protection Program)

and Michael Barej. For useful comments we thank

Michael Barej (ZMB) and two anonymous reviewers.

Bonn zoological Bulletin 63 (1): 1-14

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14

Philipp Wagner et al.

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Gray JE (1842) Description of some new species of Reptiles,

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Hamilton AC, Taylor D ( 1991) History of climate and forests in

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(1999) International Code of Zoological Nomenclature. Inter-

national Trust for Zoological Nomenclature, 4lh edition, 106

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Leache AD, Rodel MO, Linkem CW, Diaz RE, Hillers A, Fuji-

ta MK (2006) Biodiversity in a forest island: reptiles and am-

phibians of the West African Togo Hills. Amphibian & Rep-

tile Conservation 41: 22-45

Leache AD, Fujita MK, Minin V, Bouckaert R. (2014) Species

delimitation using genome-wide SNP data. Systematic Biol-

ogy. doi : 1 0. 1 093/sysbio/syu0 1 8

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ations with distance and time in the volcanic islands of the

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Planetary Science Letters 123:119-138

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Museum ofNatural History Chicago 22: 1-122

Loveridge A (1947) Revision of the African lizards of the fam-

ily Gekkonidae. Bulletin of the Museum of Comparative Zoo-

logy 98: 1-469

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APPENDEX

Material examined

Hemidactylus biokoensis sp. n. Equatorial Guinea. CAS

207624, 207663-4, 207777-8 from Bioko Island; ZFMK

9339-49 from Bioko Island (Ureca); ZFMK 9350-52 from

Bioko Island (San Carlos).

Hemidactylus coalescens sp. n. Cameroon. ZFMK

87679-82 from Nkoelon. Gabon. ZFMK 73188 from

Mokabane; ZFMK 20698 from Kama River. PR Congo.

USNM FS246343 from Impfondo; ZFMK 65482 from be-

tween Oyo and Bokouele.

Hemidactylus eniangii sp. n. Cameroon. AMNH 45332

from Koupe-Manengouba Division; ZFMK 59514 from

MT Kupe; ZFMK 61356 from Mt. Yuhan; ZFMK 61398-

99 from Bonjongo; ZFMK 69603 from Nyasoso; ZFMK

77893 from Ekomtolo. Nigeria. MVZ 253213-15 from

Erokut Park entry gate. Cross River National Park.

Hemidactylus fasciatus BMNH XXI. 24. a, from an un-

known locality. BMNH (unknown) from “W. Africa”

[holotype of Leiurus ornatus Gray, 1845], Ghana.

LSUMNS 86983, 87027, 87037 from Asutnura; MVZ

252492-6 from Birim North District; MVZ 245300-7,

245437 from Ankasa National Park. Liberia. ANSP 745 1 -

53 from Liberia [syntypes of Hemidactylus formosus Hal-

lowell, 1857],

Hemidactylus ituriensis DR Congo. AMNH 10270 from

Batama; AMNH 10271-72 from Avakubi; AMNH 10273

from Akenge; AMNH 10274 from Medje; AMNH 10275

from Panga.

Hemidactylus kyaboboensis sp.n. Ghana. MVZ 245291-

7, 245299, 245436, 249695-704 from Kyabobo National

Park. Togo, ZFMK 19922, from Faille de Baftlo [=fault

proper near Baftlo (9.282399, 1 .2 1 662 1 )]; ZFMK 93689,

from Missahdhe, near Kpalime.

Bonn zoological Bulletin 63 (1): 1-14

©ZFMK

Bonn zoological Bulletin 63 ( 1 ): 15-114

June 2014

Murine rodents (Rodentia: Murinae)

of the Myanmar-Thai-Malaysian peninsula and Singapore:

taxonomy, distribution, ecology, conservation status,

and illustrated identification keys

Uraiporn Pimsai12, Malcolm J. Pearch3, Chutamas Satasook12, Sara Bumrungsri2 & Paul J.J. Bates3

'Princess Maha Chakri Sirindhorn Natural Histoiy Museum, Prince ofSongkla University >, Hat Yai. Songkhla Province, Thai-

land 90112. E-mail: u.pimsai@gmail.com:

department of Biology, Faculty of Science, Prince of Songkla University \ Hat Yai, Songkhla Province, Thailand 90112:

sHarrison Institute, Bower-wood House, 15 St Botolph’s Road, Sevenoaks, Kent, TNI 3 3AQ, UK: E-mail:

pjjbates2@hotmail. com

Abstract. Based on field surveys undertaken between 2010 and 2013, museum studies in Thailand and the UK, and an

extensive literature review, this paper provides information on the 28 species and 12 genera of murine rodents currently

known from peninsular Myanmar, Thailand and Malaysia and Singapore. It incoiporates a detailed summary of past re-

search, 1851-2013, of the Murinae in the study area and includes descriptive characters of the external, cranial and den-

tal morphology and measurements for each of the rodent species. It lists and maps the 93 murine taxa described from the

peninsula, 84 of which are currently considered to be synonyms at species level. Each of the 389 different localities on

the 28 distribution maps is numbered and linked to its source, either literature or museum specimen, and listed in the on-

line gazetteer. The global conservation status of each species is obtained from the IUCN Red List. Remarks are made,

where data are available, on the ecology, karyology, fossil history, sperm morphology, phylogeny, and taxonomic histo-

ry and ambiguities. Recommendations are made for further research. A series of illustrated matrix keys is provided to as-

sist with the identification of all the murine genera and species within the study area.

Key words. Taxonomy, distribution, identification keys, karyology, ecology, conservation status.

INTRODUCTION

Witli 2,277 species, the order Rodentia represents approx-

imately 42 % of all mammal diversity (Musser & Carleton

2005). Geographically widespread and highly adaptable,

rodents occupy a vast array of diverse ecological niches.

They impact on the composition, structure, and succes-

sion of vegetation and fulfil many important ecosystem

services, including assisting with nutrient cycling and the

dispersal of seed and spores. Through their burrowing ac-

tivities, they mix and aerate soils and with their high bio-

mass, they provide an essential prey base for many pred-

ator species (Witrner 2004).

A minority of species causes significant problems to

man. For example, in Asia, it is considered that in any one

particular area between 5 and 1 0 % of rodent taxa are ma-

jor agricultural pests (Aplin et al. 2003). As such, they eat

crops in the field, typically reducing yields of rice by 5

to 10 % (Aplin et al. 2003), 6 % in pineapples (Joomwong

2007) and 5 % in oil palms (Aplin et al. 2003). They eat,

spoil and contaminate stored food and post-harvest loss-

es of 20 % are not unusual. They also carry diseases that

can be transmitted either directly or indirectly to humans

and livestock (Aplin et al. 2003; Chaval et al. 2010;

Received: 18.02.2014

Accepted: 18.04.2014

Witmer 2004). Some of the most virulent diseases borne

by rodents in Southeast Asia include: hantavirus, leishma-

nia infection, leptospirosis, scrub typhus, toxoplasmosis

and viral haemorrhagic fevers (Chaval et al. 2010; Her-

breteau et al. 2012, http://www.ceropath.org/research/ ro-

dent bomediseases).

Recognising the importance of rodents, both as biolog-

ical entities and because of their association with man, the

current paper, which is based on the unpublished work of

Pimsai (2012), seeks to provide a baseline for future re-

search of murines in peninsular Myanmar, Thailand,

Malaysia and Singapore. The study area is illustrated in

dark grey in Fig. 1 and all subsequent maps. It should be

noted that although the northern boundary is clearly an ar-

tifact of geography, corresponding to the northern limit of

peninsular Thailand, it also has zoogeographical signifi-

cance approximating as it does to the Indochinese-Sunda-

ic transition zone, although it is recognised that the exact

location of this zone varies amongst different taxa and has

been interpreted in different ways by different authors

(Woodruff 2003, Woodruff &Turner 2009, Hughes et al.

2011).

Corresponding editor: R. Hutterer

16

Uraipom Pimsai et al.

Fig. 1. Study area: peninsular Myanmar, Thailand and

Malaysia, and Singapore. Localities from which specimen data

were obtained are mapped and listed in the Gazetteer.

The paper focuses on the Murinae as this is the most

speciose subfamily worldwide, with 561 species and 126

genera, witlun the largest Rodent family, the Muridae, 730

species and 1 50 genera (Musser & Carleton 2005). With-

in the study area, the Murinae, the rats and mice, have the

highest population sizes, the greatest biomass, the great-

est impact on man of all the rodents, and the richest di-

versity: 28 species and 12 genera (sensu Musser & Car-

leton 2005) - Bandicota bengalensis (Gray). B. indica

(Bechstein), B. savilei Thomas, Berylmys berdmorei

(Blyth), B. bowersii (Anderson), Chiropodomys gliroides

(Blyth), Hapalomys longicaudatus Blyth, Lenothrix canus

Miller, Leopoldamys ciliatus (Bonhote), L. sabanus

(Jentink), Maxomys inas (Bonhote), M. rajah (Thomas),

M. surifer (Miller), M. whiteheadi (Thomas), Mas mus-

culus Linnaeus, M. caroli Bonhote, Niviventer cameroni

(Chasen), N. cremoriventer (Miller), N.fulvescens (Gray),

Pithecheir parvus Kloss, Rattus andamanensis (Blyth), R.

annandalei (Bonhote), R. argentiventer (Robinson &

Kloss), R. exulans (Peale), R. norvegicus (Berkenhout),

R. tanezumi (Temminck), R. tinmanicus (Miller), and Sun-

da my s muelleri (Jentink).

As a baseline, the paper seeks to put the complex tax-

onomy of the Murinae from the study region into context.

This subfamily includes taxa from two tribes, the Murini

and Rattini, and one ill-defined group, the ‘Murinae in -

certis sedis ’ as defined by Lecompte et al. (2008). All 93

taxa described from the area are listed in Table 1 . Of these,

44 were originally described as new species and 49 as sub-

species. Forty-two were described from peninsular

Malaysia (as understood today), 30 from peninsular Thai-

land, 19 from peninsular Myanmar and 2 from Singapore.

Most were named at the start of the 20"’ Century, 65 be-

tween 1 900 and 1919 and 19 between 1931 and 1941 with

only six, from 1960 onwards. The most recent was de-

scribed from Thailand in 1989. All but nine ( Berylmys

berdmorei, Leopoldamys ciliatus, Maxomys inas, M. su-

rifer, Niviventer cameroni, N. cremoriventer, Pithecheir

panms, Rattus annandalei, and R. tiomanicus — highlight-

ed in bold in Table 1 ) are considered today as synonyms,

at species level. To further aid understanding of the tax-

onomy, the type locality of each taxon is plotted on the

relevant distribution map.

The distribution maps are the first for all murine rodents

from the study region that seek to give a higher level of

geographical resolution by featuring spot localities rather

than the more familiar shaded maps as available in Mar-

shall (1988), Corbet & Hill (1992), and Francis (2008).

Previously, some spot maps were available, as part of larg-

er taxonomic studies, for certain species such as Ha-

palomys longicaudatus (Musser 1972), Chiropodomys

gliroides (Musser 1979), Sundamys muelleri (Musser &

Newcomb 1983) and Bandicota (Musser & Brothers

1994).

Literature sources for the maps were drawn from a large

number of references dating back to Blyth (1851). These

references, together with many others directly relating to

murine rodents in the study area, are reviewed and sum-

marised below. It is hoped that this review will provide a

valuable starting point for all who wish to conduct fur-

ther research on murine rodents in the study area.

The majority, approximately 60 % of the 1 15 publica-

tions included in the review is concerned with aspects of

taxonomy. This is especially the case for papers and books

published in the first one hundred years (1851-1950). Dur-

ing this time, all 40 publications were either exclusively,

or primarily taxonomic and 30 included descriptions of

new murine taxa (Table 1 ). Post 1 950, researchers in the

peninsula began to investigate additional aspects of murine

rodents, including reproduction, behaviour, ecology,

karyology, palaeontology, phylogeny, and the transfer of

diseases between rodents and man.

A particular aim of this paper (inter alia ) is to comple-

ment and support exciting new research looking at the

phylogeny of rodents in the region based on combined mi-

tochondrial and nuclear markers, for example Latinne et

al. (2012, 2013b). The paper seeks to support the idea of

a multi-approach to rodent identification as evinced by

Chaval et al. (2010) by providing a detailed summary of

the alpha taxonomy of the Murinae. It is hoped that the

paper will facilitate future research of rodent taxonomy

in Southeast Asia, and especially amongst in-country stu-

dents and zoologists based in Thailand, Malaysia and

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Murine rodents of the Myanmar- Thai-Malaysian peninsula and Singapore

17

Myanmar, and will help promote studies that combine both

morphometric and molecular approaches.

It is anticipated that the paper will assist those who wish

to investigate further the taxonomy of species, the status

of which has been questioned in a series of papers and on-

line publications, for example Berylmys berdmorei, Sun-

damys muelleri, and Rattus tiomanicus (in Musser & New-

comb 1 983 ), Leopoldamys sabanus (in Musser & Carleton

2005 and Balakirev et al. 2013) and Niviventer fulvescens

(in Musser & Carleton 2005 and Balakirev et al. 2011),

Maxomys surifer (in Gorog et al. 2004, Aplin et al. 2008c

and Latinne et al. 2013b), and Rattus tanezumi (Heaney

& Molur 2008 and Aplin et al. 2011). It is also in response

to papers such as Francis et al. (2010), which although pri-

marily concerned with bats, suggest that the number of

mammal species currently recognised in Southeast Asia

may represent only 50 % of the real diversity. In addition,

it supports Waengsothom et al. (2009) who highlight the

need for further rodent research in Thailand.

This paper seeks to complement the taxonomic work of

close colleagues working on other small mammal taxa in

the region. This team of young taxonomists, based in the

Prince of Songkla University (Thailand), National Univer-

sity of Laos, the Royal University of Phnom Penh (Cam-

bodia), and the Institute of Ecology and Biological Re-

sources (Vietnam), have studied extensively a range of

taxa in mainland Southeast Asia. Together with members

of the Harrison Institute, UK and other collaborating in-

stitutions, they have described several new species of bat

(Douangboubpha et al. 2011, Soisook et al. 2013a and

2013b; Thong et al. 2012) and a new Dying squirrel

(Sanamxay et al. 2013). Additionally, they have undertak-

en many studies of small mammal diversity and distribu-

tion (extant and fossil) in the same area (Furey et al. 2012;

Ith et al. 201 1; Kingsada et al. 201 1; Pearch et al. 2013;

Thomas et al. 2013).

Correct identification of species is of primary impor-

tance to many studies (Chaval et al. 2010). Some species

are considered agricultural pests whilst others, even with-

in the same genus, are harmless or even beneficial, feed-

ing on harmful invertebrates (molluscs and insects) and

invasive weeds. Some species are the reservoir of zoonot-

ic diseases deleterious to man, whilst others, closely sim-

ilar in external morphology are not. To meet the challenges

of murine identification within the study area, we have de-

vised a series of keys that together, help with the identi-

fication of all 28 murine species. Those for the species are

included in a series of Tables in the relevant sections of

the text. Those for the genera are in Tables 25A, B & C.

The diagnostic characters included in the matrix keys have

been extracted from the generic and species descriptions.

These matrices include a range of external, cranial and

dental characters. Much of the information has been drawn

from our own observations but none would have been pos-

sible without the detailed and inspiring work of past tax-

onomists and foremost amongst these are the studies of

Musser (including Musser 1972, 1973a, 1973b, 1979,

1981; Musser et al. 1979; Musser & Brothers 1994; and

Musser & Newcomb 1983).

Similarly, the ecological summaries in the text draw

heavily on the studies of a range of previous authors, no-

tably those of Medway (1969) and a series of papers by

Harrison (dating from 1950 to 1966) and Lim (1966 to

1975). Conservation status is drawn from the on-line

IUCN Red List (http://www.iucnredlist.org/). Of the 28

species, all but six are listed as of ‘Least Concern’. The

exceptions are one ‘Endangered’ species, Hapalomys long-

icaudatus\ four ‘Vulnerable’ species, Maxomys rajah M.

whiteheadi, Niviventer cameroni, and N. cremoriventer;

and one “Data deficient’ species, Pithecheir parvus . How-

ever, each of the 28 species plays an important role in the

ecosystems of the peninsula and each in its own way is

an important subject for future research.

There are great opportunities for a whole range of fu-

ture studies, not just in the peninsula but in Southeast Asia

generally, including taxonomy (incorporating both mor-

phometries and genetic studies), phylogeny, phylogeog-

raphy and palaeontology (looking at evolutionary histo-

ries), ecology and behaviour (especially linked to ecosys-

tem services), as well as those concentrating on the role

of rodents in disease transmission. It is our intention that

the current study will provide a baseline to promote and

facilitate this work in the Myanmar, Thailand and

Malaysian peninsular and Singapore.

LITERATURE REVIEW OF MURINE RODENT

RESEARCH 1851-2013

The first publication relevant to the current study is Blyth

(1851) who described Mus berdmorei (= Berylmys berd-

morei) from the Mergui (= Myeik) Archipelago (no exact

locality). Subsequently, he published on two murines from

Pinang (= Penang), Malaysia (Blyth 1 865); one was a new

taxon, Mus rama (Cantor in Blyth, 1865) (= Mus muscu-

lus) and the other was Mus setifer {setifer is currently re-

ferred to Bandicota indica , but these specimens are more

probably referable to B. bengalensis). Three years later,

Peters (1868) described a new taxon of pencil-tailed tree

mouse, Chiropodomys penicillatus (= C. gliroides); the

type locality is thought to be the Malaysian peninsula, al-

though no details were given (Corbet & Hill 1992).

The first synthesis of mammal records from the region

was provided by Flower (1900) who reported on 160

mammal species collected from Thailand and Malaysia.

He included seven murine taxa from the Thai-Malay

peninsula. Miller (1900a) described Mus tiomanicus (=

Rattus tiomanicus) and Mus obscurus (renamed M. pul-

lus Miller, 1901, = Rattus exulans) from Tioman Island.

In the same year. Miller ( 1900b) published the first nra-

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Uraiporn Pimsai et al.

jor paper dedicated to the peninsular murines. He de-

scribed seven new rat taxa, which had been collected in

Trong (= Trang Province, Thailand). Of these, two are still

recognised as valid species today: Mus cremoriventer (=

Niviventer cremoriventer) and M. surifer (= Maxomys su-

rifer ). The other five are treated as synonyms (sensu Muss-

er & Carleton 2005): Mus vociferous (= Leopoldamys sa-

banus), M. ferreocanus (= Berylmys bowersii ), M. validus

(= Sundamys muelleri), M. asper (= Maxomys white he a-

di), and M. pellax (= Maxomys rajah). All the type spec-

imens are deposited in the United States National Muse-

um.

Miller (1900c) reported on fourteen mammal species

collected on the Batang Islands (today in Satun Province,

Thailand) and Pulo Lankawi (today in Kedah Province,

Malaysia). He described four new taxa, three geograph-

ical races and one species, of murines, Mus vociferous lan-

cavensis (= Leopoldamys sabanus), Mus surifer flavidu-

lus (= Maxomys surifer), Mus surifer butangensis (= Max-

omys surifer) and Mus pannosus (= Rattus tanezumi).

Whilst Miller was publishing in America, in Britain,

Bonhote (1900) described an isolated, endemic species,

Mus ciliata (= Leopoldamys ciliatus), from Perak in

Malaysia. He also recorded a number of additional local-

ities for a range of murine taxa, including Mus mettada

[,«'c] (= Millardia meltada) from Bukit, Jalor, Malaysia.

This latter record is probably mistaken as the species is

currently considered to be restricted to the Indian Subcon-

tinent (Musser & Carleton 2005).

In 1900-1901, an extensive collection of mammals was

made in the islands of the Mergui Archipelago (= Myeik

Archipelago) off the west coast of Tennasserim (=

Tanintharyi Province), Burma (= Myanmar). The speci-

mens were presented to the United States National Mu-

seum. They included many murines, of which Miller

(1903a) subsequently described nine as new species. To-

day, all are considered to be synonyms. Three, Mus

matthaeus, M. stridulus and M. Incas are referable to

Leopoldamys sabanus ; Mus gilbiventer is referable to Ni-

viventer cremoriventer, and Mus luteolus, M. umbridor-

sum, M. bentincanus , M. casensis , and M. domelicus are

referable to Maxomys surifer. In the same paper, on the

basis of nine specimens from Tioman Island, off the east

coast of Malaysia, Miller described one new murid tax-

on, Mus stridens (= Leopoldamys sabanus).

Meanwhile, Bonhote (1903b) named a new species of

rat from Bukit Besar, Jalor, Malaysia, Mus bukit (= Ni-

viventer fulvescens). In a further paper, based on a collec-

tion from northern peninsular Malaysia, Bonhote ( 1 903a)

recorded an additional four rat species and described an-

other three, of which one Mus annandalei (= Rattus an-

nandalei) is still recognised as valid. Both Mus jaloren-

sis (= Rattus tiomanicus) and Mus griseiventer (= Rattus

tanezumi) are today treated as synonyms. A new taxon,

Mus jar ak, named by Bonhote (1905) from Malaysia, is

currently referred to Rattus tiomanicus. In the following

year, Bonhote (1906) provided additional data on ten

murid taxa from southern peninsular Malaysia. This pa-

per included the description of two new taxa, Mus inas

(= Maxomys inas) and Mus klossi (= Maxomys whitehea-

di). Thomas (1907) described two new species from

Pinang (Penang), Malaysia, Gunomys varillus and G. var-

ius, both are currently referred to Bandicota bengalensis.

Kloss (1908a) described two new murine taxa, Mus vil-

losus (= Rattus annandalei) from Singapore and Mus su-

rifer microdon (= Maxomys surifer) from Tioman Island,

Malaysia. His subsequent paper on a provisional list of the

mammals from the peninsular region was an update of

Flower ( 1900) and included 31 murid taxa (Kloss 1908b).

Specimens collected from Tarutao Island (today in Sat-

un Province, Thailand) and remitted to the Natural His-

tory Museum, London, included one new subspecies of

murine, described by Thomas & Wroughton ( 1 909) as Mus

vociferans tersus (= Leopoldamys sabanus). Meanwhile,

based on a collection from the Trengganu Archipelago, off

the north-eastern coast of the Malaysian peninsula, Kloss

(1911a) described two new subspecies, Mus surifer gran-

dis and M. s. flavigrandis, both of which are today treat-

ed as synonyms of Maxomys surifer, more detailed de-

scriptions of these two taxa, including measurements, were

subsequently included in Kloss (1911b). Robinson &

Kloss (1911a) described two new taxa, Mus rattus rumpia

(= Rattus tiomanicus) and Mus surifer leonis (= Maxomys

surifer) from Malaysia and Singapore respectively whilst

Robinson & Kloss (1911b) included an additional new

Malaysian murine taxon, Mus muelleri foederis (= Sun-

damys muelleri). The following year, Robinson (1912) de-

scribed a further two new taxa, Epimys surifer pemangilis

and E. s. aoris, both of which are also treated today as syn-

onyms of Maxomys surifer. The specimens were deposit-

ed in the Selangor Museum.

Miller (1913) published a further review of ‘Malaysian’

mammals. It included fifteen new taxa of murine rodents

from peninsular Myanmar, Thailand and Malaysia. Sev-

en new taxa were published from the Mergui (= Myeik)

Archipelago in Myanmar. All are now considered to be

synonyms with three, Epimys vociferans clarae, E. v. in-

sularum. and E. stentor, currently referred to Leopoldamys

sabanus and four, Rattus rattus fortunatus, R. r. exsul, R.

r. insulanus, and R. r. dentatus referred to R. tanezumi. A

further two taxa, Epimys gracilis (marginally extralimi-

tal to the current study region) and E. lepidus were col-

lected from the Tennasserim mainland (= part of

Tanintharyi Division, Myanmar) both are currently con-

sidered to be synonyms of Niviventer fulvescens. Two taxa

were described from what is today Satun Province, Thai-

land. One, Epimys solus (= N. cremoriventer), is from

Tarutao Island and the other, E. pannellus (= R. tanezu-

mi) from Butang Island. Of the four taxa from peninsu-

lar Malaysia, three, Epimy rattus vicalana, E. roa, and E.

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

19

tingius are now referred to Rattus tiomanicus and one, R.

victor , to Sundamys muelleri.

Robinson & Kloss (1914) reported on a collection of

specimens obtained from Thailand and Malaysia. Three

new taxa of murids were described from Koh Samui, Thai-

land. All are today considered to be synonyms, namely,

Epimys jerdoni pan (= Niviventer fulvescens), E. surifer

spurcus (= Maxomys surifer) and E. remotus (= R. an-

damanensis). The two new taxa from peninsular Malaysia

were Epimys orbus (= N. fulvescens) and E. surifer man-

icalis (= M. surifer). The specimens were donated to the

Federated Malay States Museum.

Kloss ( 1916a) provided additional data on the distribu-

tion of four rodent taxa, which today are referred to Max-

omys surifer Rattus tanezumi, Sundamys muelleri and

Berylmvs bowersii. In two subsequent papers, Kloss

(1916b) included the description of a new subspecies,

Pithecheirus melanurus parvus, which is currently recog-

nised as a distinct species, Pithecheir parvus, and Kloss

(1916c) described Epimys surifer eclipsis (= Maxomys su-

rifer). Robinson & Kloss (1916a) reported on six murine

taxa from Kedah Peak.

Meanwhile, in peninsular Myanmar, Wroughton (1915)

provided distribution and ecological data for nine murine

taxa in southern Tenasserim (= Tanintharyi Division). Hin-

ton (1919), in his extensive paper on the house rats of the

Indian Subcontinent and Myanmar, described one new tax-

on of murid rodent, Rattus rattus tikos (= Rattus tanezu-

mi), also from Tenasserim. Gyldenstolpe (1917) reported

on the mammals collected on two Swedish expeditions to

Thailand, including the peninsula and Gyldenstolpe

(1919) summarised the mammals which had been record-

ed from Thailand, including many rodent taxa from the

peninsula. Kloss (1919) included additional localities of

Rattus rajah surifer (= Maxomys surifer) from peninsu-

lar Myanmar-Thailand and R. r. neglectus (= R. rattus)

from peninsular Thailand. Lindsay (1926) reported on a

collecting trip to the islands of the Mergui (= Myeik) Ar-

chipelago, which included records of five murine species.

Chasen & Kloss (1931) discussed aspects of Rattus tax-

onomy of specimens originating from islands in the Straits

of Malacca. The paper included the description of a new

taxon, Rattus rattus payanus (= R. tiomanicus) from Pu-

lau Paya, off the west coast of Malaysia. Kloss (1931) de-

scribed a new taxon, Rattus canus malaisa (= Lenothrix

canus) from near Kuala Lumpur.

Chasen (1933) published on Malay Rattus. He includ-

ed descriptions, ecological information and distribution da-

ta from Malaysia and Thailand for three taxa which are

today referred to Rattus tiomanicus. R. tanezumi and R.

argentiv enter . Chasen (1936) reviewed some new and ex-

isting specimens of Gunomys (= Bandicota bengalensis)

known from Penang Island, Malaysia. In the same year,

Tate (1936) undertook a taxonomic review of the Muri-

dae, with a study area extending from mainland Southeast

Asia to New Guinea. Chasen (1937) provided information

on a range of murines from the islands off the west coast

of peninsular Thailand and described six subspecies of

Rattus tanezumi, namely: panjius, alangensis, lontaris,

kadanus, moheius, and pipidonis.

In Chasen (1940), an extensive work on ‘Malaysian’

mammals (in reality it included taxa from Malaysia, Thai-

land and Indonesia), 35 pages were devoted to the murid

rodents. He included lists of species, subspecies, and syn-

onyms. In addition, he named a number of new taxa, in-

cluding Rattus rapit cameroni, which is today recognised

as a distinct, endemic species, Niviventer cameroni, from

the Cameron Highlands, peninsular Malaysia. Other new

taxa included: Rattus rattus robinsoni (= R. tanezumi ); R.

r. perhentianus and R. r. pemanggis (both = R. tiomani-

cus); R. sabanus dictatorius and R. s. salanga (both =

Leopoldamys sabanus); and R. surifer puket, R. s. telibon,

R. s. muntia and R. s. pidonis (all = Maxomys surifer). Sub-

sequently a supplement was published by Ellerman &

Morrison-Scott (1955).

Sody (1941) reported on a collection of rats from the

Indo-Malayan-Australian region, with particular empha-

sis on murines from Indonesia. The paper included one

new taxon from the current study region, Rattus rattus

chaseni (= R. argentiventer) from Krian, Perak, Malaysia.

Meanwhile Ellerman ( 1941 ) published his monograph on

the families and genera of living rodents with the murids

included in volume 2. The exhaustive information con-

tained in this treatise included a summary of generic char-

acters and a list of subspecies and synonyms, many of

which originated from peninsular Myanmar, Thailand and

Malaysia.

In the 1950s, there was considerable interest in rodent

behaviour, reproduction, and links to human disease.

Working mostly in peninsular Malaysia, Harrison pub-

lished prolifically (Harrison & Traub 1950; Harrison &

Lim 1950 and Harrison 1952a, 1952b, 1954a, 1954b,

1955). Harrison (1956a) examined seasonality in rodent

breeding rhythms and looked at age-weight ratios for

eleven murine taxa from Malaysia to determine survival

rates. He also recorded Bandicota indica from peninsu-

lar Malaysia and for the first time reviewed the discrim-

inating characters between this species and B. bengalen-

sis (Harrison 1956b). Harrison (1957a) looked at habitats

and micro-habitats of various murine rodents in Malaysia.

Additional papers by Harrison (1957b, 1957c, 1957d,

1958, 1961 and 1962) looked at various aspects of the be-

haviour, ecology and parasites of murine rodents.

Hill (1960) undertook a detailed review of Robinson’s

collections of ‘Malaysian’ mammals, which had been de-

posited in the Natural History Museum, London’s collec-

tion. Despite the title, these specimens, including many

rodents, originated not only from Malaysia but also from

Thailand, Myanmar and Indonesia. Descriptions and

measurements of the different taxa were included, numer-

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20

Uraipom Pimsai et al.

ous localities were listed, and four new taxa were de-

scribed, Rattus rattus pharus, R. r. sribuatensis, R. r. ka -

banicus, and R. r. terutavensis, all of which are now re-

ferred to R. tiomanicus.

Dhaliwal (1961), based on Searle & Dhaliwal (1961),

reviewed the distribution and habitats of five species of

Rattus (today referred to R. annandalei, R. exulans, R.

norvegicus, R. tanezumi, and R. tiomanicus) in Singapore

Island. Subsequently, he discriminated between two Rat-

tus taxa, diardii (= R. tanezumi ) and jalorensis (= R.

tiomanicus) (Dhaliwal 1962 and 1963).

Medway (1964a) undertook a review of Rattus inas (=

Maxomys inas) with information on its ecology, distribu-

tion and external and cranial characters. He compared

specimens from peninsular Malaysia with those from

Sabah. In the same year, he published on the marmoset

rat, Hapalomys longicaudatus, from Malaysia (Medway

1964b). Rudd ( 1965) compared the weights of 10 murid

taxa in the State of Selangor, examining individual and

sexual variation. Harrison (1966) summarised nearly 20

years of research undertaken by the Institute of Medical

Research, Kuala Lumpur in his rather informal book on

the mammals of Malaysia and Singapore. Medway & Lim

(1966) reviewed the taxonomy of three Rattus taxa:

tiomanicus, jalorensis and diardii.

Lim et al. ( 1965) looked for the presence of parasitic

nematodes in 14 species of murid rodent in the Kuala

Lumpur area. Lim (1966) studied the predation of mol-

luscs by wild rats since it was considered probable that

land slugs and snails were intermediate hosts in the life-

cycle of a nematode, which is the cause of eosinophilic

meningoencephalitis in man. Medway ( 1 967) provided de-

tailed information on the breeding habits of Chiropodomys

gliroides based on specimens collected from Selangor

State, Malaysia.

Yong ( 1968) provided karyological data on four species

of murines collected from peninsular Malaysia, namely:

Rattus bowersii ferreocanus (= Beiylmys bowersii ), R.

muelleri validus (= Sundamys muelleri ), R. edwardsi cil-

iatus (= Leopoldamys ciliatus) and R. sabanus vociferans

(= L. sabanus). Yong ( 1969a) and Yong (1969b) includ-

ed the karyology of R. berdmorei (= B. berdmorei) from

Thailand but without exact locality data. Yong (1969c) re-

ported on rats from Kedah Peak, Malaysia and Harrison

(1969) reported on the abundance and population densi-

ty of mammals, including murine rodents, in Malayan low-

land forests.

Meanwhile, Medway (1969) published his monograph

on ‘The wild mammals of Malaya’, providing informa-

tion on the distribution, identification, subspecies, habits

and reproduction of all mammals from Malaysia and Sin-

gapore, including the murine rodents.

Yong (1970) included data on the external morpholo-

gy, skull anatomy, habitat, breeding behaviour, karyolo-

gy, serology and immunohaematology of Rattus edwarsi

(specimens subsequently referred to Leopoldamys cilia-

tus) and R. sabanus (L. sabanus) from peninsular

Malaysia. Yong & Dhaliwal ( 1 970) discussed variation in

pelage colour in Rattus bowersii (= Beiylmys bowersii)

from Malaysia. Marshall & Nongngork (1970) studied

seven rat species on Koh Samui, Thailand with informa-

tion on the discriminating characters and their ecology.

Lim (1970) provided information on the ecology, diet,

food preferences, and endoparasites of four species of

murid rodent, Rattus sabanus (= Leopoldamys sabanus),

R. muelleri (= Sundamys muelleri), R. bowersii (=

Beiylmys bowersii) and R. edwardsi (here considered to

be Leopoldamys ciliatus) based on specimens collected

throughout much of peninsular Malaysia.

Muul & Lim ( 1971 ) provided some valuable informa-

tion on the distribution and ecology of four lesser known

murid rodents, Rattus edwarsi (here considered to be

Leopoldamys ciliatus ), R. annandalei, Lenothrix canus,

and Pithecheir parvus in Malaysia. Lim et al. ( 1 97 1 ) pub-

lished a review of the small mammals of Penang Island,

Malaysia which included information on ten murid taxa.

Yong (1971) described a new taxon, Rattus tiomanicus

tenggolensis, from Pulau Tenggol, Malaysia.

Musser (1972) provided a detailed review of the genus

Hapalomys including information on taxonomy, ecology,

and distribution in Myanmar, Thailand and Malaysia.

Yong et al. (1972) looked at the karology of diardii (= R.

tanezumi) and tiomanicus and jalorensis (= R. tiomanicus).

Musser ( 1 973a) undertook a detailed review of Rattus cre-

moriventer (= Niviventer cremoriventer) and included in-

formation on taxonomy, distribution, and ecology.

Markvong et al. (1973) studied the karyology of murids

from throughout Thailand and included a rare record of

Rattus berdmorei (= Beiylmys berdmorei) from peninsu-

lar Thailand.

Yong (1973) undertook a study of the chromosomes of

Chiropodomys gliroides from Malaysia. Lim & Muul

(1975) studied the rare arboreal murine, Pithecheir

parvus, in peninsular Malaysia and included information

on the diagnostic characters, size, preferred habitats, dis-

tribution, habits of captive specimens, diet, and helminth

infection. Langham & Ming (1976) included the first

record of Mus caroli from peninsular Malaysia with in-

formation on the diagnostic characters and ecology. Med-

way & Yong ( 1976) studied aspects of rat systematics in

peninsular Malaysia.

Marshall (1977b) provided a summary of the murid ro-

dents of Thailand for Lekagul & McNeely ( 1977). He in-

cluded illustrated keys, some synonyms, information on

reproductive status, photographs of live animals and

skulls, some measurements, short descriptions, distribu-

tion maps, and karyology. In the same year, Marshall

( 1977a) provided a detailed review of the genus Mus in

Asia, including the Myanmar-Thai-Malaysian peninsula,

with illustrated keys, information on karyology, distribu-

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

21

tion, diagnostic characters and ecology. Chan ( 1977) and

Chan et al. (1978 and 1979) looked at protein variation

and systematics in Malaysian rats. Tweedie (1978) pro-

vided a brief introduction to the rodents of Malaysia with

a few measurements and some information on habits.

Musser et al. (1979) undertook a detailed revision of the

genus Maxomys, with much data relevant to peninsular

Myanmar, Thailand and Malaysia. In the same year, Muss-

er ( 1979) also undertook a detailed study of the arboreal

rat genus Chiropodomys, with information on the taxon-

omy, ecology and distribution. Fain et al. (1980) studied

the parasitic mite fauna of Malaysian rodents. Yong et al.

(1982) included information on the karyology of Ha-

palomys and Pithecheir. Musser (1981) published a de-

tailed taxonomic paper which focused on two genera Ni-

viventer and Leopoldamys with further comments on

Lenothrix and Maxomys.

Abe ( 1983) reviewed the taxonomy of Niviventer from

Thailand and included bukit in the synonymy of N. fiul-

vescens. In the same year, Musser & Newcomb (1983)

published a monograph on two genera, Berylmys and Sun-

damys, and also provided detailed information on 12 oth-

er genera known from the region, including 5 that are en-

demic to the Sunda Shelf. Langham (1983) studied the

small mammals of peninsular Malaysia, including Kedah

Peak.

Breed & Yong (1986) looked at the sperm morphology

of 19 taxa of murid rodents from Malaysia. Marshall

(1988) in the second edition of Lekagul & McNeely

(1988) once again provided taxonomic, ecological, and

distribution data on 36 species and 8 genera of murine ro-

dents. Boonsong & Felten (1989) described a new

species of bandicoot rat, Bandicota bangchakensis (= B.

savilei) from Nakhon Si Thammarat, Thailand. (It should

be noted that the publications of the Thai naturalist Boon-

song Lekagul are sometimes referred to Boonsong and

sometimes to Lekagul).

Corbet & Hill (1992) published a taxonomic monograph

on the mammal fauna of Southeast Asia. They provided

dichotomous and character matrix keys, a comprehensive

list of synonyms and subspecies, and distribution maps.

The book includes 12 genera and 23 species of Murinae

from peninsular Myanmar, Thailand and Malaysia and

Singapore. Musser & Brothers (1994) undertook a detailed

review of the genus Bandicota in mainland SE Asia, in-

cluding taxonomy, ecology and distribution. Three species

were recognised as occurring in the study area, B. ben-

galensis (with a very restricted range), B. savilei and B.

indica.

Chaimanee (1998) produced an invaluable, detailed

monograph of the Plio-Pleistocene rodents of Thailand,

which also included much information relevant to the ex-

tant species, especially their dentition. Chaimanee &

Jaeger (2001 ) subsequently reviewed the evolution of Rat-

tus and incorporated fossil data originating from penin-

sular Thailand. Gorog et al. (2004) looked at the phylo-

geography of three murine species, Maxomys surifer,

Leopoldamys sabanus, and M. whiteheadi in the Sunda

shelf. Francis (2008) published a guide to the mammals

of South-east Asia. Maps and illustrations were provided

for 26 of the murine species from the study region and

were further supported with brief information on diagnos-

tic characters, measurements, taxonomic notes, distribu-

tion, and ecology (two additional murine species were dis-

cussed but not illustrated).

Waengsothorn et al. (2009) included data about the ro-

dent specimens held in the ‘Centre for Thai National Ref-

erence Collections (CTNRC)’. They identified and

mapped geographical areas of Thailand that have been

well studied in the past and suggested priority areas for

field research in the future. Most recently, there have been

a series of papers on rodent-borne diseases in Thailand

(Jittapalopong et al. 2008, 201 1; Herbreteau et al. 2012).

Meanwhile Pages et al. (2010) looked at Rattini phyloge-

ny, using DNA sequence information, to determine

species boundaries. The results of this latter paper are de-

signed to assist with medical health projects relating to

murine rodents. Most recently, Pimsai (2012) undertook

a study of the murine rodent fauna of peninsular Thailand

and Malaysia, which is the basis for current paper. Achma-

di et al. (2013) looked at the phylogeny, diversity and bio-

geography of Southeast Asian Maxomys, including spec-

imens from peninsular Malaysia and Latinne et al. (2013b)

looked at the diversity and phylogeny of Murinae associ-

ated with limestone karst throughout Thailand, including

the peninsula. Pearch et al. (2013) undertook a review of

small mammal fossil faunas from Thailand including ro-

dents from peninsular Thailand.

METHODS

Species. The taxonomy of the 28 species and 12 genera

follows Musser & Carleton (2005). Differences to this tax-

onomic opinion are noted in the text and particularly in

the ‘Taxonomic notes’ for each species.

Material. Much of the data are based on the literature.

However, additional specimens were collected in Thailand

using cage traps and aluminium live traps, which were set

both on the ground and in trees. The methodology fol-

lowed Aplin et al. (2003). This new material was deposit-

ed as voucher specimens (dry skins and skulls) in the zo-

ological collection of the Princess Maha Chakri Sirind-

hom Natural History Museum, Prince of Songkla Univer-

sity, Thailand (PSUNHM).

An additional 165 specimens of 18 species were stud-

ied from 65 localities in Thailand and Malaysia (Pimsai

2012, Figure 18). The vast majority (61) of these locali-

ties were in Thailand, including 1 8 in peninsular Thailand,

with only four localities in peninsular Malaysia. These

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Uraipom Pimsai et al.

base of

vibrissae/

whiskers

cheek

throat

tail, bicoloured

ventral surfaci

under side/

muzzle supraorbital

vibrissae crown of head

pinna (ear)

nape of neck

dorsal surface/ upper parts

/ midline of back

' -j

line of /

demarcation

base/ proximal

end of tail '

' tip/ terminal/

distal end of tail

under parts hind hind li

A

Fig. 2. (A): External characters of a murine rodent (Maxomys surifer ), as referred to in the text; (B): arrangement of mammae

in an adult female murine rodent (after Aplin et al. 2003, Figure 4.4); (C): the structure of the underside of a left hind foot ( Leopoldamys

sab anus).

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

23

specimens were held in the Centre for Thai National Ref-

erence Collections (CTNRC), Thailand Institute of Sci-

entific and Technological Research (TISTR); the Harri-

son Institute, UK (HZM); and the Natural History Muse-

um, London (BMNH).

Descriptions. Generic and species descriptions are based

on personal observation (where specimens were available

to the authors) and on the literature. Literature sources in-

cluded original descriptions of the different taxa and sub-

sequent taxonomic studies and monographs. The princi-

pal literature sources are listed at the beginning of each

description.

Generic descriptions are restricted to the characters of

species found within the study area. They do not include

any additional characters of species which are found ex-

clusively extralimitally. If only one species in a genus is

found in the study area, the generic description is omit-

ted.

Common terms used in the description of the external

characters are illustrated in Fig. 2A. Where data are avail-

able, the arrangement of the mammae of each species is

given, the terminology of which is given in Fig. 2B and

follows Aplin et al. (2003). The structure of the hind feet

for each genus is illustrated where specimens were avail-

able to the authors with sufficient resolution of structure

to permit an accurate drawing. The nomenclature of the

pads is illustrated in Fig. 2C. The different hair types, in-

cluding ‘spines’ and guard hairs are illustrated in Fig.

3A-C. The section of tail of Bandicota indica shows the

arrangement of the rows of scales, typical of many murine

species (Fig. 3D). A photograph of Maxomys surifer is in-

cluded to show a clearly defined demarcation on the flanks

between the dorsal and ventral pelage (Fig. 3E); the pho-

tograph of the Bandicota indica specimen illustrates a

pelage type that has no clear definition between the up-

per and lower parts (Fig. 3F).

Descriptions of the cranial and dental characters for each

genus are supported in the text by a series of 1 2 drawings.

Technical terms of features used in the generic/specific de-

scriptions are lettered and numbered and linked to the

drawings. This method has been adopted to assist all those

using the paper who are not familiar with cranial and den-

tal morphology. To assist further, the nomenclature of the

cranial structures is also illustrated in Fig. 4A and the den-

tal structures in Fig. 4B.

In general, the skull and teeth of young adult specimens

(where available) were used for illustrative purposes, as

it was considered that these were the most useful in help-

ing to identify specimens caught in the field. However, it

is appreciated that the morphology of the cusp patterns

changes through time with young individuals often hav-

ing much more clearly defined cusps than older individ-

uals. Examples of these changes are illustrated in Fig. 5

for four different species (Berylmvs bowersi, Maxomys su-

rifer, Rattus tanezumi, and Sundamys muelleri). In addi-

tion, the dentition of a young Lenothrix canus is includ-

ed in Fig. 4B, which can be related to the older individ-

ual in Fig. 1 8B. It is important that age-related differences

in tooth morphology are borne in mind when using den-

tal characters for identification purposes.

All drawings of the skulls and dentition were made by

UP with a camera lucida and a stereo-microscope. One

specimen of one species for each genus is illustrated.

Measurements. In the original study, Pimsai (2012) took

a series of measurements from all specimens available to

her in three Thai collections (see Material above). How-

ever, this original study was primarily focused on differ-

ences between genera and was less concerned with spe-

cific differences. For the current study, it was considered

that the number of specimens measured was too small and

their geographical origin too restricted for these measure-

ments to be considered representative for many of the

species. In addition, for some taxa, such as Niviventer

cameroni, Maxomys inas, and Pithecheir parvus, no meas-

urements were available in the initial study. Therefore,

measurements included here are from a wide variety of

literature sources, including many type descriptions. Usu-

ally more than one set of measurements is included for

each species. These have been selected to represent dif-

ferent populations from different geographical areas and/or

because they include additional measurements that are not

included in the primary sources. All sources are listed in

the caption for each Table.

It is appreciated that different researchers use different

measuring techniques. In consequence, it is impossible to

define measurements so that they exactly correspond to

the method used by all the authors. However, the defini-

tions given below follow what we consider to be best prac-

tice. For the cranial measurements (Fig. 6), we have pri-

marily followed Musser (1979) and Harrison & Bates

(1991). More measurements are included than those re-

ferred to directly in the current text. It is believed that the

additional definitions will assist those working with sup-

porting literature that does not include clear definitions of

measurements. For the external characters, we have fol-

lowed Aplin et al. (2003).

Head and body length (HB): from the tip of nose to the

distal end of the anus;

Tail length (T): from the middle of the anus to the tip

of the tail (not including hairs);

Ear length (E): taken inside the ear from the bottom of

the notch to the furthest point along the rim/margin of the

pinna;

Hind foot length (HF ): from the back of the heel to the

tip of the longest toe (not including the claws);

Mass: whole body mass of the specimen (in grams);

Greatest skull length (GSL): the greatest antero-poste-

rior length of the skull, taken from the most projecting

point at each extremity, irrespective of what structures

form these points;

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Uraipom Pimsai et al.

Fig. 3. Hair and tail of Murine rodents. (A): short (underfur) and long (overfur) hairs of Rattus tanezumi; (B): spines of Max-

omys surifer ; (C): guard hairs of Bandicota inclica ; (D): rows of scales on the tail of Bandicota indiccr, E: M. surifer - clear line

of demarcation on the flank between upper and lower pelage; F: Bandicota indica - no line of demarcation on the flank between

upper and lower pelage. Not to scale.

Occipital nasal length (ONL): the distance from the tip

of the nasals to the posterior margin of the occiput (often

but not always the same as GLS, especially in rodents with

procumbent incisors);

Condylobasal length (CBL): from an exoccipital

condyle to the anterior alveolar margin of the most for-

wardly projecting upper incisor tooth;

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

25

Fig. 4. Nomenclature of (A): cranial

structures (based on Bandicota indica)

and (B): dental structures — left upper

molars, M'-M\ (on left) and left lower

molars, M^Mj, on right of a young

Lenothrix canus (following Musser &

Newcomb 1983, Figure 2).

Skull: AP: angular process; BO: basioc-

ciput; BS: basisphenoid; CDP: condylar

process; CRP: coronoid process; EC:

exoccipital condyle; F: frontal; FM:

foramen magnum; HP: hamular process;

IF: incisive foramina; IL: incisor, low-

er; IP: interparietal; IRP: incisor root

process; IU: incisor, upper; L: lachry-

mal; MA: mastoid; MAX: maxilla;

IVIDC: mandibular (lower) molars; MS:

mesopterygoid space; MXC: maxillary

(upper) molars; N: nasals; NC: naso-

lachrymal capsules; O: orbit; P: palate;

PAR: parietal; PM: premaxilla; PPF:

posterior palatine foramina; R: ramus of

half mandible; RO: rostrum (premaxil-

lae and nasals); SO: supraoccipital;

SOU: supraoccipital ridge; SPMC: su-

perior posterior mastoid chamber; SR:

supraorbital ridge; SRZ: squamosal

roots of zygoma; TB: tympanic bulla; Z:

zygomatic arch; ZP: zygomatic plate.

Dentition: Cusps on upper molars are

numbered according to Miller’s (1912)

scheme and referred to in the text with

the prefix ‘f ; a-cen: anterocentral cusp;

a-lab: anterolabial cusp; ale: anterior

labial cusplet; a-ling: anterolingual

cusp; ed: entoconid; hd: hypoconid; md:

metaconid; pc: posterior cingulum; pd:

protoconid; pic: posterior labial cusplet.

Zygomatic breadth (ZB): the greatest breadth across the

zygomatic arches, regardless of where this point is situ-

ated along the length of the arches;

Interorbital breadth (IB): the least distance, as viewed

dorsal ly, across the frontal bones between the orbital fos-

sae;

Length of rostrum (LR): the distance from the tip of the

nasal bones to the posterior margin of the zygomatic notch

(the anterior edge of the dorsal maxillary root of the zy-

gomatic plate);

Breadth of rostrum (BR): the greatest breadth across the

rostrum;

Breadth of braincase (BB): the breadth of the braincase

measured just above the squamosal roots of each zygo-

matic arch;

Height of braincase (HBC): the distance from the top

of the braincase to the ventral surface of the basisphenoid

bone;

Breadth of zygomatic plate (BZP): the least distance be-

tween the anterior and posterior edges of the zygomatic

plate;

Bonn zoological Bulletin 63 ( 1 ): 15-114

Length of nasals (LN): the distance from the anterior

tip of the nasal bones to the most posterior suture between

the nasal and frontal bones, measured parallel to the sur-

face of the nasals;

Length of diastema (LD): the distance from the poste-

rior alveolar margins of the upper incisors to the anterior

alveolar margins of the first upper molars;

Length of incisive foramina (LIF): the distance from the

anterior edge of one of the foramina to its posterior edge;

Palatal length (PL): the distance from the anterior alve-

olar margins of the incisors to the posterior edge of the

palatal bridge;

Post palatal length (PPL): the distance from the poste-

rior margins of the palatal bridge to the anterior margin

of the foramen magnum;

Breadth of bony palate at first molars (BBPM1): the least

distance between the lingual edge of the alveolus of the

first molar and the lingual edge of the alveolus of the op-

posite molar;

Breadth of bony palate at third molars (BBPM1): the

least distance between the lingual edge of the alveolus of

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Uraipom Pimsai et al.

y«S

Fig. 5. Changes in dental mor-

phology with age. A-H: left upper

molars (M'-M1); I-P: left lower

molars (Mj— M3). In each pair,

subadult teeth are on the left and

adult teeth on the right. (A/I):

Be?ylmvs bowersi, subadult,

(HZM 1.40111), Thailand; (B/J):

B. bowersi, adult, (54-1287 CT-

NRC), Thailand; (C/K): Maxomy

surifer, subadult, (54-5587 CT-

NRC), Thailand; (D/L): M. surifer,

adult, (54-2919 CTNRC), Thai-

land; (E/M): Rattus tanezumi,

subadult (54-5032 CTNRC), Thai-

land; (F/N); R. tanezumi, adult

(54-4016), Thailand; (G/O): Sun-

da my s muelleri, subadult, (54—642

CTNRC), Thailand; (H/P): 5.

muelleri adult (54 4392 CTNRC),

Thailand. Scale = 2 mm.

the third molar and the lingual edge of the alveolus of the

opposite molar;

Length of bullae (LB): the length of the bullae, antero-

posteriorly;

Length of maxillary toothrow (ALM'-M3): the distance

from the anterior edge of the alveolus of the first upper

molar (M1) to the posterior edge of the alveolus of the third

upper molar (M3);

Length of mandible (ML): the greatest distance from the

most posterior part of the condyle to the most anterior part

of the mandible, including the lower incisors/ not includ-

ing the incisors (dependent on author);

Length of mandible toothrow (ALM,-M3): the distance

from the anterior edge of the alveolus of the first lower

molar (M, ) to the posterior edge of the alveolus of the third

lower molar (M3).

Distribution maps are based on literature records and on

museum specimens studied personally by the authors or

accessed through MaNIS (http://manisnet.org /portals,

html). Each location on each map is included in the on-

line gazetteer and is also linked to its source (museum

specimen or literature), which is listed in Appendix 1. Ab-

breviations of collections in the appendix are as follows:

AMNH: American Museum of Natural History, New York,

USA; BMNH: The Natural History Museum London (for-

merly the British Museum of Natural History), UK;

FMNH: Field Museum of Natural History, Chicago, USA;

NHM Thailand: The Natural History Museum, Thailand

(specimens formerly in the Thailand Institute of Scientif-

ic and Technological Research [TISTR]); PSUNHM:

Prince of Songkla University Natural History Museum,

Hat Yai, Thailand; TTU: Texas Tech University, Lubbock,

USA; UMMZ: University of Michcgan, Museum of Zo-

ology, USA; USNM: United States National Museum

(Smithsonian), Washington. The maps and appendix were

compiled by MP. The gazetteer was prepared by UP and

MP.

Data for karyology are taken from the literature. Infor-

mation on the fossil record is from Chaimanee (1998),

Musser & Carleton (2005) and Pearch et al. (2013).

Conservation status is accessed from the web-based

IUCN Red List and is individually referenced for each of

the 28 species. Information on the ecology of the differ-

ent species is based on personal observation in peninsu-

lar Thailand and on a wide variety of published references.

SYSTEMATIC SECTION

The descriptions of the 1 2 genera are provided to assist

with the identification of murine rodents within the Myan-

mar-Thai-Malaysian peninsula and Singapore. Therefore

(as noted in the Methods section above), they are restrict-

ed to the characters of species found within the study area.

These descriptions omit characters of species that are en-

tirely extralimital to this region. Where only one species

in a genus is found in the study area, the generic descrip-

tion is omitted and the description is restricted to the

species. Information provided for the genera and 28

species, including morphological characters, distribution

and ecology, is based on personal observation (where da-

ta were available) and literature sources, which are listed

in the relevant sections.

The generic and species matrix keys are based on the

descriptions included below. They have been designed to

assist those working in the field with live or newly sacri-

ficed specimens as well as those conducting research in

museums and/or laboratories with prepared voucher ma-

terial.

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

27

Fig. 6. Cranium and teeth of Leopoldamys sabanus (54-1319 CTNCR) showing limits of cranial measurement, which are de-

fined in the text.

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Uraiporn Pimsai et al.

Genus Bandicota

Bandicoot rats

Bandicota Gray 1873: 418; type species B. gigantea Hardwicke,

in B. indica (Bechstein).

Remarks. According to Musser & Carleton (2005), there

are three species of extant Bandicota (B. indica, B. ben-

galensis and B. savilei) currently recognised, all of which

occur in the study area of peninsular Myanmar, Thailand

and Malaysia. A detailed study of the genus and its species

is provided by Musser & Brothers ( 1994). Synonyms are

listed in Musser & Carleton (2005).

Description (based on personal observation and Musser

& Brothers 1994). The body size ranges from medium to

large (Table 2). The tail is nearly always shorter than the

head and body, almost hairless without a terminal tuft, uni-

coloured, brown to brownish-black (Fig. 3D). The dorsal

pelage is grey-brown to almost black (Fig. 5 IF). The fur

is long and coarse, without spines but with long guard

hairs, which are most conspicuous on the lower black (Fig.

3C). The ventral pelage is slightly paler than the dorsal.

However, there is no clear line of demarcation in colour

between the hairs present on the upper and lower body.

The hind feet have six or seven relatively small plantar

pads (Fig. 7C) including interdigital pads ( 1 ), a very small

outer metatarsal pad (2), and a kidney-shaped inner

metatarsal pad (3). Females of B. indica and B. savilei

have six pairs of mammae; females of B. bengalensis have

up to 20 mammae.

In the skulk the rostrum is short and broad; the tips (Fig.

7Ai) of the nasals (ii) are triangular and about in line with

the anterior surface of the incisors (xv) in B. indica and

B. savilei, although they are posterior to the procumbent

incisors (xxiv) of B. bengalensis. The interorbital area is

narrowed (iii). The braincase is medium sized with promi-

nent ridges (iv), which begin at the the supraoccipital (vi)

and terminate in the interorbital region. The zygomata (v)

are almost parallel or only slightly convergent

anteriorly. The palate (xiii) is long, its length varies be-

tween species, and its posterior border (x) extends beyond

the last (M3) upper molars (ix). Therefore the postpalatal

length, the distance from the back edge of the palate to

the ventral margin of the foramen magnum (xii) is con-

siderably shorter than the palatal length. The incisive

foramina (vii) are elongated and often narrowed posteri-

orly; their posterior margins (viii) extend towards or pass

slightly behind the front of the upper molar toothrows (B.

indica and B. savilei) (xiv); in B. bengalensis, they extend

further posteriorly between the first molars (M1). The tym-

panic bullae are relatively large (xi). In each mandible, the

coronoid process (xix) and the emargination (xxi) between

the condylar (xx) and angular processes (xxii) are well-

developed; the lower incisor root forms a very prominent

process on the mandible (xxiii).

The upper incisors are opisthodont in B. indica and B.

savilei and procumbent in B. bengalensis (Fig. 7) their

enamel layers are orange, relatively pale in B. bengalen-

sis. In the upper and lower molars, the initial cusp pattern

found in juveniles is quickly lost through wear and sub-

sequently appears as a series of simple laminae. The first

upper molar (M1) is relatively large; the first lamina com-

prises cusps 1 1 , t2, and t3 (a), which are fused to form a

plate; the second lamina also has t4, t5 and t6 (c) fused,

although t4 (b) may look semi-separate in younger indi-

viduals; in the third lamina, cusps t8, t9 are fused mak-

ing an oval wear surface (d). In the second upper molar

(M:), cusp t3 is absent; sometimes 1 1 (e) is fused with t4,

t5, t6 (f) to form a plate (as illustrated in Fig. 7B), alter-

natively cusp tl is prominent and not fused; on the third

lamina (which may appear to be the second lamina, de-

pending on the status of 1 1 ), cusps t8 and t9 (g) are fused

making an oval wear surface. In the third upper molar

(M3), cusps tl , t4, t5, t6 (h) are fused making a large part

of the molar; the second lamina, comprises t8 (i) only.

In the mandibular toothrow, the first lower molar ( M , )

includes an anterocentral (j), anterolingual (1) and antero-

labial cusp (k), which are fused together making a large

lamina; the protoconid cusp (m) is fused with the meta-

conid (n); the hypoconid (o) is fused with the entoconid

(p); the posterior cingulum is small (q) (absent in B. ben-

galensis). In the second lower molar (M2), the anterolabi-

al cusp (not seen in the specimen illustrated in Fig. 7B),

the protoconid (r) and metaconid (s) and are fused; the

hypoconid (t) is fused with the entoconid (u); the poste-

rior cingulum (v) is small (absent in B. bengalensis). The

third lower molar (M3) is not greatly reduced when com-

pared with M2; the anterolabial cusp (w) is present and

fused with the protoconid (x) and metaconid (y); the pos-

terior lamina comprises the entoconid cusp (z).

Key to species for the three species currently known from

peninsula Myanmar, Thailand, and Malaysia is included

in Table 3.

Bandicota bengalensis

Lesser bandicoot rat

Arvicola bengalensis Gray 1835 (in 1830-35): pi. 21; Bengal,

India.

Gunomys varius Thomas 1907: 204; Georgetown, Penang Island,

Malaysia.

Gunomys varillus Thomas 1907: 205: Georgetown, Penang Is-

land, Malaysia.

Description (based on personal observation and Chasen

1936 and Musser & Brothers 1994). Extralimitally this is

the smallest of the three species of Bandicota, although

specimens from Penang appear to be quite large (Table 2).

The tail, which is shorter than head and body length, is

virtually naked (although contrary to this, according to

Thomas 1907, in specimens from Penang, the tail is ‘well-

clothed’ in coarse hairs 2-3 mm in length). It is uni-

coloured, brownish to grey brown, with small scales, about

10-12 rows of scales/cm. The fur on the upperparts of the

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j

i

n

P

Fig. 7. Skull and dentition of Bandicota indi-

ca (54—2806 CTNRC), Ko Khram Yai, Chon Buri

Province, Southeast Thailand. (A): dorsal, ven-

tral and lateral view of skull and mandible (Scale

= 5 mm); (B): occlusal view of upper (left) and

lower (right) left molars (Scale = 2 mm); (C): left

hind foot of B. indica (PSUZC-MM2012.199),

Phato Watershed, Chumphon Province, peninsu-

lar Thailand (Scale = 5 mm). D: lateral view of

anterior part of skull of Bandicota bengalensis

(HZM 1.39650) Mandalay Division, Myanmar.

Indicative numbers (i-xxiv) and letters (a-z) are

explained in the text ‘Description’ for the genus

Bandicota. M: mouth; lab: labial.

body is harsh in texture, and dark, an admixture of browns,

greys, lighter buff, and black (based on extralimital spec-

imens); it is described as mixed black and cream-buff,

without brown tints; slaty grey at the base in the descrip-

tion of i ’arias from Penang, Malaysia. There are moder-

ately long, dark brown/black guard hairs. The head and

dorsal line are particularly heavily pencilled in black. The

underparts are dark grey (extralimital specimens) or al-

ternatively in the description of varius from Penang, they

are dull grizzled greyish, with slaty grey bases and dull

whitish tips. There is no clear demarcation on the flanks

between the upper and lower pelage. The hindfeet are

shorter (in comparison to B. indica ); brown to brownish

grey in colour. There are 1 0 to 20 mammae (normally be-

tween 14 and 17): one pectoral, one postaxillary, one ab-

dominal, and two inguinal pairs; higher counts are due to

one additional postaxillary pair and variation in the num-

ber of abdominal mammae.

In extralimital material, the skull is smaller than those

of B. savilei and B. indica (skulls illustrated in Musser &

Brothers 1994, Figures 5 and 6). This is especially appar-

ent in the shorter occipitonasal length (ONL), length of

nasals and shorter length of the maxillary molars

(ALM'-M3); however, in material from Penang skull size

can be quite large (Table 2). Unlike in B. savilei , when

viewed from above, the nasals do not conceal the nasal

opening or the upper incisors. The upper incisors, which

are pale orange to creamy in colour, are procumbent (Fig.

7Dxxiv) whereas they are opisthodont in B. savilei and B.

indica (Fig. 7Axv).The incisive foramina of 5. bengalen-

sis are relatively long and extend posteriorly between the

first upper molars (M * ). In the posterior part of the brain-

case, the posterolateral surface of the supraoccipital (Fig.

7Axvi) is greatly reduced with the supraocciptal ridge

(xviii) fused with the dorsal third of the lambdoidal ridge

(xvii). In B. savilei and B. indica, these two ridges are

clearly separate (see Musser & Brothers 1994, Figure 8

for further details). In extralimital material, the crown

width of the first molar (M1) of B. bengalensis (2.41 mm,

2.13-2.75 mm) averages narrower than that of B. savilei

(2.81 mm, 2.52-3.16 mm) and B. indica (3.38 mm,

3.02-3.65 mm) (Musser & Brothers, 1994, Table 3). The

posterior cingulum (Fig. 7Bq and v) is absent in the first

and second lower molars (M, and M2) in B. bengalensis

but is present in B. indica and B. savilei.

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Uraipom Pimsai et al.

Karyology (based on extralimital specimens from five lo-

calities in India, Gadi & Sharma 1983). 2N= 42; NF= 58,

59 or 60; with six metacentric, two subtelocentric and 12

acrocentric chromosomes; the X chromosome is variably

a submetacentric, acrocentric or subtelocentric; the Y chro-

mosome is an acrocentric or submetacentric.

Sperm morphology. Information on the sperm morphol-

ogy of a specimen from Penang, Malaysia is included in

Breed & Yong (1986).

Fossil history. No fossil specimens of B. bengalensis are

known from the study area (Pearch et al. 2013).

Taxonomic notes. The two taxa, varius and varillus, de-

scribed from Penang, Malaysia (considered to be inadver-

tent human introductions, Chasen 1936; Musser & Car-

leton 2005 and others) are referred to B. bengalensis

(Agrawal & Chakraborty 1976; Musser & Brothers 1994;

for a contrary view of the taxonomy see Boonsong & Fel-

ten 1989).

Distribution and conservation status. Bandicota ben-

galensis is essentially a South Asian species with a range

that extends from Pakistan, India, Sri Lanka, Nepal, and

Bangladesh to Myanmar. Isolated populations in Penang

Island, Malaysia and Indonesia are thought to be human

introductions (Musser & Carleton 2005). In the study area,

its range is restricted to peninsular Myanmar and Penang

Island (Fig. 8).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, presumed large pop-

ulation, it occurs in a number of protected areas, has a tol-

erance of a degree of habitat modification, and because it

is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Aplin et al. 2008f).

Ecology and reproduction. Virtually no ecological data

are available for this species in the study area. Flower

( 1900, = Nesocia [sic] sp. inc.) noted that it had been col-

lected in gardens, outbuildings and barracks in Penang,

Malaysia, where it was common, causing damage to all

kinds of field crops and also attacking stored grain. Ex-

tral imitally, it is common in both villages and towns where

it is associated with agricultural areas. In Bangladesh, it

is usually more frequently found in higher rainfall areas.

It has a varied diet, with a preference for rice over wheat;

it also consumes molluscs and crabs. It is a good swim-

mer and causes significant damage to deep-water rice

crops. After harvest, it will frequently move from sur-

rounding areas into villages when the fields no longer pro-

vide sufficient food. It has elaborate burrow systems

(Posamentier 1989, Aplin et al. 2003). Males relocate their

burrows more frequently than females. Most feed within

an area of 1 2-40 m2 around the burrow complex, although

those living within a village generally move further (Aplin

et al. 2003). Burrows are constructed in field bunds, veg-

etable gardens and orchards and in the floors and walls

of buildings. The burrows have multiple chambers and en-

trances (sometimes as many as 12-16 per burrow) and

Fig. 8. Distribution of Bandicota bengalensis in the Myan-

mar-Thai-Malaysian peninsula and Singapore. Myanmar: 1.

Mergui (Myeik). Malaysia: 2. Georgetown, Penang Island. For

full locality details, see Gazetteer and Appendix I.

may be used for several generations. Burrow entrances are

either blocked during the day or left unblocked to facili-

tate escape. Since individuals of this species are very ag-

gressive to each other, most burrows are usually occupied

by a solitary adult male or female or females with young

(Posamentier 1989, Aplin et al. 2003). However in areas

of high density, many individuals may live in a burrow

system. In India, breeding activity peaks in the dry sea-

son. The oestrus cycle is 3-5 days and the gestation pe-

riod is 21-25 days. Sexual activity commences from about

three months of age among females, slightly later in males.

Litter size ranges from 1 to 19, with a mean of 6.2 young

(Aplin et al. 2003).

Bandicota indica

Greater bandicoot rat; Large bandicoot rat

Mus indicus Bechstein 1800: 497; Pondicherry, India.

Description (based on personal observation and Musser

& Brothers 1994). This is the largest of the three species

of Bandicota (Table 2). The tail averages shorter than head

and body length but is relatively longer that those of B.

savilei or B. bengalensis. It is virtually naked, unicoloured,

dark brown to black, with larger scales, about eight rows

of scales/cm (Fig. 3D). The fur on the upperparts of the

body is dark, blackish brown; the very long black guard

hairs are particularly well-developed over the rump (Fig.

3C, Fig. 5 IF). The underparts are dark brownish grey;

darker than in B. savilei and B. bengalensis. There is no

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clear demarcation on the Hanks between the upper and

lower pelage. The hindfeet are much larger in compari-

son to the other two species and darker in colour, dark

brown to black. There are 12 mammae (one pectoral, two

postaxillary, one abdominal, and two inguinal pairs).

The skull averages larger than those of B. savilei and

B. bengalensis (skulls illustrated in Musser & Brothers

1994, Figures 5 and 6 and in Marshall 1988, page 429)

in all measurements but is particularly larger in occipi-

tonasal length (ONL), nasal length (LN), length of di-

astema (LD), breadth of bony palate (BBPM3) and length

of the maxillary molar row (ALM'-M3) (Table 2). The in-

cisive foramina extend to the anterior margin, or just pos-

terior to the anterior margin of the first upper molars (M1).

Unlike B. bengalensis, in the posterior part of the brain-

case, the posterolateral surface of the supraoccipital (Fig.

7Axvi) is not greatly reduced (see Musser& Brothers

1 994, Figure 8 for further details). In extralimital materi-

al, the crown width of the first upper molar (M1) in B. in-

dica (3.38 mm, 3.02-3.65 mm) exceeds that of B. ben-

galensis (2.41 mm, 2.13-2.75 mm) and B. savilei (2.81

mm, 2.52-3.16 mm) (Musser & Brothers 1994, Table 3).

The posterior cingulum (Fig. 7Bq and v) is present on the

first and second lower molars (Mj and M2); it is absent

in B. bengalensis.

Karyology (based on extralimital specimens from five lo-

calities in India, Gadi & Sharma 1983). 2N= 42; NF= 66;

there are seven metacentric, four subtelocentric, and nine

acrocentric chromosomes; the X chromosome is a sub-

metacentric; the Y chromosome is a submetacentric. Oth-

er extralimital specimens from Calcutta had a formula of

2N=44 (42-47); NF= 72, 74 (Gadi & Sharma 1983).

Fossil history. No specimens of B. indica are known from

peninsular Thailand although the taxon has been record-

ed from a late middle Pleistocene site in north-eastern

Thailand (Pearch et al. 2013).

Taxonomic notes. All specimens from the study area are

referred to the nominate race.

Distribution and conservation status. Bandicota indi-

ca is known from India, Nepal, Myanmar, southern Chi-

na, and Indochina. It has been introduced into the

Malaysian peninsula and Indonesia (Musser & Carleton

2005). In the study region, it is known from peninsular

Thailand and Malaysia (Fig. 9).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, tolerance of a broad

range of habitats, presumed large population, and because

it is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Aplin et al. 2008b).

Ecology and reproduction. In peninsular Thailand, B. in-

dica was found in a range of habitats including second-

ary evergreen forest in Pet Buri Province and agricultur-

al land in Trang Province, including rice fields (UP un-

published data). In peninsular Malaysia, B. indica was re-

ported as ‘attacking rice fields’ in Kedah and Perlis (Har-

Fig. 9. Distribution of Bandicota indica in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: 1. Pa La

U; 2. Ban Na Pum; 3. Phato Watershed Conservation and Man-

agement Unit; 4. Klong Phraya Wildlife Sanctuary; 5. Khao Ying

Mee; 6. Bang Chak; 7. Tan Ta Ya Phi Rom Temple; 8. Kam

Phaeng Phet. Malaysia: 9. Perlis; 10. Kuala Jerlun. For full

locality details, see Gazetteer and Appendix I.

rison 1956b). Extralimitally, B. indica is found in field and

village/urban habitats and is especially common around

water sources. In most areas the population density is low

and it can be considered only as a minor pest to agricul-

ture. Indeed, where B. indica lives in low densities, its pre-

dation of invertebrates, including molluscs and crabs, may

outweigh the limited damage it causes to crops. Howev-

er, where it lives in high densities it can cause significant

damage to rice and potato crops and also poultry and

stored foodstuffs. When captured, it exhibits great feroc-

ity (Aplin et al. 2003).

In addition to human habitats, B. indica is also known

from uncultivated marshy areas and forest patches. It is

an excellent swimmer, allowing it to exploit a wide range

of both aquatic and terrestrial foods, including molluscs,

crustaceans, water lily fruit, water hyacinths, insects, earth-

worms, and field crops such as rice, vegetables, fruits and

nuts. Its burrow systems range from short tunnels through

to elaborate and extensive complexes with many cham-

bers and entrances, that latter are sometimes below wa-

ter. Large burrow complexes may cause considerable dam-

age to buildings, dams and roadways. They contain nu-

merous adults. Normally, individuals feed close to the bur-

row entrances. In India, breeding is shown to occur

throughout the year, with a higher incidence of pregnan-

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Uraipom Pimsai et al.

cy in the dry season. Litter size is 1 to 8 with a mean of

4.8, although larger litter sizes are also reported. The mean

gestation period is 23 days (Aplin et al. 2003).

Bandicota savilei

Burmese bandicoot rat; Savile’s bandicoot rat

Bandicota savilei Thomas 1916b: 641; Mount Popa, Myanmar,

about 2500 feet.

Bandicota bangchakensis Boonsong & Felten 1989: 202; Bang

Chak, Amphoe Muang, Nakhon Si Thammarat Province, Thai-

land, 8°30’N 100°00’E

Description (based on Musser & Brothers 1994). This is

a medium-sized species, intermediate between the small-

er B. bengaleitsis (based on extralimital specimens) and

the larger B. indica (Table 2). The tail averages shorter

than head and body length; it is relatively much shorter

than in B. indica. It is virtually naked, unicoloured, brown-

ish to greyish brown, with small scales, about 10-12 rows

of scales/cm. The fur on the upperparts of the body is less

coarse than that of B. indica and B. bengaleitsis and is

paler, brownish in colour. There are long guard hairs. The

underparts are grey buff, paler than in B. indica and B. ben-

galensis. There is no clear demarcation on the flanks be-

tween the upper and lower pelage. The hindfeet are rela-

tively short in comparison to those of B. indica. They are

brown or brownish grey. There are 1 2 mammae (one pec-

toral, two postaxillary, one abdominal, and two inguinal

pairs).

In extralimital material, the skull is intermediate in size

between the smaller B. bengaleitsis and the larger B. in-

dica (skulls illustrated in Musser & Brothers 1994, Fig-

ures 5 and 6). Unlike in B. bengaleitsis, the nasals con-

ceal the nasal opening and the upper incisors. The upper

incisors are opisthodont; they are procumbent in B. ben-

galensis. The incisive foramina extend posteriorly between

the first upper molars (M1). Unlike in B. bengaleitsis, the

posterolateral surface of the supraoccipital (Fig. 7Axvi)

is not greatly reduced (see Musser & Brothers 1994, Fig-

ure 8 for further details). In extralimital material, the crown

width of the first upper molar (M1) (2.81 mm, 2.52-3.16

mm) is intermediate between the smaller B. bengaleitsis

(2.41 mm, 2.13-2.75 mm) and the larger B. indica (3.38

mm, 3.02-3.65 mm) (Musser & Brothers 1994, Table 3).

The posterior cingulum (Fig. 7Bq and v) is present on the

first and second lower molars (M, and M2); it is absent

in B. bengalensis.

Fossil history. Within the study area, B. savilei has been

recorded from the late middle Pleistocene locality of Khao

Naphung in Thailand. The taxon is known from two fur-

ther sites in Thailand, one in the west and one in the north-

east of the country (Pearch et al. 2013).

Taxonomic notes. Musser & Brothers ( 1994) included B.

bangchakensis from peninsular Thailand as a synonym of

B. savilei; a view followed by Musser & Carleton (2005).

Distribution and conservation status. Bandicota savilei

Fig. 10. Distribution of Bandicota savilei. in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: l.PakTho

Khao Jeen; 2. Khao Rai; 3. Ban Na Pum; 4. Bang Chak; 5. Kam

Phaeng Phet. For full locality details, see Gazetteer and Appen-

dix 1.

is known from Myanmar, Thailand, and Indochina. In the

study region, it is known from peninsular Thailand (Fig.

10).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ in view of its “wide distribution, tolerance of a broad

range of habitats, presumed large population, and because

it is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Aplin 2008).

Ecology and reproduction. This is an adaptable species,

present primarily in degraded habitats and regarded as a

major agricultural pest in certain areas (Aplin 2008). Ex-

tralimitally, B. savilei is abundant in rice fields in Viet-

nam, although it avoids areas that frequently flood. This

is in contrast to B. indica , which is more tolerant of aquat-

ic habitats. An apparently ‘natural’ population of B. sav-

ilei is recorded from ‘grass beneath teak forest’ in Thai-

land (Marshall 1988). Large burrows are constructed in

bunds and other elevated areas. Maize seems to be one of

the favoured crops found in its diet. Few details are avail-

able concerning breeding behaviour. In Vietnam, a high

proportion of females were found to be pregnant at the end

of the dry season. In Myanmar, females have between five

and eleven embryos (Aplin et al. 2003).

Genus Berylmys

Berylmys rats. White-toothed rats

Berylmys Ellerman 1947; 261; type species Epimys manipulus

Thomas.

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Fig. 11. Skull and dentition of Berylmys bowersii

(54-1287 CTNRC), Doi Suthep-Pui, Chiang Mai

Province, northern Thailand. (A): dorsal, ventral and

lateral view of skull and mandible (Scale 5 mm);

(B): occlusal view of upper (left) and lower (right)

left molars of B. bowersii ( HZM . 1 .40 111), Surat

Thani, peninsular Thailand (Scale 2 mm); (C): left

hind foot of B bowersii (PSUZC-MM20 12.200),

Pak Jam, Trang Province, peninsular Thailand

(Scale 5 mm). Indicative numbers (i-xviii) and ( 1-3)

and letters (a-ae) are explained in the text ‘Descrip-

tion’ for genus Berylmys. M: mouth; lab: labial.

Remarks. According to Musser and Carleton (2005), there

are four extant species of Berylmys. Two species are ex-

tralimital to the current study, namely B. mackenziei

(Thomas), which occurs in north-east India, Myanmar,

China and Vietnam and B. manipulus (Thomas), which is

found in north-east India, northern and central Myanmar

and southern China. Meanwhile, two species occur in the

study area: B. berdmorei, which is essentially an Indochi-

nese species and B. bowersi, which is found in both the

Indochinese and Sundaic subregions (Musser & Carleton

2005). However, the taxonomy of B. bowersi , in particu-

lar, remains problematical. As noted below, Musser &

Newcomb (1983) found considerable morphological

variation amongst specimens referred to the species whilst

Pages et al. (2010) and Latinne et al. (2013b) recognised

two distinct genetic lineages. Meanwhile, in Vietnam, Tran

et al. (2009) described a new genus, Pseudobetylmys ,

which is considered by them to be closely allied to

Berylmys.

Description (based on personal observation and Musser

& Newcomb 1983; restricted to the two species occurring

in the study area). Body size ranges from medium to large

(Table 4). Tail colour varies between species, that of B.

bowersi usually has a distinct white tip, which may ex-

tend for up to two-thirds of its length whereas the tail of

B. berdmorei is either uniformly dark brown or sometimes

slightly paler below. Tail length averages considerably

shorter than head and body length in B. berdmorei but av-

erages longer than head and body length in B. bowersi.

The pelage is dense, smooth and stiff but not spiny; the

hairs are rather uniform in length giving a brush-like tex-

ture (Fig. 5 IE). On the upper surface, the pelage is dark

grey turning slightly browner in older individuals (espe-

cially in B. bowersi). The ventral pelage is white. The up-

per and lower surfaces are sharply demarcated from each

other on the flanks. The ears are large and dark brown.

The hind feet are long and narrow with 6 medium-sized

pads (Fig. 1 1C); the interdigital (1), outer metatarsal (2)

and inner metatarsal (3) pads are without ridges. Sexual-

ly adult males have cutaneous glandular areas in the mid-

line of the stomach and inguinal region, which are stained

with sebaceous secretions. Females of have four pairs of

mammae in B. bowersi and five pairs in B. berdmorei.

The morphology of the skull in three different age class-

es (adult, young adult and juvenile) is illustrated for B.

bowersi in Musser and Newcomb (1983, Figure 16). The

nasals (Fig. llAii) are long, their anterior margins (i) are

bluntly pointed or triangular. The braincase has weak lat-

eral ridges in B. bowersi (iii). The ridges are better devel-

oped in B. berdmorei. The nasolachrymal canals are high-

ly inflated (xiii). The zygomata (iv) are convergent, nar-

rower anteriorly than posteriorly. The diastema (from the

back of the upper canines to the front of the first upper

molar) is very long, almost 30 % of skull length. The in-

cisive foramina (v) are long; the posterior margins (vi) of

the foramina are mostly situated in front of the anterior

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Uraipom Pimsai et al.

margins of the first upper molar (M1) (vii), occasionally

they are level with the anterior margins or in a small mi-

nority of cases posterior to the front faces of M1. The pos-

terior border (viii) of the palate (xi) is about equal to, or

even before, the posterior borders of the last upper mo-

lars (M3). The distance from the back edge of the palate

to the ventral margin of the foramen magnum (x) (post-

palatal length) is shorter than palate length. The tympan-

ic bullae (ix) are variable in size. The supraoccipital (xii)

is large and slopes forward, especially in B. berdmorei.

In each half mandible, the coronoid process (xiv) is well-

developed and the posterior root of the incisor (xviii) is

characteristically large, especially in B. berdmorei. The

emargination (xvi) between the condylar (xv) and the an-

gular (xvii) processes is well defined.

The enamel layers of the upper and lower incisors are

white, pale yellow or pale orange. The upper incisors are

orthodont or slightly procumbent. In the first upper mo-

lar (M1), the first two rows of cusps form V-shaped lam-

inae; in young individuals (as illustrated in Fig. 1 1 B), cusp

tl (a) is separate from cusps t2 and t3 (b), which are fused;

in older individuals cusps 1 1 , t2 and t3 are all fused (Fig.

5B); cusps t4 (c), t5 (d) and t6 (e) are fused; there is one

large cusp posteriorly (f), which represents cusps t8 and

t9 (cusp t7 is absent). In the second upper molar (M2), tl

(g) is large and prominent; t3 is always absent in B. bow-

ersi and very occasionally present in B. berdmorei ; cusps

t4 (h) is separate in very young individuals but is usual-

ly fused with t5 (i) and t6 (j); the third row comprises cusps

t8 and t9 (k), which are fused. The third upper molar (M3)

is rather small; tl (1) is prominent; in young individuals,

cusp t4 (m) is separate from t5 and t6 (n), which are fused;

the last row has only t8 (o); sometimes cusp t4, t5, t6, and

t8 are all fused (Fig. 5B).

In the mandibular toothrow, the first lower molar (M,)

is without an anterocentral cusp; the anterolabial cusp (p)

is fused with the anterolingual cusp (q); in younger indi-

viduals, they are separate from the protoconid (r) and

metaconid (s) but in older individuals all four cusps are

fused; a posterior labial cusplet (t) is usually present and

fused with the hypoconid (u) and the entoconid (v); the

posterior cingulum (w) is visible in younger individuals.

In the second lower molar (M2), the protoconid (x) and

metaconid cusps (y) are fused; an inconspicuous posteri-

or labial cusplet is sometimes present (not illustrated); the

hypoconid (z) and entoconid cusps (aa) are fused; a pos-

terior cingulum is present in younger individuals (ab). The

third lower molar (M3) is rather small; the anterolabial

cusp is absent; the protoconid (ac) is fused with the meta-

conid cusp (ad); the posterior lamina is constituted only

by the entoconid cusp (ae).

Key to species for the two species currently known from

the Myanmar-Thailand-Malaysia peninsula is included in

Table 5.

Berylmys berdmorei

Berdmore’s berylmys; Berdmore’s white toothed rat

Mus berdmorei Blyth 1851: 173; Mergui (Myeik), Myanmar

Description (based on Musser & Newcomb 1983 and

Francis 2008). This is a medium-sized, robust rat (Table

4). The tail is always shorter than the head and body

length, averaging about 79 %, based on extralimital spec-

imens from southeast Thailand; it is dark brown above;

dark brown or greyish-white underneath; it lacks a pale

tip. The fur on the upper parts of the body is iron grey.

The under parts are white. The upper surfaces of the front

and hind feet are white or grey and there are six plantar

pads. Females have ten mammae, one pectoral pair, two

postaxillary pairs and two inguinal pairs. The skull is char-

acterised by its high braincase and large tympanic bullae

(Fig. llAix), although in the holotype of mullulus

(Thomas 1916a) from Myanmar, the bullae are smaller and

the upper incisors slightly procumbent. Photographs of the

skull of B. berdmorei are included in Musser & Newcomb

( 1983, Figures 17, 18, and 26) and Marshall ( 1988, 442).

Karyology (based on specimens from Thailand,

Markvong et al. 1973, Musser & Newcomb 1983). 2N=

40, FN= 66; there are seven pairs of metacentric chromo-

somes, one pair of submetacentric chromosomes, five

pairs of subtelocentric chromosomes, and six pairs of te-

locentric chromosomes; the X and Y sex chromosomes are

telocentrics. A subsequent study of one female specimen

from Thailand by Badenhorst et al. (2009) produced an

essentially similar result, 2N= 40, FN= 64, with the kary-

otype comprising two large, four medium-sized and one

small submetacentric pairs; one large, five medium-sized

and one short acrocentric pairs of chromosomes. The Y

chromosome was a small acrocentric.

Fossil history. No specimens are known from peninsular

Thailand although the taxon has been recorded from a late

middle Pleistocene site in north-eastern Thailand (Pearch

et al. 2013).

Taxonomic notes. According to Musser & Newcomb

( 1983), the geographical variation in this species is little

understood. Specimens from Vietnam and Southeast Thai-

land appear to average larger than those from southern

Myanmar but very little is known about specimens from

this latter region. The taxon mullulus (Thomas 1916a)

from Thagata (just extralimital to the study area) is pos-

sibly a synonym of B. b. berdmorei. However, the holo-

type of B. b. berdmorei, which is in the collections of the

Zoological Survey of India, is apparently badly damaged.

Virtually nothing is known of the species within the study

area of peninsular Myanmar, Thailand, and Malaysia.

Distribution and conservation status. Berylmys berd-

morei is an essentially Indochinese species, including

southern China, Thailand, Lao PDR, Cambodia, and

southern Vietnam. In the study area, it occurs in peninsu-

lar Myanmar and Thailand, north of the Isthmus of Kra

(Fig. 12).

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35

Fig. 12. Distribution of Berylmys berdmorei in the Myanmar-

Thai-Malaysian peninsula and Singapore. Myanmar: 1 . Nathe

Mine; 2. Mergui [Myeik] (no exact location). Thailand: 3.

Phetchaburi. For full locality details, see Gazetteer and Appen-

dix I.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “ in view of its wide distribution, tolerance of a de-

gree of habitat modification, presumed large population,

and because it is unlikely to be declining fast enough to

qualify for listing in a more threatened category” (Aplin

et al. 2008g).

Ecology and reproduction. Little is known about this rat

and nothing is recorded from the study area. Extralimi-

tally, it is a terrestrial species that is known to inhabit

forests from near sea level to at least 1 ,400 m, but is most

common in upland areas where it is reported to favour

swampy habitats. It is occasionally an agricultural pest,

but generally avoids human habitations (Aplin et al.

2008g). In Southeast Thailand, it was found in lowlands

at about 50 metres, where it was uncommon and sparse-

ly distributed in swampy forests and marshy grass (Mar-

shall 1988). In Vietnam, it was found in swampy forests

and marshes from sea level to over 1000 metres (Lunde

& Son 2001 ) and on Con Son Island, it was collected in

forest at 300 metres (Van Peenan et al. 1 970). In Lao PDR,

it was moderately abundant in Luang Prabang Province.

Here it lived both on the valley floor and adjacent slopes,

where its burrows were located in small bamboo thickets

and in fields planted with cassava and sweet potatoes; it

is variously reported as damaging these crops. It was al-

so found in an irrigated rice-field complex. Its burrows are

several metres in length with two entrances and one cen-

tral chamber. Burrows are occupied by individuals of both

sexes (Aplin et al. 2003).

Berylmys bowersi

Bowers’s berylmys, Bowers’s rat

Mus bowersi Anderson 1879: 304; Hotha, Kakhyen Hills, west

Yunnan, China, 4500 feet.

M. ferreocanus Miller 1900b: 140; Trang, peninsular Thailand,

3000 feet.

Description (based on personal observation and Musser

& Newcomb 1983). This is the largest species of

Berylmys (Table 4). The tails of specimens from the study

area are dark above and below and usually have an ex-

tensive white tail tip of between half and two-thirds of the

tail length. However, a recently collected specimen from

Surat Thani Province has only the slightest indication of

a pale tip (UP unpublished data) (Fig. 5 IE). The pelage

is brownish grey on the upperparts; it is white on the un-

derparts and is sharply demarcated on the flanks. The hind

feet are dark above. Yong & Dhaliwal (1970) report an

aberrant individual that had yellow pelage. There are eight

mammae (one pectoral pair; one postaxillary pair and two

inguinal pairs). The skull is characterised by its larger size

in comparison to B. berdmorei but with relatively small-

er tympanic bullae (although the bullae of B. berdmorei

mullulus are also small). The upper incisors are orthodont,

not procumbent as in B. berdmorei. Photographs of the

skull of B. bowersi are included in Musser & Newcomb

(1983, Figures 24 and 25) and Marshall (1988, p. 445).

Karyology (based on specimens from Thailand and

Malaysia, Musser & Newcomb 1983). 2N= 40, FN= 66;

there are seven pairs of metacentric chromosomes, one

pair of submetacentric chromosomes, five pairs of subte-

locentric chromosomes, and six pairs of telocentric chro-

mosomes; the X and Y sex chromosomes are telocentrics.

A study of one male and one female specimen from Thai-

land by Badenhorst et al. (2009) produced a comparable

result, 2N= 40, FN= 64, with the karyotype comprising

two large, four medium-sized and one small submetacen-

tric pairs; and one large, five medium-sized and one short

acrocentric pairs of chromosomes. The X chromosome

was a medium-sized acrocentric; the Y chromosome was

a small acrocentric.

Sperm morphology. Information on the sperm morphol-

ogy of three specimens from peninsular Malaysia is in-

cluded in Breed & Yong (1986).

Fossil history. No specimens are known from peninsular

Thailand (Pearch et al. 2013).

Taxonomic notes. Specimens from south of the Isthmus

of Kra are referred to B. b. ferreocanus and are differen-

tiated from Indochinese specimens by their larger skull

size, relatively longer palate but shorter tympanic bullae;

the tails have a greater proportion of white distally (Muss-

er & Newcomb 1983). Pages et al. (2010) and Latinne et

al. (2013b) recognise two distinct genetic lineages, one

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Uraipom Pimsai et al.

Fig. 13. Distribution of Berylmys bowersii in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: 1. Maprit;

2. Rajjaprabha Dam; 3. Klong Phraya Wildlife Sanctuary; 4.

Nakhon Si Thammarat Province; 5. Pak Jam; 6. Trang; 7. Yala

Province; 8. Narathiwat Province. Malaysia: 9. Kedah State;

10. Maxwell’s Hill; 1 1. Taiping; 12. Fraser’s Hill; 13. Kampong

Janda Baik; 14. Ulu Gombak, 20lh mile Pahang Road; 15. Ke-

pong; 16. Kuala Lumpur; 1 7. Ulu Langat. Not located: Yunong

Jang. For full locality details, see Gazetteer and Appendix I.

sular Thailand, it was collected in Trang at an altitude of

about 920 m (3000 feet) (Miller 1900b). In peninsular

Malaysia, it is more abundant in highlands over 615 m

(2000 feet) than in lowland areas (Harrison 1954a, 1957a,

1957b and Musser & Newcomb 1983) and most speci-

mens are known from 1000-1500 m altitude (Lim 1970,

Francis 2008). It tends to replace Sundamys muelleri in

upland areas (Harrison 1954a). According to Medway

( 1969), it is generally found in gullies and valley bottoms

on hillsides, rarely on the crests or ridges. Its abundance

and population density were discussed by Harrison

( 1 969). It seems to be equally abundant in dry and wet ar-

eas (Lim 1970). In contrast to Bandicota indica, it appears

to be a docile animal (Medway 1969). Its diet is mostly

restricted to vegetable matter (Harrison 1954a) although

Medway (1969) suggests that it may also eat slugs and

snails and Lim (1970) also found insects and even fish

scales in the stomach of one individual. Its mean weight

after ten months was estimated to be 350 g (Harrison

1956a). It is the least arboreal of all the Berylmys (Muss-

er & Newcomb 1983) and its nest sites are found on the

ground, in crevices among rocks, in holes in fallen logs,

in holes in the banks of forest pathways, along forest

streams, and in the holes at the bases of trees in the dry

parts of the forest floor (Lim 1970). Litter size is 2 to 5

with a mean of 4. Typically it lives for about six months

in the wild (Harrison 1955; Medway 1969). Extralimi-

tally, in Lao PDR, B. bowersi appears to be strictly for-

est dwelling, although it is occasionally trapped in crop-

ping areas (Aplin et al. 2003).

from northern Thailand and one from Kanchanaburi

Province and peninsular Thailand. However, the level of

genetic difference between these two lineages is lower

than that between B. bowersi and B. berdmorei (Latinne

et al. 2013b).

Distribution and conservation status. Berylmys bower-

si has a range that extends from northeast India to Myan-

mar, south China, Thailand, Lao PDR, Vietnam, and

Sumatra. In the study area, it occurs in peninsular Thai-

land and Malaysia (Fig. 13).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “as the species is widespread and locally abundant,

at least in areas with suitable tree cover. It is present in

several protected areas, and they are not likely to be de-

clining at a rate to warrant listing in a threatened catego-

ry” (Aplin et al. 2008a).

Ecology and reproduction. In peninsular Thailand, B.

bowersi was collected in secondary evergreen forest in

Surat Thani Province (approx. 400 m. a.s.l.) whilst oth-

ers were trapped in lowland evergreen forest (UP unpub-

lished data). In peninsular Malaysia, B. bowersi is a ter-

restrial species, which is most common in primary forest

and which has also found in lower numbers in disturbed

primary forest and secondary forest (Lim 1970). In penin-

Genus Chiropodomys

Tree mice

Chiropodomys Peters 1868: 448; type species C. penicillatus Pe-

ters (in C. gliroides).

Remarks. Chiropodomys are small arboreal mice, which

are relatively common in Southeast Asia. Six extant

species have been described, of which only one, C.

gliroides , occurs in the study region of the Myanmar-Thai-

Malaysian peninsula. With the exception of C. gliroides ,

the other five taxa have relatively restricted distributions.

Synonyms of the extant species are included in Musser

(1979) and Musser & Carleton (2005).

Description. The generic description is omitted since there

is only one species of this genus in the study area.

Chiropodomys gliroides

Pencil-tailed tree mouse

Mus gliroides Blyth 1856: 721; Cherrapunji, Khasi Hills, Assam,

India.

Chiropodomys penicillatus Peters 1868: 448; probably peninsu-

lar Malaysia.

Description (based on personal observation and Musser

1979). This is a small arboreal rat with a long tail (Table

6), which has a well-defined tuft of hairs at the tip. The

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fur is thick, short (6-8 mm in length), and soft. Guard hairs

are abundant but short, scarcely extending beyond the

dense coat. In specimens from peninsular Malaysia, the

pelage is brightly coloured, reddish brown on the upper

parts, with an orange-red (ochraceous) Hank stripe. The

underside of the head and belly is white and is clearly de-

marcated from the upper surface. The head is wide and

the face is short. The eyes are large and prominent; they

are encircled by a dark ring and are densely surrounded

by thin lashes. The cheeks are buffy; the facial whiskers

are long. The ears are thin, brown and finely haired. In

specimens from north of the Isthmus ofKra (10° 30’N),

the upper pelage tends to be paler; there is usually no flank

stripe; and the belly fur is tinged with cream. Specimens

from Bankachon in southern peninsular Myanmar exhib-

it intermediate characters between the northern, Indochi-

nese form, and the southern, Sundaic form. The fore feet

are white. The hind feet are white variably tinged with

brown. Both the fore and hind feet are highly specialised

for climbing (drawings of fore and hind feet are includ-

ed in Musser 1979, Figure 13). In the hind foot (Fig. 14C),

the first digit (hallux) (4) of the each hind foot is much

shorter than the others and ends in a large fleshy pad (3)

in which a small, nearly tlat nail is embedded; the other

digits are slender and tenninate in smaller pads, which

have short, thin, sharp claws. The fifth toe (2) appears to

be always flexed and capable of being bent over towards

the hallux. The interdigital ( 1 ) and inner metatarsal pads

(5) are grossly enlarged; the outer metatarsal pad (3) is

crescent shaped. Both the palmar (fore foot) and plantar

(hind foot) pads are naked and unpigmented. The tail is

long but not prehensile; it is hairy and tipped with a well-

developed tuft of hairs (see Marshall 1988, page 424),

which are about 4-5 mm in length. It is uniformly brown

or dark brown in colour. Females have two pairs of mam-

mae in the inguinal region.

The skull is wide and short. The rostrum is short and

broad; the anterior margins of the nasals (Fig. 14Ai) are

bluntly rounded. The interorbital area (ii) is broad and the

lachrymal bones (iii) are well-developed. The braincase

is wide and rounded with prominent supraorbital ridges

(iv). The palate (xi) is broad; its posterior border (viii) ex-

tends just beyond the last (M3) upper molars (vii). There-

fore the distance from the back edge of the palate to the

ventral margin of the foramen magnum (x) (postpalatal

length) is shorter than the palatal length. The incisive

foramina (v) are short and wide in specimens from the

Malay Peninsula and islands on the Sunda Shelf and long

and thin in material from Indochina (as illustrated here).

When long, their posterior margins (vi) extend to, or

slightly pass the front margins of, the first molar teeth ( M 1 )

(xii). The tympanic bullae (ix) are moderately developed.

Fig. 14. Skull and dentition of Chi-

ropodomys gliroides, (no specimen number

recorded), Phu Phan District, Sakon Nakhon

Province, north-east Thailand. (A): dorsal,

ventral and lateral view of skull and mandible

(Scale = 5 mm); (B): occlusal view of upper

(left) and lower (right) left molars (Scale =

2 mm); (C); left hind foot of C. gliroides

(modified from Hill I960) (Scale = 5 mm).

Indicative numbers (i-xvii) and ( 1-5) and let-

ters (a-av) are explained in the text ‘Descrip-

tion’ for C. gliroides. M: mouth; lab: labial.

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Uraiporn Pimsai et al.

Each half mandible is short with a small coronoid process

(xiii) and a shallow emargination (xv) between the condy-

lar (xiv) and angular processes (xvi); the posterior root of

the lower incisor (xvii) is indicated but not well-developed.

Photographs of the skull are available in Musser & New-

comb 1983, Figure 102 and Marshall 1988, page 425).

The enamel layers of the upper and lower incisors are

either orange or pale yellow and without grooves; the up-

per incisors are orthodont. The upper and lower molars

are relatively small in relation to the size of the skull; all

upper molars have three roots. In the first upper molar

(M1), cusps tl and t2 (Fig. 14Ba) and t3 (b) are variably

fused or separated in different individuals; on the second

row, cusps t4 (c), t5 (d) and t6 (e) are all separate from

each other in younger individuals; on the third row, cusps

t7 (f), t8 (g) and t9 (h) are also separate; a posterior cin-

gulum (i) is present. The second upper molar (M:) includes

a large and prominent anterior lingual cusp (tl ) (j) and an-

terior labial cusp (t3) (k); on the second row, cusps t4 (1),

t5 (nr) and t6 (n) are separate; on the third row, cusps t7

(o), t8 (p) and t9 (q) are separate; the posterior cingulum

(r) is present; sometimes cusp t8 fuses with the posterior

cingulum. The third upper molar (M3) is elongated; the an-

terior lingual cusp (tl ) (s) is large and prominent; cusp t3

is absent; cusp t4 (t) is separate from t5 and t6 (u), which

may be fused; in the last row, t8 and t9 (v) are fused; a

posterior cingulum (w) is present.

In the mandibular toothrow, all the lower molars have

two large roots. The first lower molar (Mt) has an ante-

rocentral cusp (x); the anterolabial cusp (y) is separate

from the anterolingual cusp (z); the protoconid (aa) is sep-

arate from the metacomd cusp (ab); an anterior labial cus-

plet (Cv2) (ac) and a posterior labial cusplet (Cv3) (ad)

are present; the hypoconid (af) is larger than the entoconid

cusp (ag); labial cusplet 4 (Cv4) (ae) and labial cnsplet 5

(Cv5) (ah) are also present; the posterior cingulum (ai) is

well-developed. In the second lower molar (M2), the an-

terolabial cusp is large and prominent (aj); the protoconid

(ak) is similar in size to the metaconid cusp (al); labial cus-

plets Cv3 (am), Cv4 (an), and Cv5 (aq) are present; the

hypoconid (ao) is larger than the entoconid cusp (ap); the

posterior cingulum (ar) is well-developed. The third low-

er molar (M3) is relatively large; the anterolabial cusp (as)

is present; the protoconid (at) is the same size as the meta-

conid cusp (au); the posterior lamina comprises the ento-

conid cusp (av) only.

Karyology. 2N= 42; there are 1 8 pairs of acrocentric au-

tosomes and two pairs of metacentric-submetacentric au-

tosomes, the X chromosome is a metacentric and the Y

chromosome is a subacrocentric (Yong 1973; Yong et al.

1982).

Sperm morphology. Information on the sperm morphol-

ogy of a specimen from peninsular Malaysia is available

in Breed &Yong ( 1986).

Fossil history. In Thailand, it is known from 12 late

Fig. 15. Distribution of Chiropodomys glimides in the Myan-

mar-Thai-Malaysian peninsula and Singapore. Myanmar: 1.5

miles south of Karathuri; 2. Bankachon. Thailand: 3. Trang

Province. Malaysia: 4. Kedah Peak; 5. Gunong Kutu; 6.

Gunong Tahan; 7. Teloni River; 8. Malaysian peninsular (no ex-

act locality); 9. Gunong Mengkuang; 10. Gunong Benom; 11.

Bukit Kutu; 12. Semangko Pass; 13. Bentong Forest; 14. 161*1

mile of Pahang Road, Kuala Lumpur; 15. Jaram; 16. Bukit

Lagong; 17. Gornbak Forest Reserve; 18. Kepong Forest Re-

serve; 19. Ulu Gornbak; 20. Kuala Lumpur; 21. Tanjong Rabok;

22. Ulu Langat; 23. Ulu Langat Forest Reserve; 24. Bukit Man-

dol; 25. Tamok; 26. Bakok. For full locality details, see Gazetteer

and Appendix I.

Pliocene to late middle Pleistocene sites and one Holocene

site, including six localities in the peninsula (Pearch et al.

2013).

Distribution and conservation status. Chiropodomys

gliroides has a range that extends from western China to

northeast India, Myanmar, Thailand, Lao PDR, Vietnam,

Malaysia, Indonesia. In the study area, it is known from

peninsular Myanmar, Thailand and Malaysia (Fig. 15).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, presumed large pop-

ulation, it occurs in a number of protected areas, has a tol-

erance of a degree of habitat modification, and because it

is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Lnnde et al. 2008a).

Ecology and reproduction. This is primarily an arbore-

al rodent, although it also descends to the ground (Med-

way 1969). According to Hill (1960) it has been collect-

ed at altitudes ranging from 585 metres (1900 feet) to 1630

metres (5300 feet) in peninsular Malaysia, where it occurs

in most types of forest but is most common in areas in

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which bamboo is abundant (Medway 1969). By day, it

nests in hollow trees or the intemodal spaces of large bam-

boos. These bamboos are of any age, from standing green

stems to old dead lengths lying on the ground. To reach

the intemodal space of an intact standing bamboo, it gnaws

a neat circular entry hole of about 25 mm in diameter.

Leaves of a variety of different plants are brought into the

intemodal space as nesting material (Medway 1969).

Gestation is thought to last 21 days. The young are

weaned at 1 7 days of age and reach adult size and appar-

ent maturity within 100 days (Medway 1967). Pregnan-

cies are unevenly distributed through the year, with ac-

cording to Harrison ( 1 955) most ( 1 8 %) occurring between

January-March. However, Medway (1967) found 20 %,

50 %, and 36 % of females collected in Selangor were

pregnant in December, January and April respectively,

with pregnant females found in all months except for June

and July. Litter size is 1 to 3 individuals (mean 2.2). Adult

size is attained at about 100 days (Medway 1969).

Genus Hapalomys

Marmoset-mice

Hapalomys Blyth 1859: 296; type species H. longicaudatus

Blyth.

Background. Hapalomys is a Southeast Asian arboreal rat

(illustrated in Abramov et al. 2012, Figure 2). According

to Musser & Carleton (2005) there are two living

species, H. longicaudatus and H. delacouri Thomas. H.

delacouri is a small bodied species and is confined to

Hainan Island (South China), north Lao PDR and south-

ern Vietnam. The larger H. longicaudatus has wider dis-

tribution and is known from Myanmar, southwest and

peninsular Thailand, and peninsular Malaysia. However,

Abramov et al. (2012), on the basis of new karyotypic da-

ta, suggest that specimens from northern Lao PDR and

Vietnam may be referable to a third species, H. pasquieri

Thomas. Several new fossil species of Hapalomys have

been described from Thailand (Chaimanee 1998) and Chi-

na (Zheng 1993). The geographical origin of the genus is

not clear (Musser & Newcomb 1983).

Description. The generic description is omitted since there

is only one species of this genus in the study area.

Hapalomys longicaudatus

Greater marmoset rat

H. longicaudatus Blyth 1859: 296; Sitang River, Tenasserim,

Myanmar.

Description (based on personal observation and Musser

1972). This is a medium-sized rat (Table 7), which is well

adapted for climbing in bamboo and small branches (see

Musser 1972, Figures 1 and 2 for photographs of live an-

imals). Fur on the upperparts is thick, soft and woolly; it

is greyish-brown or brown. There is an ochraceous (yel-

lowish-orange-brown) strip on the flanks, which separates

the dorsal surface from the essentially white underparts,

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although in some individuals there may be a pale yellow-

ish-brown tinge on the hairs in the axillary, inguinal and

abdominal areas. The face is dark, greyish-brown, includ-

ing the cheeks. The ears are dark brown and relatively

small compared to the size of the head. The feet are large,

adapted for gripping and characteristic in shape (not il-

lustrated here; photographs included in Musser 1972, Fig-

ures 4 and 5). The hind feet, which are brown with paler

hairs on the toes, are broad and wide with long, well-

spaced digits; there are large, ridged, interdigital and

metatarsal pads on the soles of feet, and broadened pads

near the tip of the toes. The first digit (hallux) is semi-op-

posable, with a flattened nail instead of a pointed claw.

The tail is longer than head and body, brown throughout,

well haired on the distal third, and ending in a tuft of hairs.

Females have four pairs of mammae, including one pec-

toral, one postaxillary, and two inguinal pairs.

In the skull, the rostrum is medium sized and wide; the

nasals (Fig. 16Aii) scarcely extend in front of the incisors

(xiv) and have blunt tips (i). The nasolacrimal capsules

(iii) are large and inflated. The interorbital area (v) is wide.

The braincase is broad, inflated, and, in adults, has promi-

nent, well -developed ridges (iv) that form a flange of bone

overhanging the supraorbital region. The posterior border

(x) of the palate (xiii) is about in line with the posterior

borders of the last upper molars (M3) (ix), or in some cas-

es anterior to the posterior borders of M\ The distance

from the posterior border of the palate to the ventral mar-

gin of the foramen magnum (xii) (postpalatal length) is

shorter than palatal length. The incisive foramina (vi) are

narrow; their posterior margins (vii) are situated well in

front of the anterior margins of the first molars (M1) (vi-

ii). The tympanic bullae (xi) are large and globular. Each

half mandible is medium-sized, elongate and with a small

coronoid process (xv); the lower incisor root forms a

prominent process on the mandible (xvi). Photographs of

the skull are available in Musser & Newcomb 1983, Fig-

ure 102 and Marshall 1988, page 423).

The enamel layers of the incisors are orange. The up-

per incisors are orthodont relative to the rostrum. The up-

per and lower molars have a characteristic morphology.

The anterior border of the first upper molar (M1) has two

cingular conules (Fig. 16Ba and b); on the first row, cusps

tl (c), t2 (d) and t3 (e) are separate from each other; on

the second row, cusps t4 (f), t5 (g) and t6 (h) are separate;

on the third row, cusps t7 (i), t8 (j) and t9 (k) are sepa-

rate. In the second upper molar (M2), the anterior lingual

cusp (tl ) (1) and anterior labial cusp (t3) (m) are promi-

nent; on the second row, cusps t4 (n), t5 (o) and t6 (p) are

separate; on the third row, cusps t7 (q), t8 (r) and t9 (s)

are separate. In the third upper molar (M3), the anterior

lingual cusp (tl) (t) is prominent; cusp t3 is absent; cusps

t4 (u) and t5 (v) are separate; in the last row, t8 (w), t9

(x) and the posterior cingulum (y) are fused.

In the mandibular toothrow, the first lower molar ( M , )

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Uraipom Pimsai et al.

Fig. 16. Skull and dentition of Hapalomvs longicaudatus

(BM.63. 1163), Kelanton, peninsular Malaysia. (A): dorsal, ven-

tral and lateral view of skull and mandible (Scale = 5 mm); (B):

occlusal view of upper (left) and lower (right) left molars (Scale

= 2 mm). Indicative numbers (i-xvi) and letters (a-aw) are ex-

plained in the text ‘Description’ for H. longicaudatus. M: mouth;

lab: labial.

has a small anterocentral cusp (z), which is fused to the

anterolingual cusp (aa) in Fig. 16B; the anterolabial cusp

(ab) is separate; a labial cusplet (Cv3) (ac) is present; the

protoconid (ad) is separate from the metaconid cusp (ae);

a labial cusplet (Cv5) (af) is present; the hypoconid (ag)

is separate from the entoconid cusp (ah); there are two

small but well-developed cusplets (ai and aj) on the pos-

terior border of the tooth. In the second lower molar (M2)

a labial cusplet (Cv3) (ak) is present; the protoconid (al)

is separate from the metaconid cusp (am); a labial cusplet

(Cv5) (an) is present; the hypoconid (ao) is separate from

the entoconid cusp (ap); there are two small cusplets (aq

and at ) on the posterior cingulum. In the third lower mo-

lar (M3), the anterolabial cusp is absent; the protoconid

(as) is the same size as the metaconid cusp (at) and the

hypoconid (au) is the same size as the entoconid cusp (av);

there is a posterior cingulum (aw).

Karyology (based on specimens from peninsular

Malaysia, Yong et al. 1982). 2N= 50; there are 23 pairs

of acrocentric autosomes and one pair of subacrocentric

autosomes, the X chromosome is a metacentric and the Y

chromosome a subacrocentric.

Sperm morphology. Information on the sperm morphol-

ogy of a specimen from peninsular Malaysia is included

in Breed & Yong (1986).

Fossil history. Fossil specimens of H. longicaudatus from

the middle Pleistocene are known from four localities in

Thailand, including two sites in the peninsula (Pearch et

al. 2013).

Taxonomic notes. Musser ( 1 972) undertook a detailed re-

view of the genus, including a comparison of H. longi-

caudatus with H. delacouri. Currently, H. longicaudatus

is considered to be monotypic with no subspecies de-

scribed.

Distribution and conservation status. According to

Musser & Carleton (2005), H. longicaudatus is known

from Southwest China, southeast Myanmar, and southwest

Thailand. However, Lunde (2008) referred the Chinese

population to H. delacouri. Within the study area, it has

been recorded from relatively few localities in peninsu-

lar Myanmar, Thailand and Malaysia (Fig. 17).

The conservation status of H. longicaudatus is listed by

IUCN as ‘Endangered’ because “its area of occupancy is

highly restricted to undisturbed bamboo habitat within

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Fig. 17. Distribution of Hapalomys longicaudatus in the

Myanmar-Thai-Malaysian peninsula and Singapore. Myanmar:

l.Bankachon. Thailand: 2. Pattani. Malaysia: 3. Belatop: 4.

Ulu Kelantan; 5. Kelanta; 6. Genting Semangko (Semangko

Pass); 7. Kuala Lumpur (near). For full locality details, see

Gazetteer and Appendix I.

evergreen lowland forest and is probably less than 500

km2, its distribution is severely fragmented, and the ex-

tent of its remaining habitat is probably declining”. It is

thought to be possibly extinct in Thailand and Myanmar

(Aplin & Lunde 2008a).

Ecology and reproduction. According to Medway

( 1969), H. longicaudatus is an arboreal species associat-

ed with the bamboo Gigantochloa scortechinii. During the

day, it sleeps in nests in the intemodal spaces of standing

green, or dead, bamboo stems. Access to these spaces is

through holes, which the rat gnaws in the outer wall of

the bamboo; these entrances, which characteristically have

a 10 mm strip of bamboo skin removed from around them,

can be at any height above the ground. In contrast to the

holes made by Chiropodomys, those of H. longicaudatus

are larger, some 35 mm in diameter. The nests are lined

exclusively with leaflets of bamboo. The diet consists en-

tirely of parts of bamboo, with the flowering shoots be-

ing especially favoured. Further information on the behav-

iour, ecology and diet of this species in Malaysia is in-

cluded in Medway ( 1 964b). Little is known of the species

in Thailand, although a specimen from Khwae Noi Riv-

er in southwestern Thailand was caught in bamboo jun-

gle (Musser 1972). Similarly, a specimen from Bankachon

in peninsular Myanmar was also found among bamboos,

together with Chiropodomys (Musser 1972).

Genus Lenothrix

Lenothrix rats

Lenothrix Miller 1903b: 466; type species L. canus Miller.

Background. The genus Lenothrix includes only one

species, the Sundaic rodent, Lenothrix canus.

Generic description. The generic description is omitted

since the genus is monospecific.

Lenothrix canus

Sundaic lenothrix; Grey tree rat; Hoary rat

Lenothrix canus Miller 1903b: 466, pi. 18; Tuangku Island, Ban-

jak Islands, Sumatra.

Rattus canus malaisia Kloss 1931: 105; near Kuala Lumpur, Se-

langor, Malaysia.

Description (based on Miller 1903b, Kloss 1931, Muss-

er 1981, Musser & Newcomb 1983). This is medium-large

arboreal rat (Table 8) with a body mass of 81-220 g (Muul

& Lim 1971). The fur, which is without spines, is char-

acterised by its woolly texture. The underfur is fine, dense

and woolly, about 12 mm in length in the mid-part of the

back. There are also slender, straight cylindrical hairs

which are about 25 mm in length and in addition some

weak, flattened hairs, which are most abundant on the

flanks and underparts, but are mostly inconspicuous. The

colour of the upper surface is grey or grey-brown. It is

darkest in the midline of the back, where there are some

black hairs. It is palest on the shoulders and flanks. The

demarcation between the upper surface and the belly is

reasonably well-defined on the flanks. The belly is

creamy-white or creamy-buff. The sides of the muzzle are

white; the vibrissae are shiny black. The ears are relative-

ly small and blackish, virtually naked except for some

minute hairs. The fore and hind feet are white. The hind

feet (not illustrated), which are brown with some white

hairs, are short, broad and adapted for climbing in trees.

Most of the plantar surface of each foot is taken up by

large interdigital and metatarsal pads that are textured with

high ridges and deep grooves. The pads at the end of the

digits are swollen, especially on the first toe (the hallux),

which has a small nail-like claw; the other digits have

short, recurved, sharp claws. The two outer digits are se-

mi-opposable. The tail is very long but not prehensile. It

is black at the base and has an extensive white tip, some-

times up to half or more of its length. It is covered with

very short hairs, which turn white and increase in size to-

wards the tip, where they are some 5 mm in length. In the

mid-part of the tail, there are 1 1 rings/cm, but the rings

are not clearly defined over much of the tail. According

to Musser (1981), there are ten mammae: one pectoral

pair; two postaxillary pairs, and two inguinal pairs. How-

ever, Kloss (1931) suggested that there are eight mammae:

three pectoral pairs and one inguinal pair.

In the skull, the rostrum is long and the anterior mar-

gins of the nasals (Fig. 1 8 Ai ) are rounded or blunt. The

interorbital area (ii) is relatively narrow whereas the or-

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Uraiporn Pimsai et al.

Fig. 18. Skull and dentition of Lenothrix canus (BM.49.630),

Kuala Lumpur, peninsular Malaysia. (A): dorsal, ventral and lat-

eral view of skull and mandible (Scale = 5 mm); (B): occlusal

view of upper (left) and lower (right) left molars (Scale = 2 mm).

Indicative numbers (i-xx) and letters (a-ag) are explained in the

text ‘Description’ for/,, canus. M: mouth; lab: labial.

bital area (iii) is large. The braincase has very well-devel-

oped lateral ridges (iv), especially over the posterior part

of the orbit, where the ridges are flange-like. The zygo-

matic plates (xv) are narrow; the zygomata (v) are con-

siderably broader than the braincase. Squamosal roots

(xiv) of the zygomata are set in the mid-part of the sides

of the braincase. The posterior margin (ix) of the palate

(xii) ends before or in line with the last upper molars (M1)

(viii); the distance from the posterior margin of the palate

to the ventral margin of the foramen magnum (xi) (post-

palatal length) is shorter than the palatal length. The in-

cisive foramina (vi) are short and wide; their posterior

margins (vii) end well in front of the anterior margin of

the first molars (M1) (xiii). The tympanic bullae are mod-

erately developed (x). Each half mandible has a large coro-

noid process (xvi); the posterior root of the lower incisor

(xx) and the emargination (xviii) between the condylar

(xvii) and angular (xix) processes are clearly defined. Pho-

tographs of the skull are available in Musser (1981, Fig-

ure 44) and Musser & Newcomb ( 1983, Figure 94).

The enamel layers of the incisors are white or pale yel-

low; the upper incisors are opisthodont or almost ortho-

dont relative to the rostrum. In young individuals most of

the cusps are separate from each other (Fig. 4B); in old-

er individuals (Fig. I8B) they become fused. In the first

upper molar (M1), cusps tl, t2 and t3 (a) are separate in

young individuals but become fused with age; on the sec-

ond row, cusps t4 (b) t5 and t6 (c) are variably fused in

older individuals; cusp t7 (d) is separate from cusps t8 and

t9 (e), which are fused. There is a small posterior cingu-

lum visible in young individuals (Fig. 4B). In the second

upper molar (M2), the anterior lingual cusp (tl ) (f) is large

and prominent and there is a small t3 (g); on the second

row, cusps t4, t5 and t6 (h) are fused; on the third row,

cusp t8 and t9 (i) are fused in older individuals; there is

also a small posterior cingulum visible in younger spec-

imens. The third upper molar (M1) is narrower than the

second upper molar (M2) but only a little shorter; the an-

terior lingual cusp (tl ) (j) is large and prominent; cusp t3

is very small and in older individuals is fused with cusps

t4, t5 and t6 (k) (Fig. 1 8B); the last row has t8 and t9 (1)

are fused together; the posterior cingulum is well-devel-

oped (m).

In the mandibular toothrow, the first lower molar (M,)

has an anterocentral cusp (n), an anterolabial cusp (o) and

an anterolingual cusp (p); there is a small anterior labial

cusplet (q); the protoconid (r) is separate from the meta-

conid cusp (s) in younger individuals; the posterior labi-

al cusplet (t) is separate in younger individuals (Fig. 4B)

and fused with the hypoconid (u) and entoconid (v) in old-

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43

Fig. 19. Distribution of Lenothrix canus in the Myanmar-Thai-

Malaysian peninsula and Singapore. Malaysia: 1 . Muka Head

Forest Reserve, Penang Island; 2. Lerek; 3. Kuala Berang; 4.

Gunong Benom; 5. Bukit Kutu; 6. Ulu Gombak; 7. Bukit

Lagong; 8. Subang; 9. Bukit Lanjang Forest Reserve; 10. Kuala

Lumpur; 1 1 . Pahang Road, 16 miles N.E. of Kuala Lumpur; 12.

Kepong, 8 miles N.W.of Kuala Lumpur; 13. Sungei Buloh; 14.

Bukit Mandol; 15. Kemidak; 16. Tamok; 17. Kudong. For full

locality details, see Gazetteer and Appendix I.

er individuals (Fig. 18B); the posterior cingulum (w) is

well-developed. In the second lower molar (M2), the an-

terolabial cusp (x) is prominent; the protoconid (y) is fused

with the metaconid cusp (z); the hypoconid (aa) is sepa-

rate from the entoconid cusp (ab) in younger individuals

(Fig. 4B); the posterior cingulum is well-developed (ac).

In the third lower molar (M3), the anterolabial cusp is pres-

ent (Fig. 4B) or absent (Fig. 18B); the protoconid (ad) is

separate from the metaconid cusp (ae) in younger individ-

uals; the posterior lamina comprises a very small

hypoconid (af) and a large entoconid cusp (ag) (the two

are fused in the older specimen in Fig. 18B).

Karyology (based on specimens from Malaysia, Yong

1969a; Musser 1981). 2N = 46, FN = 63 (in males) and

62 (in females); there are three pairs of small metacentric

chromosomes, five pairs of subtelocentric chromosomes,

14 pairs of telocentric chromosomes; the X chromosome

is a submetacentric and the Y chromosome a telocentric.

Sperm morphology. Information on the sperm morphol-

ogy of two specimens from peninsular Malaysia is includ-

ed in Breed & Yong (1986).

Fossil history. No specimens are known from Thailand

(Pearch et al. 2013).

Taxonomic notes. Although Lenothrix was treated as a

subgenus of Rattus (Ellerman 1949, Medway 1969), sub-

sequent authors (Misonne 1969; Medway & Yong 1976)

followed Miller (1903b) and included it in its own sepa-

rate genus. Musser (1981) provided an exhaustive list of

distinguishing characters. Kloss ( 1931 ) described a new

subspecies L. c. malaisia, to which he referred specimens

from peninsular Malaysia and Sarawak. However, the va-

lidity of this race is not clear and requires further study.

Distribution and conservation status. Lenothrix canus

is a Sundaic species, which is found in peninsular

Malaysia, Tuankul Island, and Borneo, including Sarawak,

Sabah, and southwest Kalimantan (Musser & Carleton

2005). Its distribution in the study area is illustrated in Fig.

19.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ since it has a “wide distribution, presumed large pop-

ulation, occurrence in a number of protected areas, toler-

ance to some degree of habitat modification, and because

it is unlikely to be declining at nearly the rate required to

qualify for listing in a threatened category” (Ruedas et al.

2008b).

Ecology and reproduction. This is primarily an arbore-

al rodent, which in peninsular Malaysia has been collect-

ed from the foothills of Selangor (Medway 1969) where

it was common in the forests of Bukit Lagong, Bukit Ku-

tu, and Bukit Lanjang and also in the lowland secondary

forests at Subang and Meru. In addition, it was abundant

in the Kampong rubber plantations on Bukit Mandol and

was trapped in primary and disturbed primary forests in

Pahang and Johor (Muul & Lim 1971). Extralimitally, it

is considered to be a common species in primary rain for-

est in the Mount Kinabalu National Park in Sabah (Wells et

al. 2004); it was also collected in primary and secondary

forest in Sarawak (Kloss 1931 ). Mean litter size is 3.0 with

a range of 2 to 6 (Harrison 1955). Average life span in the

wild is five months (Harrison 1956a).

G e nu s L eopoldamys

Long-tailed giant rats

Leopoldamys Ellerman 1947: 261; type species Mus sabanus

Thomas.

Remarks. Leopoldamys is a rather large and semi-arbo-

real rat, which according to Musser & Carleton (2005) is

represented by six living species, of which two, L. sabanus

and L. ciliatus, are found in the study area of peninsular

Myanmar, Thailand and Malaysia. The distribution of the

genus is mapped in Musser (1981, page 265). With the ex-

ception of L. sabanus, and to a lesser extent L. edwardsi,

the other four taxa appear to have relatively restricted

ranges. Leopoldamys milled (Robinson & Kloss) is cur-

rently thought to be restricted to the Langbian Mountains

of Vietnam; Leopoldamys neilli (Marshall) is endemic to

central and northern Thailand; L. siporanus (Thomas) is

endemic to the Mentawai Archipelago of Indonesia; and

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Uraipom Pimsai et al.

L. ciliatus is confined to peninsular Malaysia and the high-

lands of Sumatra.

However, for an alternative view see Balakirev et al.

(2013). They recognise 8 species. Three, L. ciliatus, L.

siporanus and L. milleti, have geographical ranges com-

parable to those outlined in Musser & Carleton (2005).

However, they consider L. sabanus to be restricted to Bor-

neo; specimens formerly referred to L. sabanus in the low-

lands of the Malacca Peninsula, northward to southwest-

ern Thailand are assigned to L. vociferous (Miller); and

specimens from the lowlands of eastern and central In-

dochina are now included in L. revertens (Robinson &

Kloss). The taxon neilli is considered to be a junior syn-

onym of L. herberti (Kloss). Leopoldamys edwardsi has

a range that includes China and northern Vietnam (north-

ward of 21°N). However, it should be noted that this new

taxonomic treatment is contrary to that of Latinne et al.

(2012, 2013a, 2013b).

Description (based on personal observation and Musser

1981; restricted to the two species [sensu Musser & Car-

leton 2005] occurring in the study area). Leopoldamys is

characterised by its large size, long tail, which greatly ex-

ceeds head and body length (Tables 9 and 11), and smooth

fur, which is without spines. The guard hairs on the back

are soft and about equal in length to the overfur, or only

slightly longer. The dorsal pelage is sharply demarcated

from the pale underparts, which are white to creamy white

(Fig. 5 ID). The tail is usually unicoloured, brown (some-

times slightly paler below) in L. ciliatus and bicoloured

in L. sabanus. When bicoloured, its upper surface is

brown, sometimes mottled brown and the lower surface

is pale, tending to white (above and below) towards the

tip. The hind feet are long and slender, each with six plan-

tar pads (Fig. 20C); the interdigital pads (1) are medium

sized; the outer metatarsal pad (2) is rounded and the in-

ner metatarsal pad (3) is elongated. Females have eight

mammae: one pectoral pair, one postaxillary pair, and two

inguinal pairs (Musser 1981), although according to Med-

way (1969) there are two pectoral pairs and two inguinal

pairs.

The skull is long and narrow. The nasals are long with

rounded, blunt anterior margins (Fig. 20Ai). The interor-

bital area is broad (vi) and each lachrymal bone (ii) is

small. In lateral view, the braincase is low; it has well-de-

veloped supraorbital ridges (iii), which extend posterior-

ly to the supraoccipital (v). The zygomata (iv) are virtu-

ally parallel to each other. The zygomatic plates (xv) do

not extend much beyond the anterior root (xvi) of each

zygoma; meanwhile the squamosal root (xvii) is set high

on the side of the braincase. The palate (xiii) is wide and

scored by shallow palatine grooves; its posterior border

(x) ends before, or in line with, the last upper molars (M1)

(ix). The distance from the back edge of the palate to the

ventral margin of the foramen magnum (xii) (postpalatal

length) is shorter than the palatal length. The incisive

foramina are short and broad (vii); their posterior margins

(viii) end well in front of the anterior margin of the first

upper molars (xiv). The tympanic bullae are small (xi) and

pressed against the squamosal bones (xviii). Each half

mandible has a small coronoid process (xx) and a shal-

low emargination (xxii) between the condylar (xxi) and

angular processes (xxiii). The process of the lower inci-

sor root (xxiv) is little developed.

The upper incisors (xix) are strongly opisthodont and

the enamel layers of the upper and lower incisors are bright

orange. In the first upper molar (M1), cusps tl and t2 (Fig.

20Ba) are fused with t3 (b); cusps t4, t5 and t6 (c) are al-

so fused; on the third row, cusps t8 (d) and t9 (e) are fused;

there is no posterior cingulum. In the second upper mo-

lar (M:), cusp 1 1 (f) is rounded and large; cusp t3 is most-

ly, but not always, absent; cusps t4, t5 and t6 (g) are fused;

there is a large t8 (h). In the third upper molar (M3), the

anterior lingual cusp (tl) (i) is large and prominent; cusp

t3 is absent in L. ciliatus but occasionally present in some

specimens of L. sabanus from Malaysia; cusps t4, t5 and

t6 (j ) are fused; the last row comprises only t8 (k); in old-

er individuals, t8 becomes fused with t4, t5 and t6 (Fig.

20B); there is no posterior cingulum.

In the mandibular toothrow, the first lower molar (M,)

is without an anterocentral cusp; the anterolabial cusp (1)

and anterolingual cusp (m) are fused; in the second row,

the protoconid (n) cusp is fused with the metaconid cusp

(o); in the third row, the posterior labial cusplet (p) is sep-

arate in young individuals but otherwise is fused with the

hypoconid (q) and entoconid cusps (r); the posterior cin-

gulum (s) is well-developed. In the second lower molar

(M2), the anterolabial cusp is very occasionally present in

L. ciliatus but absent from L. sabanus (Fig. 20B); the pro-

toconid (t) is fused with the metaconid cusp (u); the pos-

terior labial cusplet (v) is always present, separate in young

individuals, it is fused in older individuals (Fig. 20B) with

the hypoconid (w) and entoconid cusps (x); the posterior

cingulum (y) is well-developed. In the third lower molar

(M3), the anterolabial cusp is absent; the protoconid (z)

is fused with the metaconid cusp (aa); the posterior lam-

ina is constituted only by the entoconid cusp (ab), although

sometimes there is also a small hypoconid.

Key to species for the two species currently recognised

in the Myanmar-Thai-Malaysian peninsula is included in

Table 10.

Leopoldamys ciliatus

Sundaic mountain leopoldamys

Mus ciliata Bonhote 1900: 879; Mount Inas, Perak, Malaya.

Description (based on Bonhote 1900, Yong 1970). This

is a large rat with a long tail (Table 9). The mean weight

of 30 males and 30 females from peninsular Malaysia was

300 g and 284 g respectively (Lim 1970), with a range

(both sexes) of 204-395 g (Muul & Lim 1971). The fur

is short and without long, protruding guard hairs. On the

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XXIV

M

M

-lab lab—

nt

y

ub

Fig. 20. Skull and dentition of Leopoldamys sa-

banus (54-1319 CTNCR), Phu Num Tok,

Sarahuri Province, central Thailand. (A): dorsal,

ventral and lateral view of skull and mandible

(Scale = 5 mm); (B): occlusal view of upper (left)

and lower (right) left molars (Scale = 2 mm); (C):

left hind foot of L. sabanus

(PSUZC-MM20 12.201 ) Phato Watershed Conser-

vation and Management Unit, Chumphon

Province, peninsular Thailand (Scale = 5 mm). In-

dicative numbers (i-xxiv) and (1-3) and letters

(a-ab) are explained in the text ‘Description’ for

Leopoldamys. M: mouth; lab: labial.

upper surface, it is dark brown or greyish, comprising a

light grey under-fur intermixed with longer hairs, which

are pale at their bases and dark at their tips; the flanks are

fawn coloured, without a reddish brown stripe. The fur

is without spines, the guard hairs are grey with black or

brown tips. The underparts are yellowish-white and clear-

ly demarcated from the flanks; frequently there is a me-

dian dark patch (brown or grey) on the chest between the

forelimbs (Medway 1969, = R. edwardsi). The ears are

moderately long, naked and uniformly rounded. The

whiskers are very long and numerous. There are two very

long supraorbital bristles/hairs, which exceed 70 mm in

length (hence the species name) and which differentiate

this species from L. sabanus (bristle length = 40 mm). The

fore and hind feet are a uniform brown, sometimes with

an irregular longitudinal whitish stripe. The hind feet are

long in comparison to those of L. sabanus. The tail is long,

significantly longer than head and body length (Medway

1969, Musser & Newcomb 1983), although in the origi-

nal description of ciliatus, the tail (315 mm) is considered

to be only slightly longer than head and body (290 mm)

(Table 9). It is usually unicoloured, brown, and with many

short, stiff bristles; occasionally specimens have an un-

pignrented patch at the tip or the entire underside pale. The

skull and teeth, which are illustrated in Musser & New-

comb ( 1983, Figure 1 15, = /.. edwardsi), closely conform

to the generic description provided above. According to

Bonhote ( 1900), the skull is similar to that of L. sabanus

but is slightly longer (where the two are sympatric) and

with a greater depth of braincase and less prominent supra-

orbital ridges.

Karyology. 2N = 42, FN = 56 (male and female); there

are three pairs of small metacentric chromosomes, four

pairs of subtelocentric chromosomes, and 1 3 pairs of te-

locentric chromosomes; the X and Y chromosomes are te-

locentrics (= Rattus edwardsi in Yong 1969a; =

Leopoldamys edwardsi in Musser 1981).

Sperm morphology. Information on the sperm morphol-

ogy of two specimens (listed as L. edwarsi ) from penin-

sular Malaysia is included in Breed &Yong ( 1986).

Fossil history. Currently, there are no records of fossil L.

ciliatus from the study area (Pearch et al. 2013).

Taxonomic notes. This taxon was formerly included in

L. edwardsi by Musser (1981) and Corbet & Hill ( 1 992).

However, Musser & Carleton (2005) considered that L.

edwardsi was restricted to the Indochinese region, where-

as specimens from the Sundaic region were referable to

L. ciliatus.

Distribution and conservation status. Leopoldamys cil-

iatus is known from the highlands of Sumatra and penin-

sular Malaysia, where its range is illustrated in Fig. 21.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ since it has a “wide distribution, presumed large pop-

ulation, it occurs in a number of protected areas, has a tol-

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Uraipom Pimsai et al.

Fig. 21. Distribution of Leopoldamys ciliatus in the Myanmar-

Thai-Malaysian peninsula and Singapore. Malaysia: 1. Kedah

Peak; 2. Mt. Inas; 3. Gunong Tahan; 4. Gunong Benom; 5. Buk-

it Kutu; 6. Gunong Menkuang; 7. Bukit Fraser (above Semangko

Pass); 8. Gunong Ulu Kali; 9. Gunong Bunga Buah; 10. Ulu

Gombak, 20,h mile [of] Pahang Road; 11. Ulu Langat Forest Re-

serve; 12. Jelebu District. For full locality details, see Gazetteer

and Appendix I.

erance of a degree of habitat modification, and because it

is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Musser et al. 2008a).

Ecology. In peninsular Malaysia, L. ciliatus occurs in

montane forests with a rather peaty floor, which is car-

peted with mosses and has many crevices at the base of

trees and rocks (Yong 1970, = R. edwardsi ). It is mainly

confined to altitudes in excess of 1 075 metres (3500 feet)

and as such does not usually overlap with L. sabanus

(Yong 1970, Lim 1970, = R. edwardsi). The lowest alti-

tude it was encountered was at 615 metres (2000 feet) at

Kampong Janda Baik in Pahang (Yong 1970). Lim (1970)

suggests that it is restricted to primary forests and although

primarily a ground-dweller, also climbs trees; its nests

were found in tree holes (nearly two metres above the

ground) and in fallen logs. He also suggested that it prefers

drier, well-drained areas although it may visit wetter ar-

eas. It is a terrestrial and possibly arboreal, omnivorous

species which occurs in primary and degraded tropical

moist forest. It occurs in the montane forests of Sumatra

and peninsular Malaysia usually above 1000 m (Musser

et al. 2008a). Its diet includes insects, fruit and vegetable

matter (Lim 1970).

Leopoldamys sabanus

Indomalayan leopoldamys; Long-tailed giant rat

Mus sabanus Thomas 1887: 269; Mount Kinabalu, Borneo, In-

donesia.

Mus vociferans Miller 1900b; 138; Trang, peninsular Thailand,

1000 feet.

M. stridens Miller 1903a: 28; Tioman Island, East Malaysia.

M. stridulus Miller 1903a: 29; Bentinck Island, Mergui Archi-

pelago, Myanmar.

M. matthaeus Miller 1903a: 29; St Matthew Island, Mergui Ar-

chipelago, Myanmar.

M. lucas Miller 1903a: 30; St Luke Island, Mergui Archipela-

go, Myanmar.

E. stentor Miller 1913: 19; James Island, Mergui Archipelago,

Myanmar.

Mus vociferans lancavensis Miller 1900c: Langkawi Island,

Malaysia.

M. v. ter sus Thomas & Wroughton 1909: 535; Teratau Island,

southwest Thailand.

Epimys v. insularum Miller 1913: 19; Domel Island, Mergui Ar-

chipelago, Myanmar.

E. v. clarae Miller 1913: 20; Clara Island, Mergui Archipelago,

Myanmar.

R. s. dictatorius Chasen 1940: 165; Penang Island, Malaysia.

R. s. salanga Chasen 1940: 166; Junk Seylon Island, Southwest

Thailand.

Description (based on personal observation and Miller

1900b; Miller 1900c; Miller 1903a; Thomas and

Wroughton 1909; Miller 1913; Chasen 1940 and follows

the taxonomy of Musser & Carleton 2005). This is a large

rat with a very long tail (Tables 9 and 10). The pelage on

the upper surface is short and the black guard hairs, which

are some 15 mm in length, scarcely extend beyond the

over-fur. The colour varies from darker forms (vociferans,

tersus, matthaeus ) to paler forms ( stridulus , lucas, lan-

cavensis). In general, the mainland form ( vociferans ) has

grey under-fur on the dorsal surface, which is overlain with

golden-orange/yellow hairs, which are grizzled with black

(Fig. 5 1 D). The black is most conspicuous on the back and

the rump and least apparent on the flanks. There is a clear

division between the orange-brown flanks and the ventral

surface. The belly and the inner side of the legs are dull

white to yellowish-white to the hair bases (vociferans) or

creamy buff (lancavensis, stridulus), although in some

forms the yellow is more pronounced (stridens,

matthaeus). The ears are rounded. The muzzle is compa-

rable to the upper surface in colour and the cheeks are or-

ange-buff, like the flanks. The whiskers are black and long.

The supraorbital bristles are about 40 mm in length. The

fore and hind feet are white, irregularly marked with

brown. The footpads are well-developed for climbing. The

tail is characteristically long, always much longer than the

head and body (Tables 9 and 1 1). In general it averages

more than 140 % of head and body length, although there

is some individual and possible geographical variation.

The tail is bicoloured in the basal half, dark brown above

and whitish (unpigmented) below. The half nearer the lip

is usually, but not always, whitish (unpigmented) above

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47

Fig. 22. Distribution of Leopoldamys sabanus in the Myanmar-

Thai-Malaysian peninsula and Singapore. Myanmar: l.Tavoy

Island; 2 King’s Island; 3. Ross Island 4. Bentinck Island; 5.

Kisseraing Island; 6. Domel Island; 7. Malcolm Island; 8. Clara

Island; 9. Sullivan Island; 10. James Island; 1 1 . Bankachon; 12.

St. Luke Island; 13. Hastings Island; 14. St. Matthew Island; 15.

Victoria Island. Not located: Heifer Island. Thailand: 16.

Maprit; 17. Chumpann; 18. Phato Watershed Conservation and

Management Unit; 19. Tha Chang; 20. Rajjaprabha Dam; 21.

Khao Nong; 22 .Khao Tha-Phet; 23. Ban Plai Nam; 24. Klong

Phraya Wildlife Sanctuary; 25. Krung Ching; 26. Khao Luang;

27. Nongkok; 28. Junk Seylon Island; 29. Telok Poh, Pulau Pan-

jang; 30. Klong Tun Sai; 31. Khao Phu Khao Ya National Park;

32. Lam-ra; 33. Pasir Raja, Pulau Lontar; 34. Trang; 35. Pulau

Telibon; 36. Teratau Island; 37. Ban Thou; 38. Ban Ya Kan.

Malaysia: 39. Kaki Bukit; 40. Langkawi Island; 41 . Sungei Kil-

im; 42. Kedah Peak; 43. Penang Island; 44. Ulu Selama; 45. Ulu

Trengganu; 46. Maxwell’s Hill; 47. Taiping; 48. Gunong Ijan;

49. Pangkor Besar Island; 50. Genting; 5 1 . Gunong Benom; 52.

Binting Bidai; 53. Jenka; 54. Klang Gates; 55. Ginting Bidai;

56. 16th. mile of Pahang Road, Ulu Gombok; 57. Fraser’s Hill;

58. Kuala Kubu; 59. Genting Simpah; 60. Templer Park; 61.

Kampong Janda Baik; 62. Ulu Gombak, 201'1 mile [ot] Pahang

Road; 63. Bukit Lagong Forest Reserve; 64. Bukit Lanjang; 65.

Kampong Janda Buah; 66. Kuala Lumpur; 67. Cheras; 68. Ulu

Langat; 69. Bukit Lantar; 70. Bukit Tangga; 71. Tioman Island;

72. Karang; 73. Mount Pulai. For full locality details, see

Gazetteer and Appendix I.

and below. It is coarsely, conspicuously and uniformly an-

nulated with 7 to 8 rings of scalcs/cm in the mid-part and

with numerous short stiff bristles. The skull (Fig. 20A) es-

sentially conforms to the generic description above, with

some individual and geographical variation. Photographs

of the skull are available in Musser & Newcomb 1983,

Figure 115 and Marshall 1988, page 484).

Karyology (based on specimens from Malaysia, Yong

1969a; Musser 1981). 2N = 42, FN = 54 (male and fe-

male); there are two pairs of small metacentric chromo-

somes, four pairs of subtelocentric chromosomes, and 14

pairs of telocentric chromosomes; the X and Y chromo-

somes are telocentrics.

Sperm morphology. Information on the sperm morphol-

ogy of four specimens from peninsular Malaysia is includ-

ed in Breed & Yong (1986).

Fossil history. There are seven localities of fossil L. sa-

banus in Thailand including two late middle Pleistocene

sites in the peninsula (Pearch et al. 2013).

Taxonomic notes. According to Musser (1981) and Muss-

er & Carleton (2005), considerable morphological varia-

tion is apparent between populations from the Indochinese

and Sundaic subregions and among insular populations

from the Sundaic subregion. This may indicate the pres-

ence of more than one species. Subsequently, Latinne et

al. (2013b) suggested that L. sabanus (sensu stricto) is es-

sentially confined to the Sundaic subregion.

Distribution and conservation status. Leopoldamys sa-

banus, as currently understood, is widespread. It ranges

from Bangladesh to Myanmar, Thailand, Indochina and

Indonesia. Its range in the Myanmar-Thai-Malaysian

peninsula is illustrated in Fig. 22. For a contrary view of

its distribution in Thailand, see Latinne et al. (2013b), who

suggested that it was restricted to southern Thailand with

a northern limit of Kanchanaburi Province.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ since it has a “wide distribution, presumed large pop-

ulation, and because it is unlikely to be declining fast

enough to qualify for listing in a more threatened catego-

ry. Additional taxonomic studies will reveal this taxon to

be comprised of several species for which a review of the

Red List assessment will be necessary” (Lunde et al.

2008b).

Ecology and reproduction. Leopoldamys sabanus is

mostly nocturnal and unlike L. ciliatus is mainly found in

lowland forest, especially evergreen forest habitats up to

1200 metres although it has been found above this eleva-

tion at Maxwell’s Hill, Perak (up to 1380 metres) and on

the slopes of Mount Kinabalu (up to 3100 metres) in

Sabah, Malaysia (Marshall 1988; Francis 2008; Lunde et

al. 2008b). Yong (1970) states that it seems to be confined

to dipterocarp forests with a hard substratum carpeted with

fallen leaves and the trees without crevices at the base. Lim

( 1970) suggests that its altitudinal range does not overlap

with that of L. ciliatus and that it prefers disturbed primary

forest and secondary forest to pristine primary forest; its

nests were found in tree holes and in burrows in the

ground. According to Medway (1969), it is mostly ground

dwelling and favours well drained areas. However, it is

able to climb freely and Gorog et al. (2004) and Wells et

al. (2004) noted that it was a semi-arboreal rat that for-

ages on the ground as well as in the canopy and amongst

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Uraipom Pimsai et al.

lianas. According to Lim (1970), the diet consists of in-

sects, fruit, and other vegetable matter (as well as snails).

In South Asia, it is a nocturnal and subterranean species,

which occurs in tropical, subtropical, wet montane tem-

perate forests (Molur et al. 2005).

The young are born in short burrows dug into sloping

hillsides. There are one or two entrances and the nest

chamber is lined with broad leaves, most of which are col-

lected from surrounding shrubs and woody undergrowth.

In Selangor, Malaysia pregnant females were recorded in

all months, but most frequently from July to September

and least frequently from January to March. Mean litter

size is 3.1 (1 to 7) (Medway 1969).

Genus Maxomys

Maxomys rats

Maxomys Sody 1936: 55; type species Mus bartelsii Jentink.

Remarks. Maxomys is an essentially Sundaic genus of ro-

dents. With the exception of M. rajah, M. surifer, M. white-

headi. , the other 14 species have relatively restricted ge-

ographical ranges. Eleven are variously distributed in the

large and small islands on the Sunda shelf and the Thai-

Malay peninsula, south of the Isthmus of Kra. Only one

of these, M. surifer , extends north of the isthmus into In-

dochina. A single species, M. moi Robinson & KJoss, is

endemic to Laos and Vietnam. One species, M. pagensis

Miller, occurs on the Mentawai Islands, off the fringe of

the Sunda Shelf. Four species, M. hellwaldii Jentink, M.

dollmani Ellerman, M. musschenbroekii Jentink, and M.

wattsi Musser, are found on Sulawesi and are the only

members of this genus found to the East of Wallace’s line

(Musser et al. 1979). Four species, M. inas, M. rajah, M.

surifer, and M. whiteheadi , occur in the Myanmar-Thai-

Malaysian peninsula and Singapore (Musser & Carleton

2005). Aspects of the phylogeny, diversity and biogeog-

raphy of the genus were reviewed by Gorog et al. (2004)

and Pages et al. (2010). Most recently, Achmadi et al.

(201 3) recognised two putative new species from Sulawe-

si and Latinne et al. (2013b) identified four highly diver-

gent genetic lineages within M. surifer.

Description (based on personal observation and Musser

et al. 1979; restricted lo the four species occurring in the

study area). The body size ranges from small to medium

(Tabic 12). The tail is shorter to slightly longer than the

head and body. It is finely scaled in all species; it is dark

above and paler below, sometimes with the portion near-

est to the tip white (unpigmented) above and below, but

is without a terminal pencil of hairs. The dorsal pelage

over the head and body is short, soft, dense, and spiny

(Figs 2A, 3B, 3E, and 51 A). The hind feet are always long

and narrow, with smooth and naked plantar surfaces (Fig.

23C), usually with five, smooth pads (four interdigital and

one inner metarsal), lacking the outer metatarsal pad, in

M. inas and M. whiteheadi. If present, as in M. surifer and

M. rajah , this pad is small and rounded (2); the inner

metarsal pad (3) is similar in size to the interdigital pads

(1). In all local species, females have four pairs of mam-

mae: one pectoral, one postaxillary and two inguinal pairs.

In the skull, the rostrum is long and wide (Fig. 23 Aii )

and the nasals (v) extend in front of the upper incisors (xi-

ii). The anterior border of each nasal is triangular anteri-

orly (i). The lachrymal bones (iii) are relatively large. The

zygomata are virtually parallel with each other (vi). The

braincase has well-defined ridges on the dorso-lateral mar-

gins (iv). The palate is relatively short (but exceeds the

postpalatal length); its posterior border (ix) is in line with,

or slightly anterior to, the posterior borders of the last up-

per molars (M') (xii). The incisive foramina are short, wide

and heart-shaped (vii); their posterior margins (viii) are

situated well in front of the anterior margins of the first

molars (xi). The tympanic bullae are small (x). In each half

mandible, the coronoid process (xiv) is small; the ernar-

gination (xvi) is shallow between the condylar (xv) and

angular (xvii) processes.

The incisors have an orange enamel layer; the upper in-

cisors (xiii) are opisthodont. In the first upper molar (M1),

cusp tl (Fig. 23Ba) is separate in younger individuals (Fig.

5C), but subsequently fuses with cusps t2 and t3 (b); on

the second row, cusp t4 (c) may or may not (Fig. 23B) be

separate from cusps t5 and t6 (d), which are fused; the

third row has one large medial cusp which is representa-

tive of both t8 and t9 (e) (cusp t7 is missing). In the sec-

ond upper molar (M:), cusp 1 1 (f) is prominent and large;

cusp t3 (g) is usually present in M. rajah, M. surifer and

M. whiteheadi but is often absent in M. inas ; in younger

individuals, cusp t4 (h) is separate from cusps t5 and t6

(i), which are fused (Fig. 5C); in older individuals t4, t5

and t6 may be fused, as in Fig. (23B); cusps t8 and t9 (j)

are fused. In the third upper molar (M1), cusp tl (k) is

rounded and large; cusp t3 is usually absent; cusp t4 (1)

is separate from cusps t5 and t6 (m) in younger individ-

uals but fused in older individuals (Fig. 5C & D); there

is a single cusp in the third row (n).

In the mandibular toothrow, the first lower molar (M,)

comprises an anterolabial (o) and anterolingual cusp (p),

which are separate in younger individuals (Fig. 5K) but

are fused together in older Maxomys; the protoconid (q)

and metaconid cusp (r) are well-defined; there is a poste-

rior labial cusplet (s) which is separate in young individ-

uals (Fig. 5K) but is fused with the hypoconid (t) and en-

toconid cusps (u) in older individuals; there is a posteri-

or cingulum (v) in M. rajah and M. surifer but this is most-

ly absent in M. inas and M. whiteheadi. In the second low-

er molar (M2), there is usually an anterolabial cusp (w)

in all local species except M. whiteheadi; it is adjacent to

the protoconid (x) and metaconid cusps (y); in M. rajah

and M. surifer, there is also usually a posterior labial cus-

plet (z) adjacent to the hypoconid (aa) and entoconid cusps

(ab) but this is generally absent in M. inas and M. white-

headi; there is a posterior cingulum (ac). In the third low-

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

49

Fig. 23. Skull and dentition of Maxomys surifer

(54-2919 CTNCR), Pong Nam Ron District, Chan-

thaburi Province, Southeast Thailand. (A): dorsal,

ventral and lateral view of skull and mandible (Scale

= 5 mm); (B): occlusal view of upper (left) and low-

er (right) left molars (Scale = 2 mm); (C): left hind

foot of M. surifer (PSUZC-MM2012.202), Phato

District, Chumphon Province, peninsular Thailand

(Scale = 5 mm). Indicative numbers (i-xvii) and

(1-3) and letters (a-af) are explained in the text ‘De-

scription’ for Maxomys. M: mouth; lab: labial.

er molar (M3), the anterolabial cusp is absent; the rneta-

conid (ad) and protoconid cusps (ae) are fused and the pos-

terior lamina is constituted only by the entoconid cusp (af).

In M. rajah, a posterior cusplet is also usually present.

Key to species of the four species currently recognised

in the study area (sensu Musser & Carleton 2005) is in-

cluded in Table 13. However, recent research indicates that

there are additional distinct genetic lineages of Maxomys

within peninsular Thailand that may prove to be discrete

species (Latinne et al. 2013b).

Maxomys inas

Malayan mountain maxomys; Malayan mountain spiny rat

Mus inas Bonhote 1906: 9; Mount Inas, Perak, Malaysia.

Description (based on Bonhote 1906, Medway 1969,

Francis 2008). This is a medium-sized rat (Table 12) with

a mass of 55-105 g (Medway 1969). On the upper sur-

face, the fur is long and thickly interspersed with stiff, flat-

tened spines. It is a uniform, rich yellow-reddish-brown

(ochraceous rufous) grizzled with black. On the ventral

surface, it is grey washed with chestnut to pale yellow-

brown (ochraceous) to pinkish buff. The demarcation be-

tween the upper and lower surfaces is not well defined.

The upper surfaces of the fore- and hind feet are brown-

ish white. There are five plantar pads on the hind feet. In

the type description, the tail is described as being rather

shorter than head and body length (Bonhote 1906) but in

Hill (1960), Medway (1969) and Musser et al. ( 1979), the

measurements appear about equal, whereas according to

Francis (2008) it is slightly longer. The tail is almost naked

and bicoloured, dark above and paler below. The skull is

medium-sized (Table 12).

Karyology (based on specimens from Malaysia, Yong

1969a; Musser et al. 1979). 2N= 42, FN= 83; there are

ten pairs of metacentric chromosomes and ten pairs of sub-

metacentric chromosomes; the X chromosome is a meta-

centric and the Y chromosome is an acrocentric.

Sperm morphology. Information on the sperm morphol-

ogy of two specimens from peninsular Malaysia is includ-

ed in Breed & Yong ( 1986).

Fossil history. Currently, there is no fossil record of M.

inas in the study area (Pearch et al. 2013).

Taxonomic notes. Although described as a discrete

species, this taxon was treated subsequently by Chasen

(1940) as a subspecies of Rattns alticola Thomas (= Max-

omys alticola) described from Borneo. Later, it was recog-

nised again as a distinct species by Medway (1964a), Cor-

bet & Hill ( 1992) and Musser & Carleton (2005).

Distribution and conservation status. Maxomys inas is

endemic to peninsular Malaysia, where its range is illus-

trated in Fig. 24.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ since it “has a relatively wide distribution in mon-

tane parts of the Malay Peninsula (though it may occur

further north), and there are not believed to be any major

threats to the species at present” (Lunde & Aplin 2008).

Ecology and reproduction. According to Musser & Car-

leton (2005), it is essentially restricted to montane habi-

tats in peninsular Malaysia above 900 metres. In Pahang,

Malaysia, it was collected at altitudes ranging from ap-

proximately 1080 metres to 1630 metres (3500 feet to

5300 feet) (Hill 1960). Litter size in one individual was

3 (Medway 1969). Its diet includes caterpi liars (Medway

1964a, 1969).

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Uraiporn Pimsai et al.

Fig. 24. Distribution of Maxomys inas in the Myanmar-Thai-

Malaysian peninsula and Singapore. Malaysia: 1. Mt. Inas; 2.

Maxwell’s Hill; 3. Mt. Brinchang; 4. Gunong Tahan; 5. Gunong

Benom; 6. Bukit Fraser; 7. Bukit Kutu; 8. Gunong Menkuang;

9. Jelebu District. For full locality details, see Gazetteer and Ap-

pendix I.

Maxomys rajah

Rajah maxomys; Rajah spiny rat

Mus rajah Thomas 1894: 45 1 ; Mount Batu Song, Sarawak, Bor-

neo, Malaysia.

Mus pellax Miller 1900b: 147; Trang, peninsular Thailand.

Description (based on Miller 1900b, Hill 1960, Medway

1969 and Musser et al. 1979). This is a medium-large rat

(Table 12) with a mass of 80-265 g (Medway 1969). The

pelage is harsh with many conspicuous spines. The back

and crown of the head are brown ( light brown to dull buff

brown) without any reddish tinge, although the shoulders

and flanks are more ochre in colour. The midline of the

back is often, but not always, darker. However, there are

relatively few black hairs and the numerous spines are grey

not black. There is a clear division on the flanks between

the dorsal and ventral surfaces; the latter is white to the

base of the hairs and often has a dark longitudinal streak

in the midline of the belly (Medway 1969). The inner side

of the legs and upper surface of the feet are white. There

are six plantar pads, including a small outer metatarsal pad.

The face is brown and there are moderately conspicuous

grey-brown eye-rings. The ears are dark brown, and there

is a white spot between the ears in a minority of speci-

mens of the local form, pellax. The tail is reported to av-

erage slightly longer than head and body length in Hill

(1960), Medway (1969) and Pimsai (2012) and slightly

shorter in Musser et al. ( 1 979) (Table 1 2). It is bicoloured,

but not sharply, light brown above and whitish (unpig-

mented) below, the colours becoming indefinite near the

tip. There are about ten rings of scales/cm in the midpart

of the tail; the hairs are reduced in length near the tip. The

skull is medium-large (Table 12) and is essentially simi-

lar to that of M. surifer in all characters, with the possi-

ble exception of the upper molar toothrow, which is slight-

ly longer on average in M. rajah. Photographs of the skull

are included in Marshall (1988, page 449).

Karyology (based on specimens from peninsular

Malaysia, Yong 1 969a, Musser et al. 1 979). 2N= 36, FN=

56; there are six pairs of metacentric chromosomes, three

pairs of submetacentric chromosomes, and eight pairs of

acrocentric chromosomes, both the X and Y chromosomes

are metacentrics.

Fossil history. There are no records of fossil M. rajah

from peninsular Thailand (Pearch et al. 2013).

Taxonomic notes. The taxonomic status of Maxomys ra-

jah has varied over time. Some suggested that rajah and

surifer were conspecific (Ellerman & Morrison-Scott 1951

& 1955, Harrison 1957a) whilst others treated them as sep-

arate species (Chasen 1940, Sody 1941, Ellerman 1949,

Hill 1960, Yong 1972, Corbet & Hill 1992, Musser&Car-

leton 2005). Hill (1960) provides a detailed comparison

of the local race pellax with the nominate form from

Sarawak.

Distribution and conservation status. Maxomys rajah

is known extralimitally from Riau Archipelago, Sumatra

and Borneo (Musser & Carleton 2005); in the study area

it is found in southern peninsular Thailand-Malaysia and

Singapore (Fig. 25).

Its conservation status is listed by IUCN as ‘Vulnera-

ble’ because of “a population decline, estimated to be more

than 30 % over the last ten years, inferred from extensive

loss and degradation of its lowland forest habitat” (Aplin

et al. 2008e).

Ecology and reproduction. According to Francis (2008),

this is a nocturnal and predominantly terrestrial rodent. It

lives in burrows in forest, the entrances of which are loose-

ly plugged with leaves (Yong 1972). It generally occupies

higher, better drained sites than M. surifer (Harrison

1957a, Medway 1969). It is common in appropriate habi-

tat. Nor (2001) recorded 34 specimens between 500 and

1 100 m on Mount Nuang, Ulu Langat, Selangor Province,

Malaysia. Yong (1972) collected specimens on Gunong

Benom between 215 and 770 metres; it tends to avoid in-

teracting with M. surifer and the two species are seldom

found in the same location. In Trang Province, Thailand,

it was collected at an altitude of about 310 metres and had

an average litter size is 3.3 (2-5, n= 3) (Harrison 1955).

In Narathiwat Province, near the Malaysian border it was

collected in tropical rain forest at approximately 200 me-

tres a.s.l. (LIP unpublished data).

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Fig. 25. Distribution of Maxomys rajah in the Myanmar-Thai-

Malaysian peninsula and Singapore. Thailand: 1. Khao Tha-

Phet; 2. Ban Plai Nam; 3. Khao Luang; 4. Trang; 5. Hala Bala

Wildlife Research Station. Malaysia: 6. Ulu Selama; 7. Krian

Road; 8. Runuk Tanjong; 9. Jeram Kawan; 10. Pulau Pangkor

Besar; 11. Jenka; 12. Rawang; 13. Kampong Awak; 14. Ulu

Gombak, 20"' mile [ol] Pahang Road; 15. Bukit Bangkong; 16.

Bukit Lanjang Forest Reserve; 17. Klang Gates; 18. Bukit

Lagong; 19. Subang Forest Reserve; 20. Kuala Lumpur; 21.

Chera; 22. Ulu Langat Forest Reserve; 23. Bukit Dantai; 24.

Bekok; 25. Pulai. Singapore: 26. Singapore Island. For full

locality details, see Gazetteer and Appendix I.

Maxomys surifer

Indomalayan maxomys. Red spiny maxomys. Red spiny

rat

Mas surifer Miller 1900b: 148; Trang, peninsular Thailand.

Mus luteolus Miller, 1903a: 36; St Matthew Island, Mergui Is-

lands, Myanmar.

Mus umbridorsum Miller 1903a: 37; Loughborough Island, Mer-

gui Islands, Myanmar.

Mus casensis Miller 1903a: 38; Chance Island, Mergui Islands,

Myanmar.

Mus bentincanus Miller 1903a: 38; Bentinck Island, Mergui Is-

lands, Myanmar.

Mus domelicus Miller 1903a: 39; Domel Island, Mergui Islands,

Myanmar.

Mus surifer flavidulus Miller 1900c: 189; Langkawi Island,

Malaysia.

Mus surifer butangensis Miller 1900c: 190; Adang Island, Bu-

tang Islands, west Thailand.

Mus surifer microdon Kloss, 1908a: 145; Tioman I, Malaysia

(pre-occupied by Mus microdon Peters, 1852) = Epimys surifer

binominatus Kloss 1915: 223.

Mus surifer flavigrandis Kloss 1911a: 119; East Perhentian Is-

land, East Malaysia.

Mus surifer grandis Kloss 1911a: 119; Great Redang Island, East

Malaysia.

Mus surifer leonis Robinson & Kloss 1911a: 170; Singapore Is-

land.

Epimys surifer pemangilis Robinson 1912: 593; Pemanggil Is-

land, East Malaysia.

Epimys surifer aoris Robinson 1912: 594; Aor Island, East

Malaysia.

Epimys surifer manicalis Robinson & Kloss 1914: 230; Koh Pen-

nan (Island), Northeast Malaysia.

Epimys surifer spurcus Robinson & Kloss 1914: 230: Koh Samui

(Island), Thailand.

Epimys surifer eclipsis Kloss 1916c: 53; Koh Kra (Island), Thai-

land.

Rattus surifer puket Chasen 1940: 169; Junk Seylon Island, West-

ern Thailand.

Rattus surifer telibon Chasen 1940: 170; Telibon Island, West-

ern Thailand.

Rattus surifer muntia Chasen 1940: 1 70; Muk (=Muntia) Island,

Trang, Western Thailand.

Rattus surifer pidonis Chasen 1940: 171; Pipidon Island, West-

ern Thailand.

Description (following the taxonomy of Musser & Car-

leton 2005 and based on personal observation and Miller

1900b, Miller 1900c, Miller 1903a, Kloss 1908a, Kloss

1911a, Robinson & Kloss 1911a, Robinson 1912, Robin-

son & Kloss 1914, Chasen 1940, Medway 1969). This is

a medium-large rat (Tables 12 and 14; Figs 2A, 3E, and

5 1 A) with a mass of 1 39-284 g (Medway 1969). The fur

is characteristically harsh and includes numerous spiny

hairs (Fig. 3B). The underlying colour is a bright reddish

brown, which is paler in some specimens and darker in

others (Fig. 3E). These colours are variably grizzled with

black, which is heavy in some geographical forms and

may almost combine to produce a dark stripe down the

midline of the back. In some, it is darkened on the shoul-

ders, back and rump; in others it is blackish brown on the

posterior half of the back, less so on the shoulders and

head. The Banks range in colour from orange-brown to

buff; there is always less grizzling compared to the upper

surface. There is a clear division between the flanks and

the under surface, which is white to the base of the hairs

in some specimens but yellowish-white to creamy/dirty

buff in others (Fig. 3E). There is occasionally an indis-

tinct buff patch on the centre of the abdomen. The throat

has a band of darker hairs in some specimens; this ranges

in colour and size from narrow, about 1 8 mm, and yellow

to darker, yellowish-orange-buff and about 30 mm in

width. In many individuals, there is no colour band or an

indistinct band. The muzzle sometimes has a whitish or

yellowish-buff patch at the base of the whiskers. The

cheeks are yellowish-orange. There is sometimes a dark

ring of hairs around the eye. The outer surfaces of the legs

are comparable in colour to the flanks. In some individ-

uals, the fore and hind limbs are encircled with yellow-

ish-brown, separating the white inner portion from the

foot. In other individuals, this ring is only present on the

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Uraiporn Pimsai et al.

hind or fore limbs or is absent. The feet are white or dull

white. The hindfeet have six pads, including a small,

rounded outer metatarsal pad (Fig. 23C1).

The tail is variable in length (Table 12), although most-

ly it is about equal to the head and body. However, in some

populations, it is generally longer and in others, it is of-

ten shorter (Table 14). It is bicoloured, dark brown above

and pale (dull white) below (Fig. 2A). Usually, there is a

pale tip, which is equal to one quarter or more than one

third of the tail length. There are 12 rings of scales/cm in

the midpart of the tail; the rings become narrower and less

regular near the tip, where the sparse hairs become more

numerous but shorter; there is no pencil of hairs. The skull

is medium-large (Table 12) and photographs are includ-

ed in Marshall (1988, page 451).

Karyology (based on specimens from Malaysia, Yong

1969a). 2N= 52, FN= 66; there are four pairs of metacen-

tric chromosomes, two pairs of submetacentric chromo-

somes, and 19 pairs of acrocentric chromosomes; both the

X and Y chromosomes are metacentrics. For Thai speci-

mens, Markvong et al. (1973) report that there are five

pairs of metacentric chromosomes, one pair of submeta-

centric chromosomes, 19 pairs of acrocentric chromo-

somes, with the X chromosome being a metacentric and

the Y chromosome a submetacentric (Musser et al. 1979).

Sperm morphology. Information on the sperm morphol-

ogy of two specimens from peninsular Malaysia is includ-

ed in Breed & Yong (1986).

Fossil history. Fossil specimens dating back to the late

Pliocene have been collected from seven localities in Thai-

land including four in the peninsula (Pearch et al. 2013).

Taxonomic notes. It is generally accepted that Maxomys

surifer, as currently understood, is a complex of cryptic

species, requiring further taxonomic revision (Gorog et al.

2004, Aplin et al. 2008c, Achmadi et al. 2013), with

Latinne et al. (2013b) recognising four highly divergent

genetic lineages. Hill (1960) provides descriptions (most-

ly with measurements, see Table 14) and further informa-

tion on the following named forms from the study area:

aoris, binominatus, butangensis, ccisensis, flavidulus, flav-

igrandis, grandis, leonis, luteoloits, manicalis, muntia, pe-

rn angilis, pidonis, puket, spurcus, surifer and telibon. Pre-

viously the taxon surifer was included by Harrison ( 1957a)

in M. rajah, although he noted several distinguishing char-

acters between rajah pellax and rajah surifer.

Distribution and conservation status. Maxomys surifer

is widely distributed, ranging from Myanmar to Indochi-

na and south-west China to Indonesia (Musser & Carleton

2005); in the study area it is found in the Myanmar-Thai-

land-Malaysia peninsula and Singapore (Fig. 26).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ because it is a “very widespread species, it has a pre-

sumed large population, no major threats, and occurs in

many protected areas” (Aplin et al. 2008c).

Fig. 26. Distribution of Maxomys surifer in the Myanmar-Thai-

Malaysian peninsula and Singapore. Myanmar: 1. Tavoy Is-

land; 2. King’s Island; 3. Tenasserim River; 4. Thaget; 5. Ross

Island; 6. Sir John Hayes Island; 7. Bentinck Island; 8. Kisseraing

Island; 9. Dome] Island; 10. Malcolm Island; 11. Lampi Island;

12. Loughborough Island; 13. Maliwun; 14. Bankachon; 15.

Hastings Island; 16. Victoria Point; 17. St. Matthew Island; 18.

Victoria Island. Not located: Taok Plateau. Thailand: 19.

Tap-Li; 20. Maprit; 21. Marmok; 22. BanNa; 23. Tasan; 24. Lam

Nam Kra Buri National Park; 25. Ban Bang Non; 26. De Lisle

Island; 27. Koh Pennan; 28. Koh Surin Nua; 29. Chance Island

(Koh Surin Tai); 30. Koh Yam Yai; 31. Koh Samui; 32. Phato

Watershed Conservation and Management Unit; 33. Koh Rah;

34. Tha Chang; 35. Thung Chalee Wildlife Sanctuary; 36. Ban

Kok Klap; 37. Khao Nong; 38. Plai Nam; 39. Khao Lak; 40.

Khao Luang; 41. Nongkok; 42. Koh Kra; 43. Song Hong; 44.

Chong; 45. Junk Seylon [Phuket Island]; 46. Koh Yoa Yai; 47.

Lam-ra; 48. Pipidon Island; 49. Pak Jam; 50. Trang; 51. Gap;

52. Koh Muk; 53. Nam Tok Ton Te; 54. Telibon Island; 55. Koh

Tarutao; 56. Kuan Khao Wang Forest Park; 57. Khao Num Kang

National Park; 58. Nam Tok Sai Khao; 59. Na Pra Du; 60. Pu-

lo Rawi; 61. Pulo Adang; 62. Wang Bla Chan; 63. Sam Yak

A-Sen; 64. Ban Thon; 65. Goah Tanah; 66. Hala Bala Wildlife

Research Station. Malaysia: 67. Pelarit; 68. Padang Sireh; 69.

Pulo Langkawi; 70. Sungei Kilim, Pulau Langkawi; 71. Sungei

Kubong, Pulau Langkawi; 72. Dayang Bunting; 73. E. Perhent-

ian Island; 74. Great Redang Island; 75. Gurun; 76. Kedah Peak;

77. Temengoh; 78. Telok Bahang, Penang Island; 79. Ulu Sela-

ma; 80. Lenggong; 81. Maxwell’s Hill; 82. Bukit Gantang; 83.

Gunong Ijan; 84. Jeram Kawan; 85. Tanjong Hantu; 86. Kota

Tongkat; 87. Bentong; 88. Ginting Bidai; 89. Rawang; 90. Ulu

Gombak, 20th. mile [of] Pahang Road; 91. Bukit Lagong; 92.

Ayer Kring; 93. Cheras; 94. Tioman Island; 95. Pemanggil Is-

land; 96. Bukit Tampin; 97. Pulau Aur; 98. Bukit Bcsar; 99. Si

Karang; 100. Kangka Kuli; 101. Kangka Ketcho; 102. Pulai; 103.

Johore Bahru; 104. Pelepak. Singapore: 105.Changi. For full

locality details, see Gazetteer and Appendix I.

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53

Ecology and reproduction. Maxomvs surifer is locally

common to quite rare depending on the population sur-

veyed (Aplin et al. 2008c). In Thailand, it was found in a

range of habitats including primary forest in Ranong

Province and in the various islands that comprise Tarutao

National Park, Satun Province. In Trang and Surat Thani

Provinces it was found in secondary forest at approximate-

ly 126 metres a.s.l. In Chumphon and Songkhla Provinces,

it was trapped in evergreen rain forest at approximately

50 metres a.s.l. It appears to be less associated with hu-

man habitation than Rattus tanezumi (UP unpubl. data).

In peninsular Malaysia, it is common in the forests of

the lowlands and hills up to at least 1450 metres (Med-

way 1969). It is nocturnal and mostly terrestrial, living in

burrows in primary and secondary forests, and also found

in rice fields and gardens, especially if the gardens are ad-

jacent to forests, but not in heavily disturbed areas (Cor-

bet & Hill 1992, Francis 2008). It is not found in trees and

it is not commensal (Chaimanee 1998). The natural diet

consists of vegetable material, including fruits and fallen

fruit, supplemented by insects, slugs, and small verte-

brates. The young are born in nest chambers lined with

fresh green leaves cut from surrounding shrubs and un-

dergrowth. The burrows are short and have one or two en-

trances. Mean litter size is 3.3 (2-5) (Medway 1969).

Maxomys whiteheadi

Whitehead’s maxomys

Mus whiteheadi Thomas 1894: 452; Mt Kinabalu, northern Bor-

neo, Malaysia.

Mus asper Miller 1900b: 145; Trang, peninsular Thailand.

Rattus klossi Bonhote 1906: 9; Johore, Malaysia.

Description (based on personal observation and Miller

1900b, Bonhote 1906, Medway 1969). This is a small rat

(Table 12) with a mass of 35-75 g (Medway 1969). The

fur on the upper surface is a reddish-brown, grizzled with

dark brown in some specimens. The fur bases are grey.

There are numerous grey spines with dark tips; these are

particularly abundant on the back and are less numerous

on the flanks and belly. The crown of the head, shoulders

and back are darker brown; the flanks are yellow-brown

to buff. The underside, which is not sharply demarcated

from the dorsal surface is grey or dull buff, fading to buffy

grey on the chin and inner side of the legs; the hairs are

dusky at their bases. In the form klossi , the belly is ap-

parently a paler, creamy buff. There is sometimes a tawny

spot on the chest. The muzzle is brown, greyish on the

sides and there is a dark ring around the eyes. On aver-

age, the tail is shorter than the head and body (Table 12),

it is bicoloured and without a white tip; the tip is also with-

out a pencil of hairs. The hind foot usually has 5 pads; very

occasionally a small outer metatarsal pad is present. The

skull is small (Table 12); photographs are included in Mar-

shall (1988, page 447).

Karyology (based on specimens from Malaysia, Yong

1969a, Musser et al. 1979). 2N= 36, FN= 71; there are

eight pairs of metacentric chromosomes and nine pairs of

submetacentric chromosomes, the X chromosome is a

metacentric and the Y chromosome is an acrocentric.

Sperm morphology. Information on the sperm morphol-

ogy of a specimen from peninsular Malaysia is included

in Breed & Yong (1986).

Fossil history. There are no published records of fossil

M. whiteheadi from Thailand (Pearch et al. 2013).

Taxonomic notes. There is little consistent geographical

variation in this species (Robinson & Kloss 1918, Corbet

& Hill 1992).

Distribution and conservation status. Maxomys white-

headi is known extralimitally from Sumatra, Borneo and

adjacent islands (Musser & Carleton 2005). In the study

area, it is found in peninsular Thailand and Malaysia (Fig.

27).

Its conservation status is listed by IUCN as ‘Vulnera-

ble’ since it is “believed to have undergone a decline of

more than 30 % over the last ten years as inferred from

rate of forest loss in the region, particularly in the low-

lands of Sumatra and Borneo” (Aplin et al. 2008d).

Ecology and reproduction. This is a nocturnal and main-

ly terrestrial rodent, which lives in tall and secondary low-

land forests. It is also found in plantations, rice paddies

and disturbed areas but only if adjacent to forest. It is most

common in forested lowlands and foothills, but has also

been collected in mountains (Medway 1969; Aplin et al.

2008d). According to Medway (1969), it requires a canopy

of either forest trees or bushes in scrub. In Thailand, it was

collected in tropical rain forest in Narathiwat Province (ap-

proximately 200 metres, a.s.l.) and in primary forest in

Phattalung Province (approximately 95 metres a.s.l.) (UP

unpublished data). Its diet includes ants and other insects

as well as plant matter (Francis 2008). There is no signif-

icant seasonal variation in reproductive activity. Litter size

is 1-6 (mean = 3.0, n = 57) (Harrison 1955). On average,

individuals are thought to survive for about 3.5 months

in the wild (Harrison 1956a, Medway 1969).

Genus Mus

Mice

Mus Linnaeus, 1758: 59; type species M. musculus Linnaeus.

Background. According to Musser & Carleton (2005),

there are 38 species of extant Mus , which belong to four

subgenera: Coelomvs Thomas, Mas, Nannonmys Peters,

and Pryomys Thomas. With the exception of Mus muscu-

lus, which has a worldwide distribution thanks to its close

association with man, all other species are confined to

Africa, Europe and/or Asia and most have relatively re-

stricted ranges. In Southeast Asia, six species, boocluga

(Gray), cervicolor Hodgson, cookii Ryley, fragilicauda

Auffray, pahari Thomas, and shortridgei (Thomas) are

confined to the Indochinese subregion and three species,

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Uraipom Pimsai et al.

Fig. 27. Distribution of Maxomys whiteheadi in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: 1. K.hao

Tha-Phet; 2. Ban Plai Nam; 3. Khao Phu Khao Ya National Park;

4. Ban Khao Chong; 5. Trang; 6. Khao Ram; 7. Ban Lam Mai;

8. Bang Lang Dam; 9. Hala Bala Wildlife Research Station.

Malaysia: 10. Pelarit; 1 1. E. Perhentian Island; 12. Kedah Peak;

13. Temengoh; 14. Ulu Selama; 15. Lenggong; 16. Tredoh Peak;

17. Semangoh [Semangko]; 18. Gunong Semanggul; 19. Bukit

Gantang; 20. Labong Edan; 21. Bentong; 22. Ginting Bidei; 23.

Ulu Gombak, 20"’ mile [of] Pahang Road; 24. Bukit Lagong; 25.

Bukit Lanjang Forest Reserve; 26. Kepong; 27. Gombak; 28.

Subang Forest Reserve; 29. Kuala Lumpur; 30. Cheras; 31. Buk-

it Tangga; 32. Nyalas; 33. Padang Tuan; 34. Tamok Forest Re-

serve; 35. Mount Lun Chun; 36. Mount Pulai; 37. Pelepak. Not

located: Sap Patchung. For full locality details, see Gazetteer

and Appendix I.

crociduroides (Robinson & Kloss), terricolor Blyth

(probably an inadvertent human introduction), and vulcani

Robinson & Kloss, to the Sundaic subregion. A further two

species, M. musculus and M. caroli , occur in both the In-

dochinese and Sundaic subregions (both may be inadver-

tant human introductions to the Sundaic subregion) and

are present in the study area of peninsular Myanmar-Thai-

land-Malaysia and Singapore.

Description (based on personal observation and Marshall

1988; Aplin et al. 2003; restricted to the two species oc-

curing in the study area). The body size is small. The tail

is about as long, or longer in some M. caroli, as the head

and body (Table 15). Tail colour varies between species.

In M. musculus , it is uniformly dark brown, in M. caroli,

it is clearly bicolour, dark above and pale below. In the

two local species, the dorsal pelage is without spiny hairs;

it varies from dark grey to deep brown to dark brown. The

ventral pelage is uniformly white or white with grey bases

in M. caroli ; il is greyish brown and not clearly demar-

cated from the upper surface on the flanks in M. muscu-

lus (in specimens from the study region). The hind feet

(Fig. 28C) are elongated. Each has six pads; the interdig-

ital pads (1) are small and situated towards front of the

foot; the outer metatarsal pad (2) is especially small; the

inner metatarsal pad (3) is elongated. Females have five

pairs of mammae.

In the skull, the nasals (Fig. 28 Aii ) are short; their an-

terior margins (i) are rounded or blunt and usually extend

in front of the upper incisors (xii), although not in M. car-

oli. The lachrymals (iii) are small. The braincase is small

with low, indistinct, ridges (iv). The incisive foramina (v)

are long; their posterior margins (vi) usually extend well

beyond the front margins of the anterior molars (M1) (vii),

although in M. caroli, they are shorter and only reach the

anterior borders of M1. The posterior border (viii) of the

palate (xi) extends beyond the last molar (M3). The dis-

tance from the posterior border of the palate to the ven-

tral margin of the foramen magnum (x) (postpalatal length)

is slightly shorter than palatal length. The tympanic bul-

lae (ix) are medium-sized. The coronoid process (xiii) of

each half mandible is narrow and hooked backwards; the

emargination (xv) between the condylar (xiv) and angu-

lar processes (xvi) is well-developed.

The enamel layers of the incisors are orange (dark or-

ange in M. caroli). The upper incisors are opisthodont (xii)

(procumbent in M. caroli). The first upper molar (M1) is

large, equal to or exceeding the combined lengths of the

second (M2) and third molars (M3). In M1, cusp tl (a) is

sometimes fused (especially in older individuals) and

sometimes separate from t2 (b) and t3 (c), which are fused;

cusps t4 (d), t5 (e) and t6 (f) are variably separate or fused

with one another depending on the individual; posterior-

ly, there is one large medial cusp, which includes t8 (g)

and a small t9 (h) (cusp t7 is missing). In the second up-

per molar (M2), cusp tl (i) is large and prominent; a very

small cusp t3 is sometimes present or absent as in Fig.

28B; on the second row cusps t4 (j), t5 (k) and t6 (1) are

variably fused or separate, depending on the individual;

the third row comprises cusp t8 (m) and a small cusp t9

(n), which are fused in older individuals. The third upper

molar (M3) is small; cusp tl (o) is relatively large and

prominent; cusps t4, t5, t6 (p) are fused; the last row has

only 18 (q); sometimes cusp t4, t5, t6, t8 are fused mak-

ing a large part of the molar, especially in a worn tooth

(as in Fig. 28B).

In the mandibular toothrow, the first lower molar (M, )

is without a anterocentral cusp; there is an anterolabial

cusp (n), an anterolingual cusp (s), a protoconid (t) and a

metaconid (u); the hypoconid (v) and entoconid cusps (w)

are often fused; the posterior cingulum (x) is well-devel-

oped. In the second lower molar (M2), there is sometimes

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Murine rodents of the Myanmar- Thai-Malaysian peninsula and Singapore

55

B

Fig. 28. Skull and dentition of Mus musculus

(JTM6912), no data, Thailand. (A): dorsal, ven-

tral and lateral view of skull and mandible

(Scale = 5 mm); (B): occlusal view of upper

(left) and lower (right) left molars (Scale = 2

mm); (C): left hind foot ofM musculus (HZM

457.18657), Beesham, Pakistan (Scale = 5

mm). Indicative numbers (i-xvi) and (1-3) and

C letters (a-af) are explained in the text ‘Descrip-

tion’ for Mus. M: mouth; lab: labial.

an indistinct indication of an anterolabial cusp, otherwise

it is absent; the protoconid (y) is fused with the metaconid

cusp (z); the hypoconid (aa) is fused with the entoconid

cusp (ab); the posterior cingulum (ac) is well-developed.

The third lower molar (M3) is small; the anterolabial cusp

is absent; protoconid (ad) is fused with the metaconid cusp

(ae) ; the posterior lamina comprises the entoconid cusp

(af) .

Key to species for the two species of Mus currently known

from peninsula Myanmar, Thailand, and Malaysia is in-

cluded in Table 16.

Mus caroli

Ryukyu mouse; Long-tailed rice-field mouse

Mus caroli Bonhote 1902: 627; Okinawa, Ryukyu Islands.

Description (based on Bonhote 1902, Marshall 1977a,

Marshall 1988, Aplin et al. 2003). This is a small species

which has a mean mass of 1 1 .0 g (Marshall 1 977a). Char-

acteristically, its tail is distinctly bicoloured, with the up-

per part, very dark, almost black; its length is variably re-

ported as equal to, or exceeding, head and body length

(Table 15). The fur on the upperparts of the body is light

fulvous-brown (brownish-yellow) to brownish-grey. The

underparts are white with a clear line of demarcation on

the flanks. According to Bonhote ( 1902), the fur is soft

and moderately long and is entirely free from spines, al-

though Aplin et al. (2003 ) suggest that the fur on the back

and flanks contains narrow spines and varies in texture

from soft to moderately stiff. The hind feet are variously

described as white (Bonhote 1902), white or white pep-

pered with dark hairs (Aplin et al. 2003) or dark grey to

white (Marshall 1988). The chin and lips are white. The

skull has procumbent upper incisors, which are dark or-

ange. The nasal bones do not hide the incisors when

viewed from above. The incisive foramina are relatively

short; their posterior borders are about in line with the an-

terior borders of the first upper molars (M1).

Karyology. 2N= 40, based on specimens from northern

Thailand; the karyotype is illustrated in Markvong et al.

(1973, Figure 6). Badenhorst et al. (2009) reported that

M. caroli has a 2N= 40 acrocentric karyotype (NFa= 38).

Fossil history. Specimens are known from one late

Pliocene/early Pleistocene site in western Thailand (just

north of the study area) and one late middle Pleistocene

site in northeastern Thailand (Pearch et al. 2013).

Taxonomic notes. Corbet & Hill (1992) suggested that

on the basis of shared characters, such as the pure white

ventral pelage, specimens from peninsular Thailand are

possibly referable to M. caroli ouwensi Kloss from east-

ern Java.

Distribution and conservation status. M. caroli has a

range that extends from Japan to China, Vietnam, Lao

PDR, Cambodia and Thailand. Specimens from Malaysia

and Indonesia may represent inadvertent introductions by

humans (Musser & Carleton 2005). In the study area, it

is recorded from peninsular Thailand and Malaysia (Fig.

29).

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Uraipom Pimsai et al.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “because of its wide distribution, presumed large

population, occurrence in protected areas, tolerance to

some degree of habitat modification, no known major

threats, and because it is unlikely to be declining at near-

ly the rate required to qualify for listing in a threatened

category” (Aplin & Lunde 2008b).

Ecology and reproduction. Mus caroli inhabits rice fields

and other agricultural grassy areas and can subsist on in-

vertebrates (Marshall 1977a). It is most often associated

with man-made habitats and is only rarely collected in nat-

ural habitats; the latter include pine-grass savannah and

dipterocarp forest. Others have been found in grass and

sedge close to water (Marshall 1988). Information on its

occurrence in Kedah, Malaysia is included in Langham

& Ming ( 1 976). In northern Thailand, it was found in pine

savannah (Marshall 1977a). In Myanmar, it is known from

rice and hay fields and also grassy areas in deciduous for-

est (Corbet & Hill 1 992). It nests inside paddy-field dikes

and its globular nest is made of dried grass (Marshall

1977a). Litter size is 5 to 6 (Marshall 1988).

Mus musculus

House mouse

M. musculus Linnaeus, 1758: 62; Upsala, Sweden.

Description (based on personal observation and Marshall

1988; Aplin et al. 2003). This is a small species with a

mean mass of 13.1 g (Marshall 1977a). In material from

the study area, it has a unicoloured, dark brown, tail (oc-

casionally the undersurface of the tail may be slightly

paler), which is about equal in length to the head and body.

The dorsal pelage is dark greyish-brown to brown; the un-

derparts are a slightly paler greyish-brown (in local ma-

terial; elsewhere the belly may be white). In consequence,

there is no clear line of demarcation on the flanks. The

fur is soft throughout without spines. The hind feet are

brown, except for the toes which are paler. The skull (Fig.

28A) has opisthodont, orange, upper incisors and relative-

ly long incisive foramina, the posterior borders of which

are about in line with the mid-part or posterior borders of

the first upper molars (M1). The nasal bones hide the in-

cisors when viewed from above.

Karyology. 2N= 40 based on specimens from Thon Buri,

Bangkok, Thailand; there are 40 telocentric chromosomes

and the karyotype is illustrated in Markvong et al. (1973,

Figure 9).

Fossil history. There are no fossil records of M. muscu-

lus from Thailand (Pearch et al. 2013).

Taxonomic notes. According to Corbet and Hill ( 1992),

specimens from Southeast Asia are provisionally referred

to M. m. castaneus Waterhouse 1843.

Distribution and conservation status. Mus musculus is

closely associated with humans and has an almost world-

wide distribution (Musser & Carleton 2005). In the study

area, it is recorded from very few localities in peninsular

Fig. 29. Distribution of Mus caroli in the Myanmar-Thai-

Malaysian peninsula and Singapore. Thailand: 1. Prachuap-

khirikhan. Malaysia: 2. Titi Batu; 3. Teroi Tua; 4. Kampong

Perigi. For full locality details, see Gazetteer and Appendix I.

Thailand and Malaysia (Fig. 30). However, it is highly

probable that this simply reflects a lack of collecting ef-

fort and reporting rather than a natural scarcity and that

nearly every house in the major cities has a popidation of

M. musculus (Guy Musser pers. comm.).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ since it is “a widespread and abundant species that

thrives in anthropogenic habitats” (Musser et al. 2008c).

Ecology and reproduction. Mus musculus is a commen-

sal species that in Southeast Asia is essentially confined

to houses, food stores, and out-buildings, although it is oc-

casionally found in village gardens and animals pens. It

is not found in cropping areas. Its diet is very varied and

it can cause significant damage to stored foods. It nests

in a variety of sites, including burrows excavated in the

walls and floors of buildings or under piles of straw or

piles of grain bags. In Malaysia, it has an average gesta-

tion period of 20 days. Mean litter size is 4.3 infants (I

to 7). Sexual maturity is attained after 35 days (Aplin et

al. 2003; Medway 1969).

Genus Niviventer

Niviventer rats

Niviventer Marshall 1976: 402; type species Mus niviventer

Hodgson.

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Fig. 30. Distribution of Mus musculus in the Myanmar-Thai-

Malaysian peninsula and Singapore. Thailand: 1. Trang

Province. Malaysia: 2. Penang [Island]; 3. Johore Bahru. Sin-

gapore: 4. Singapore Island. For full locality details, see

Gazetteer and Appendix I.

Remarks. There are 1 6 extant species of Niviventer , which

are divided into two divisions the Niviventer andersoni-

Division and the Niviventer niviventer- Division (Musser

1981 ). The former, includes two species and is restricted

to China. The latter includes the remainder of the taxa,

most of which have geographically restricted ranges but

two, N. cremoriventer and N. fulvescens, are more wide-

spread. These two taxa, together with N. cameroni , a

Malaysian endemic, occur in the study area. Synonyms

of the extant species are included in Musser & Carleton

(2005); for a contrary view of species delimitations see

Balakirev et al. (201 1 )

Description (based on personal observation and Musser

1981; restricted to the three species occurring in the study

area). This is a medium-sized rat (Table 17). The pelage

is dense, with flattened, grooved spines and guard hairs

of variable length. The dorsal pelage is greyish-brown to

reddish-brown. The ventral pelage is uniformly white or

cream-coloured and is sharply demarcated on the flanks

from the dorsal surface. The tail is usually longer than head

and body length, with longer hairs at the tip, which form

an indistinct tuft/pencil in some species. The tail is usu-

ally bicoloured but is monocoloured in one local species,

N. cremoriventer (Fig. 5 IB). The hind foot of the genus

is usually long and slender but is short and broad in N.

cremoriventer. There are six plantar pads (Fig. 3 1C); the

interdigital ( 1 ), outer metatarsal (2) and inner metatarsal

(3) pads are all moderately developed. Females have four

pairs of mammae (one pectoral pair, one postaxillary pair,

and two inguinal pairs).

The skull is long, slender and often flattened. The nasals

(Fig. 3 1 Aii ) are long with rounded or blunt anterior mar-

gins (i) that extend in front of the upper canines (xvi). The

interorbital area is narrow (vii). There are well defined

ridges (v) on the lateral margins of the braincase (vi). The

zygomatic arches (iv) are narrow, not exceeding the width

of the braincase; the zygomatic plates (xvii) are also nar-

row; the squamosal roots (xviii) of the zygomata are sit-

uated moderately high on the braincase. The lachrymal

bones (iii) are small and inconspicuous. The palatal bridge

(xv) is short; its posterior border (xii) is situated either be-

fore, in line with, or just beyond (as illustrated in Fig. 3 1 A)

the posterior margins of the last upper molars (M') (xi).

Therefore the distance from the back edge of the palate

to the ventral margin of the foramen magnum (xiv) (post-

palatal length) is shorter than the palate length. The inci-

sive foramina (viii) are long and narrow; their posterior

margins (ix) extend to, or slightly pass, the front margins

of the first upper molars (M1) (x). The tympanic bullae (xi-

ii) are very small relative to the size of the skull. In each

half mandible, the coronoid process (xix) is small; the

emargination (xxi) between the condylar (xx) and angu-

lar (xxii) processes is relatively shallow; the lower inci-

sor root forms an indistinct process on the mandible (xxi-

ii).

The upper incisors are orthodont or opisthodont (de-

pending on species but also showing individual variation

within species, for example both types are found in N. ful-

vescens) and the enamel layer is smooth and orange. The

first upper molar (M1) is long, narrow and with a simple

cusp pattern; cusps tl, t2, t3 (Fig. 3 1 Ba ) are fused; in the

second row t4, t5 and t6 (b) are also fused; on the third

row, cusp t7 is absent; cusp t8 (c) is enlarged and there is

no posterior cingulum. In the second upper molar (M:),

cusp tl (d) is rounded and large; cusp t3 is mostly absent;

cusps t4, t5, t6 (e) are fused; on the third row, cusps t7

and t9 are absent; cusp t8 (f) is large. In the third upper

molar (M3), cusp tl (g) is rounded and large; cusp t3 is

always absent; in the youngest individuals, t4 is separate

from t5 and t6, which are separate from t8, however in

all older individuals, t4, t5, t6 and t8 (h) are fused (as il-

lustrated here).

In the mandibular toothrow, the first lower molar ( M , )

is without an anterocentral cusp; the anterolabial cusp (i)

is fused with the anterolingual cusp (j); the protoconid (k)

is fused with the metaconid cusp (1); in a minority of in-

dividuals there is a small, separate posterior labial cusp

(m) that becomes fused (as illustrated in Fig. 3 IB) with

the hypoconid (n) and the entoconid cusp (o); the poste-

rior cingulum (p) is well-developed. In the second lower

molar (M2), there is sometimes an anterolabial cusp (not

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Uraipom Pimsai et al.

M M

-lab lab-

B

Fig. 31. Skull and dentition of Niviventer

fulvescens (54-7179 CTNCR), Phuluang

Wildlife Research Centre, Loei, northern

Thailand. (A): dorsal, ventral and lateral

view of skull and mandible (Scale = 5

mm); (B): occlusal view of upper (left) and

lower (right) left molars (Scale = 2 mm);

(C): left hind foot of TV. fulvescens

(PSUZC-MM20 12.203), Na Thawi Dis-

trict, Songkhla Province, peninsular Thai-

land (Scale = 5 mm). Indicative numbers

(i-xxiii) and (1-3) and letters (a-y) are ex-

plained in the text 'Description' for Ni-

viventer. M: mouth; lab: labial.

illustrated in Fig. 3 1 B), which, when present, is fused with

the protoconid (q) and metaconid cusps (r); in a minori-

ty of individuals there is a small, separate posterior labi-

al cusp (s) that becomes fused with the hypoconid (t) and

entoconid cusps (u) in older individuals; the posterior cin-

gulum (v) is well-developed. In the third lower molar

(M3), the anterolabial cusp is absent; the protoconid (w)

and metaconid cusps (x) are fused; the posterior lamina

is constituted only by the entoconid cusp (y).

Key to species for the three species currently known from

the Myanmar-Thai-Malaysian peninsula (sensu Musser &

Carleton 2005) is included in Table 1 8. However, the find-

ings of Pages et al. (2010) and Balakirev et al. (201 1 ) ques-

tion the taxonomy of this genus, as understood here, and

in consequence, future revisions may change species def-

initions.

Niviventer cameroni

Cameron Highlands niviventer; Cameron Highlands

white-bellied rat

Rattus rapit cameroni Chasen 1940: 176; Cameron Highlands,

Pahang, Malaysia, c. 5000 ft (1524 metres).

Description (based on Chasen 1940; Medway 1969 = Rat-

tus fulvescens, and Musser 1981 = TV. rapit cameroni). This

is a medium sized rat (Table 17). The upperparts are bright

reddish brown; the fur is mixed with many spines and long

black guard hairs; the underparts are white. The tail is bi-

coloured, brown above and white (unpigmented) below

from the base to the tip (although Medway 1969 suggests

that it is dark brown on the underside near the base); the

tail is without a conspicuous tuft. The skull is large with

a long and robust rostrum and well-developed incisive

foramina; the tympanic bullae are small. The teeth are

large. Photographs of the skull are available in Musser

1981, Figure 10).

Sperm morphology. Information on the sperm morphol-

ogy of two specimens (listed as Niviventer rapit ) from

peninsular Malaysia is included in Breed & Yong ( 1986).

Fossil history. No fossil material of TV. cameroni has been

recorded from Thailand (Pearch et al. 2013).

Taxonomic notes. Niviventer cameroni was originally de-

scribed as a subspecies of rapit by Chasen (1940), a view

followed by Musser (1981) and Corbet & Hill (1992).

However, Musser & Carleton (2005) treat it as a distinct

species based on its larger cranial and dental measure-

ments and its tail, which is without a conspicuous tuft.

Distribution and conservation status. Niviventer

cameroni is endemic to the Cameron Highlands of penin-

sular Malaysia (Fig. 32).

Its conservation status is listed by IUCN as 'Vulnera-

ble' “because its extent of occurrence is less than 10,000

km’, its distribution is severely fragmented, and there is

continuing decline in the extent and quality of its forest

habitat” (Musser & Ruedas 2008).

Ecology and reproduction. The holotype was collected

at an altitude of about 1524 metres (5000 feet) in the

Cameron Highlands. It is now known from elevations

ranging from 1524 to 2012 metres in these Highlands in

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Fig. 32. Distribution of Niviventer cameroni in the Myanmar-

Thai-Malaysian peninsula and Singapore. Malaysia: 1.

Cameron Highlands. For full locality details, see Gazetteer and

Appendix I.

primary montane tropical forest (Musser & Carleton

2005). It is not known if the species can persist in disturbed

or modified habitats (Musser & Ruedas 2008).

Niviventer cremoriventer

Pencil-tailed rat. Dark-tailed tree rat.

Mus cremoriventer Miller 1900b: 144; Trang, south Thailand,

3000ft.

M. gilbiventer Miller 1903a: 35; Sullivan Island, Mergui Islands,

Myanmar.

E. solus Miller 1913: 22; Teratau Island, southwest Thailand.

Description (based on personal observation and Musser

1973a). This is a largely arboreal, small rat (Table 17) with

a mass 50-105 g (Medway 1969). The upper pelage is

dense and with fine overhairs (up to 20 mm) and numer-

ous flattened, grooved, semi-rigid spines (up to 20 mm)

and longer guard hairs (35-40 mm). The colour is bright

orange to reddish-brown or yellowish, reddish brown,

which is grizzled with black along the rump and back; it

is palest on the flanks and brightest on the shoulders and

thighs (Fig. 5 IB). The underparts are white to cream buff;

the pelage is shorter (8 to 10 mm) and includes some flat-

tened, flexible spine-like hairs. The upper and lower-parts

of the body are clearly demarcated along the flanks, where

often there is a thin, orange strip. The tail is considerably

longer than the head and body and is dark brown, above

and below (Fig. 5 IB); it is quite hairy, with a conspicu-

ous tuft/pencil of hairs extending 5 to 8 mm beyond the

tail tip. The face is short, the eyes are ringed in brown-

ish-orange fur, and the ears are large, covered in short, fine

hairs, and are pale to dark brown in colour. The upper sur-

faces of the front and hind feet have short, fine, white

hairs; the claws of the hind feet are about 4 to 5 times the

size of those of the front feet; the hind foot of N. cre-

moriventer is illustrated in Musser (1973a, Figure 1 ). Fe-

males have four pairs of mammae. The skull is small and

compact; it is illustrated in Musser (1981, Figure 10). The

rostrum is short, broad and blunt. The incisive foramina

tend to terminate in line with or before the anterior bor-

ders of M1. The tympanic bullae are very small.

Fig. 33. Distribution of Niviventer cremoriventer in the Myan-

mar-Thai-Malaysian peninsula and Singapore. Myanmar: 1.

Sullivan Island. Thailand: 2. Khlong Wan [Tapli]; 3. Tasan; 4.

Ban Bang Nawn; 5. Ban Bang Non; 6. Khlong Saeng; 7. Klrao

Luang; 8. Trang; 9. Chong; 1 0. Muang; 1 1 . Klrao Num Kang Na-

tional Park; 12. Pattani; 13. Nam Tok Sai Khao; 14. Tarutau Is-

land; 15. Koh Rawi; 16. Pulo Adang. Malaysia: 17. Pulo

Langkawi; 18. Bukit Besar; 19. Kedah Peak; 20. Penang Island;

21. Telok Bahang; 22. Temengoh; 23. Lenggong; 24. Gunong

Semanggul; 25. Gunong Ijau; 26. Maxwell’s Hill; 27. Gunung

Kerbau; 28. Kuala Berang; 29. Brinchang Peak; 30. Tanjong

Hantu; 31. Lubok Tamang; 32. Tanjong Dungan; 33. Bukit Be-

si; 34. Kuala Tahan; 35. Tahan; 36. Genting; 37. Gunung Siny-

om; 38. Tanjong Malim; 39. Genting Sempat; 40. Ulu Gombok;

41. Lubok Tamang; 42. Bukit Lagong; 43. Bukit Lagong For-

est Reserve; 44. Kuala Lumpur; 45. 27 km. north-east of Kuala

Lumpur; 46. Kepong; 47. Bukit Mandol; 48. Subang; 49. Pa-

hang Road, 16 miles north-east of Kuala Lumpur; 50. Tioman

Island. For full locality details, see Gazetteer and Appendix I.

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Uraipom Pimsai et al.

Karyology (based on specimens from peninsular

Malaysia, Yong 1969a; Musser 1973a). 2N= 46, FN= 54

with one pair of subterminal autosomes, 18 pairs of acro-

centric autosomes, and three pairs of metacentric auto-

somes; the X and Y chromosomes are acrocentric.

Fossil history. No fossil material of N. cremoriventer has

been recorded from Thailand (Pearch et al. 2013).

Taxonomic notes. Specimens from the study area are re-

ferred to the nominate form, N. c. cremoriventer. The two

island taxa, gilbiventer and solus, are discussed in detail

by Musser (1973a) who treats them as synonyms of the

nominate form. According to Latinne et al, (2013b), mi-

tochondrial markers (a 85 bp cytb fragment) clearly dis-

criminate between N. cremoriventer and N. fulvescens.

Distribution and conservation status. Niviventer cre-

moriventer is found in peninsular Thailand and Malaysia,

and some offshore islands; including the Myeik Archipel-

ago of Myanmar, Anambas Islands, Sumatra, Nias, Billi-

ton, Bangka, Borneo, Java, Bali, and small islands off the

northern tip of Sabah (Musser and Carleton 2005). Its dis-

tribution in the study area is mapped in Fig. 33.

Listed as ‘Vulnerable’ because of a “population decline,

estimated to be more than 30 % over the last ten years,

inferred shrinkage in distribution, and habitat destruction

and degradation” (Ruedas et al. 2008e).

Ecology and reproduction. On Tarutao Island and Rawi

Island, off the west coast of peninsular Thailand, N. cre-

moriventer was collected in primary forest (UP, unpub-

lished data). However, in Malaysia, it is reported as be-

ing common in secondary-growth forests but uncommon

or absent in primary forest. It is nocturnal and mostly ar-

boreal; most specimens have been trapped between two

and four metres off the ground, although occasionally they

are trapped at ground level (Musser 1973a). Its diet com-

prises vegetable matter, fruits and insects. It becomes sex-

ually mature at a weight of about 50 g; litter size varies

from 2 to 5, with a mean number of 3.7 (Harrison 1954b).

Niviventer fulvescens

Indomalayan niviventer; Chestnut white-bellied rat

Mus fulvescens Gray 1847: 18; Nepal.

Mus bukit Bonhote 1903b: 125; Bukit Besar, Jalor, Malaysia,

2500 feet.

Epimys lepidus Miller 1913: 20; Bok Pyin, South Tenasserim,

Myanmar.

Epimys orbus Robinson & Kloss 1914: 228; Kliao Nawng, Ban-

don, Southern Thailand, 3500 feet.

Epimys jerdoni pan Robinson & Kloss 1914: 229; Samui Island,

Thailand.

Description (based on personal observation and Bonhote

1903b; Medway 1969 = Rattus niviventer ; Musser 1981;

Abe 1983). Niviventer fulvescens is the medium size rat

(Table 17) with a mass of 70 g (Medway 1969). The up-

perparts have yellowish-brown to orange-brown pelage,

which is grey at the base. This is intermixed with many

short, flattened spines, which vary in colour from white

to dark brown and are variably tipped with black. There

are short black guard hairs, which are especially abundant

on the lower back. The underparts are white or yellow-

ish-white. There is a sharp line of demarcation on the

flanks. The tail is bicoloured, but not always markedly so,

dark brown above and white below; it has fine silky hairs

which are longer at the tip. The tail is usually longer, some-

times considerably longer than head and body length. The

cheeks, sides of the head and neck are yellowish-orange-

brown without dark grizzling. White extends on to the

wrists but not on to the ankles. The upper surfaces of the

fore and hind feet are dirty white, each with a brown me-

dian strip, which does not reach the base of the digits. The

skull is medium-sized with a robust rostrum and relative-

ly short incisive foramina; as with all Niviventer, the tym-

panic bullae are small. The teeth are unremarkable. Pho-

tographs of the skull are available in Musser 1981, Fig-

ure 1 1 ).

Sperm morphology. Information on the sperm morphol-

ogy of a specimen (listed as Niviventer bukit ) from penin-

sular Malaysia is included in Breed & Yong (1986).

Fossil history. Niviventer fulvescens is known from six

fossil localities in Thailand dating back to the early to late

middle Pleistocene. Three of the localities are in the

peninsula (Pearch et al. 2013).

Taxonomic notes. Corbet & Hill (1992) referred speci-

mens from north of the Isthmus of Kra to N. fulvescens

and those from the south to N. bukit ; this view was con-

trary to Abe (1983), who considered bukit to be the local

subspecies of N. fulvescens in the Thai peninsular. This

latter view was followed by Musser & Carleton (2005) but

was challenged by Balakirev et al. (2011), who treated

bukit as a discrete species with an Indochinese and Sunda-

ic distribution. Pages et al. (2010) also suggested that there

was more species diversity within fulvescens than current-

ly recognised. Over the years, N. fulvescens has been con-

fused with N. confucianus, N. niviventer, and N. tenaster.

Musser & Carleton (2005) recommend further research,

especially with DNA sequences, to determine the relation-

ships of all these closely related taxa. Latinne et al. (2013b)

discriminated between this species and N. cremoriventer

using mitochondrial markers (a 85 bp cytb fragment).

Distribution and conservation status. Niviventer ful-

vescens has a range that extends from Nepal and north-

ern India to Bangladesh, south China, Myanmar, Thailand,

and Indochina. It is also known from peninsular Myan-

mar-Thailand-Malaysia, Sumatra, Java and Bali (Musser

& Carleton 2005). Its distribution in the study area is

mapped in Fig. 34.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, presumed large pop-

ulation, it occurs in a number of protected areas, has a tol-

erance of a degree of habitat modification, and because it

is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Musser et al. 2008b).

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Fig. 34. Distribution of Niviventer fulvescens in the Myanmar-

Thai-Malaysian peninsula and Singapore. Myanmar: 1.

Bokpyin; 2. Bankachon; 3. Victoria Point. Thailand: 4. Khao

Mon; 5. 30 km. west of Chumphon; 6. Tap-Li; 7. Koh Samui;

8. Ao Ko; 9. Khao Nawng; 1 0. Khlong Saeng; 1 1 . Ban Phi Tham;

12. Khao Luang; 13. Tang Plan; 14. Nongkok; 15. Tone Sai Wa-

terfall National Park; 16. Trang; 17. Nam Tok Tamot; 18. Khao

Ram; 19. Nam Tok Sai Khao; 20. Mae Wad. Malaysia: 21.

Pelarit; 22. Kota Bharu; 23. Temengoh; 24. Lenggong; 25.

Maxwell’s Hill; 26. Taiping; 27. Mt. Brinchang; 28. Bukit Be-

sar. For full locality details, see Gazetteer and Appendix 1.

Ecology and reproduction. Niviventer fulvescens was

collected in tropical rain forest in Songkhla Province at

approximately 150 metres a.s.l. (UP unpublished data).

Marshall ( 1 988 = Rattus bukit) suggested that it was com-

mon in the lowland evergreen forest of peninsular Thai-

land; it was found in a similar habitat at Bankachon in

peninsular Myanmar (Wroughton 1915). In northern Thai-

land, N. fulvescens was found in evergreen, pine and de-

ciduous forests and on the Korat Plateau it was common

in evergreen forest and secondary forest (Marshall 1988).

According to Francis (2008), it is a mostly nocturnal

species, which is found in forest habitats, gardens and dis-

turbed areas with vegetation. It is active both on the ground

and in trees, climbing on large lianas and vines.

Genus Pithecheir

Woolly tree rats, Pithecheir rats

Pithechir [sic] Muller 1839: 12 ( nomen nudum).

Pithecheir Lesson 1840: 264; type species Pithecheir melanu-

rus Lesson.

Remarks. Pithecheir is a medium-sized arboreal rat which

is endemic to the Sundaic subregion. There are two ex-

tant species, P. parvus , which is restricted to the Malaysian

peninsula and P. melanurus Cuvier, which is found in Ja-

va ( Musser & Carleton 2005).

Description. The generic description is omitted since there

is only one species of this genus in the study area.

Pithecheir parvus

Monkey-footed rat, Malayan woolly tree rat

Pithecheir melanurus parvus Kloss 1916b: 250; Bukit Ku-

ril. Selangor, Malaya, 3400ft.

Description (based on personal observation and Kloss

1916b, Misonne 1969; Medway 1969 and Musser & New-

comb 1983). This is a medium-sized rat, which together

with P. melanurus, is one of the most arboreal murine ro-

dents of the Sundaic region. The tail is naked, uniformly

brown and prehensile distally (towards the tip); it aver-

ages 1 13 % of head and body length (Table 19). The fin-

is very long, soft and without spines. On the upper sur-

face, the pelage is brownish-red, most richly coloured on

the head and upper back; the hair bases are slate-grey. The

underparts are creamy-white. The chin, flanks and lower

parts of the hind limbs are buffy-brown. There is a clear

line of demarcation between the upper and lower surfaces.

The ears are short and translucent; whitish at the base. The

feet are brown above and highly adapted for climbing

(hence the English name 'monkey-footed rat’); the hind

feet (not illustrated) are short and broad with very large

interdigital and metatarsal pads; the first toe (hallux) is

widely separated from the others with an enlarged pad at

the tip, which has a very short claw; all four other toes

also have enlarged pads at the tips. There are two pairs of

inguinal mammae.

The skull is elongated and medium-sized. The nasals

(Fig. 35 Ai i ) extend beyond the anterior border of the in-

cisors (xii); their anterior margins (i) are rounded and

blunt. The interorbital area (iii) is wide. The braincase is

wide and when viewed from the side is of medium height;

in older individuals, it has strongly developed supraorbital

ridges (v), which form thin tlanges of bone overhanging

the orbital areas. They extend posteriorly as strong ridges

on the braincase. The zygomata (iv) are robust and broad-

er than the braincase but not as widely flared as in

Lenothrix ; the zygomatic plate is very narrow (xiii). The

incisive foramina (vi) are medium-sized and relatively

broad; their posterior margins (vii) end well in front of the

anterior margin of the first molars ( M 1 ) (viii). The poste-

rior border (x) of the palate ends before or in line with the

posterior border of the last upper molars (M5) (ix). Palatal

length slightly exceeds postpalatal length. The tympanic

bullae (xi) are very large. Each half mandible has a small

coronoid process (xiv); the lower incisor root is moder-

ately prominent (xv). Photographs of the skull are avail-

able in Musser & Newcomb ( 1983, Figure 94).

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Uraiporn Pimsai et al.

The enamel layers of the incisors are orange. The up-

per incisors are opisthodont (xii). In the first upper mo-

lar (M1), the first row comprises three separate cusps, tl

(Fig. 35Ba), t2 (b) and t3 (c); on the second row, cusps t4

(d), t5 (e) and t6 (f) are separate or variably fused, depend-

ing on the age of the individual; on the third row, cusp t7

(g) is separate from t8 (h) and t9 (i), which is small. In

the second upper molar (NT), the anterior lingual cusp (tl )

(j) is large and prominent; t3, if present, is small (not pres-

ent in the tooth illustrated in Fig. 35B ); on the second row,

cusps t4 (k), t5 (1) and t6 (m) are separate; on the third

row, cusp t7 (n) is separate from t8 (o), which is essen-

tially fused with a very small t9 (not present in the tooth

illustrated in Fig. 35B). In the third upper molar (M3); the

anterior lingual cusp (tl ) (p) is large and prominent; cusp

t3 is absent; cusp t4 ( q ) is separate from cusps t5 and t6

(r), which are fused; in the last row, cusp t7 is absent; cusps

t8 and t9 (s) are fused.

In the mandibular toothrow, the first lower molar (M,)

has an anterocentral (t), anterolabial (u) and anterolingual

cusp (v); the protoconid (w) is smaller than the metaconid

cusp (x); the hypoconid (y) is smaller than the entoconid

cusp (z); there is a posterior cingulum (aa). In the second

lower molar (M2), the anterolabial cusp is absent; the pro-

toconid (ab) and metaconid cusps (ac) and the hypoconid

(ad) and entoconid cusps (ae) are separate; there is a pos-

terior cingulum (af). In the third lower molar (M3), the an-

Fig. 35. Skull and dentition of Pithecheir parvus

(BM.73.8), Simpang Pertang, Negeri Sembilan,

peninsular Malaysia. (A): dorsal, ventral and lateral

view of skull and mandible (Scale = 5 mm); (B): oc-

clusal view of upper (left) and lower (right) left mo-

lars (Scale = 2 mm). Indicative numbers (i-xv) and

letters (a-ai) are explained in the text ‘Description’

for/' parvus. M: mouth; lab: labial.

terolabial cusp is absent; there is a protoconid (ag) and

metaconid cusp (ah); the posterior lamina comprises a

fused hypoconid and entoconid cusp (ai).

Karyology (based on specimens from Malaysia, Yong et

al. 1982). 2N= 50; 16 pairs of acrocentric autosomes, one

pair of subacrocentric autosomes and seven pairs of meta-

centric-submetacentric autosomes, the X chromosome is

a metacentric and the Y chromosome a subacrocentric.

Sperm morphology. Information on the sperm morphol-

ogy of a specimen from peninsular Malaysia is included

in Breed & Yong ( 1 986).

Fossil history. Specimens dating back to the early Pleis-

tocene have been collected from two localities in penin-

sular Thailand (Pearch et al. 2013).

Taxonomic notes. Measurements included by Kloss

( 1916b) for the type specimen are small (Table 19) and

probably represent a juvenile or very young adult.

Distribution and conservation status. Pithecheir parvus

is endemic to peninsular Malaysia (Fig. 36).

Its conservation status is listed by IUCN as ‘Data De-

ficient’ “since, although it has been recorded over a rea-

sonably wide area, it is uncertain if the species can per-

sist in modified habitats. If it can be found at disturbed

sites, it could be a Least Concern species, however, if it

is largely confined to primary or undisturbed forest, it may

qualify for a threat category” (Aplin et al. 2008h).

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63

Fig. 36. Distribution of Pithecheir parvus in the Myanmar-

Thai-Malaysian peninsula and Singapore. Malaysia: 1. Gunong

Benom; 2. Bukit Kutu; 3. Sungei Jengka Forest Reserve; 4. Sub-

ang Forest Reserve; 5. Meru Forest Reserv; 6. Bukit Perangoh;

7. Tanah Wat Forest; 8. Ayer Bangi; 9. Labis Forest Reserve. For

full locality details, see Gazetteer and Appendix I.

Ecology and reproduction. The holotype of P. parvus

was collected at an elevation of about 1050 metres (3400

feet) on Bukit Kutu in Selangor, Malaysia (Kloss 1916b).

Subsequently, it was collected from the same locality by

Muul & Lim (1971) and from other secondary and low-

land forests. The latter authors also obtained a specimen

from Gunong Benom, Pahang in primary forest at about

550 metres (1800 feet) elevation. According to Chaima-

nee (1998), P parvus lives in dense forests up to 1 600 me-

tres elevation but Aplin et al. (2008h) suggest that the up-

per limit is 1200 metres. With its prehensile tail and spe-

cialised hind feet, this is an arboreal species, which is

found in the forest understorey and in the crowns of short

palms and tree ferns. It constructs large globular nests

(Lim & Muul 1975; Musser & Newcomb 1983).

Genus Rattus

Rats

Rattus Fischer 1803: 128; type species Mus decumanus Pallax

(currently included in Rattus norvegicus).

Remarks. Rattus is the most diversified extant genus of

rodents with 64 species. Most species (59 of the 64) have

relatively restricted distributions, the exceptions are R. ar-

gentiventer, R. exulans, R. norvegicus, R. tanezumi and R.

tiomanicus , which together with R. andamanensis and R.

annandalei are the seven species currently known from

the Myanmar-Thai-Malasian peninsula (Musser & Car-

leton 2005). Synonyms of the extant species are includ-

ed in Musser et al. ( 1 979) and Musser & Carleton (2005).

Description (based on the seven species occurring in the

study area). The body size ranges from small ( exulans ) to

large ( norvegicus ) (Table 20). The tail is usually brown

above and below (Fig. 5 1C) but in R. norvegicus is slight-

ly bicoloured, paler below. Tail length averages less than

the head and body in some species, R. argentiventer and

R. norvegicus , is about equal to, or slightly exceeds it in

others, R. exulans, R. tanezumi and R. tiomanicus , and con-

siderably exceeds it in R. andamanensis and R. annan-

dalei. However, these are average measurements and there

is much individual variation and some geographical vari-

ation (see for example Tables 22 and 23). Some species,

R. andamanensi, R. argentiventer, R. exulans R. tanezu-

mi, and R. tiomanicus have pelage that includes spiny hairs

(to various degrees), whereas in others, R. norvegicus and

R. annandalei, they are absent. The dorsal pelage is grey-

brown, olive-brown, yellowish-brown/orange and variably

speckled with black. The ventral pelage is also variable

in colour, from white to yellowish-white to grey. The hind-

feet have six pads (Fig. 37C). Four are interdigital ( 1 ),

these are smooth in some species ( argentiventer ), whilst

in others they are ridged to assist with climbing (for ex-

ample tiomanicus and tanezumi ); the outer metarsal pad

(2) is smaller and the inner is kidney-shaped (3). The num-

ber of mammae varies between species; one has two ax-

illary pairs and two inguinal pairs (exulans), others have

three axillary pairs and three inguinal pairs; R. annandalei

is variable with four, five, or six pairs.

In the skull, the rostrum (Fig. 37Avi) is long and wide;

the nasals have rounded or blunt tips (i) and extend in front

of the incisors (xv), sometimes far enough to form a short

tube. The nasolacrimal capsules (ii) are large and inflat-

ed. The lachrymals (iii) are small. The braincase is medi-

um-sized. There are well-developed supraorbital ridges

(iv), which extend posteriorly along the lateral margins of

the braincase. The zygomatic arches (v) are robust and nar-

rower slightly anteriorly, such that they are broader where

they attach to the braincase. The zygomatic plate (xvi) is

naiTow. The palate (xiv) is long; the length of palate varies

between species; its posterior border (xi) extends beyond

the last upper molars (M3) (x). Therefore the distance from

the back edge of the palate to the ventral margin of the

foramen magnum (xiii) (postpalatal length) is shorter than

palate length. The incisive foramina (vii) are long and rel-

atively narrow; their posterior margins (viii) extend to, or

slightly pass, the anterior margins of the first molars (M1)

(ix). The tympanic bullae (xii) are relatively large. The

coronoid process (xvii) of the mandible and the emargina-

tion (xix) between the condylar (xviii) and angular (xx)

processes are well-developed. The process of the lower

incisor root (xxi) is moderately developed.

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Fig. 37. Skull and dentition of Rattus tanezumi

(54-4016 CTNCR), Kaeng Klioi District, Saraburi

Province, central Thailand. (A): dorsal, ventral and

lateral view of skull and mandible (Scale = 5 mm);

(B): occlusal view of upper (left) and lower (right)

left molars (Scale = 2 mm); (C): left hind foot of

R. tanezumi (PSUZC-MM20 12.204), Phato Dis-

trict, Chumphon Province, peninsular Thailand

(Scale = 5 mm). Indicative numbers (i-xxi) and

( 1-3) and letters (a-ae) are explained in the text

"Description’ for Rattus. M: mouth; lab: labial.

The enamel layers of the incisors are orange. The up-

per incisors are opisthodont (xv) or sometimes orthodont

relative to the rostrum (vi). In the first upper molar (M1),

cusp tl (Fig. 37Ba) is separate (in young individuals, see

Fig. 5E) from cusps t2 and t3 (b), which are fused; on the

second row, cusp t4 (c) is separate or fused with cusps t5

and t6 (d), which are fused; on the third row, there is a

large medial cusp, t8 (e), fused to a small lateral one, t9

(f); cusp t7 is missing. In the second upper molar (M2),

cusp tl (g) is large and prominent; most (but not all) spec-

imens have a small cusp t3 (h); in the second row, cusps

t4, t5 and t6 (i) are fused (in the youngest of individuals

with newly erupted teeth, t4 is separate from t5 and t6,

which are just fused); the third row comprises a larger t8

(j) and a smaller t9 (k), which are fused; t7 is absent. In

the third upper molar (M1), cusp tl (1) is large and promi-

nent; cusp t3 is usually absent in most species; cusps t4,

t5, t6 (m) are fused, even in young individuals; the last

row has only t8 (n); in worn teeth, sometimes cusps t4,

t5, t6, t8 are fused making a large part of the molar (as il-

lustrated in Fig. 37B).

In the mandibular toothrow, the first lower molar (M,)

is without an anterocentral cusp; in the oldest individu-

als, the anterolabial (o) and anterolingual cusps (p) and

the protoconid (q) and metaconid cusps (r) are all fused,

whereas in younger individuals they are all separate; in

the youngest individual of R. tanezumi seen, there is al-

so an anterior labial cusp (not illustrated in Figs 5M or

37B) and a posterior labial cusp (s); the hypoconid (t) is

fused with the entoconid cusp (u); the posterior cingulum

(v) is well-developed. In the second lower molar (M2), the

anterior labial cusp (w), the protoconid (x) and the meta-

conid cusp (y) are fused; a posterior labial cusplet is pres-

ent (see Fig. 5M) but may be fused with hypoconid (z)

(as in Fig. 37B), which is in turn fused with the entoconid

cusp (aa); the posterior cingulum (ab) is well-developed.

In the third lower molar (M3), the anterolabial cusp is ab-

sent; the protoconid (ac) is fused with the metaconid cusp

(ad); the posterior lamina comprises the entoconid (ae).

Key to species for the seven species of Rattus currently

known from peninsular Myanmar- Thailand- Malaysia and

Singapore is shown in Table 21. It should be noted that

differentiating between Rattus species, and especially R.

tanezumi and R. tiomanicus, is often difficult using mor-

phometric characters. This is further complicated by the

fact that the taxonomy still requires further refinement

with recent molecular studies showing that there are, for

example, at least four distinct lineages of Rattus in penin-

sular Thailand, the identification of which are not deter-

mined (Latinne et al. 2013b). If these lineages are true

species, then it is highly probable that R. tanezumi, and

possibly R. tiomanicus, are complexes of morphological-

ly cryptic species. As noted by Pages et al. (2010), fur-

ther research incorporating molecular data from the holo-

types of the different Rattus taxa, especially those of the

numerous synonyms, would be the most suitable way to

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65

determine accurately the taxonomy of this genus in South-

east Asia.

Ruttus andamanensis

Indochinese forest rat

Mus (leggada) andamanensis Blyth 1860: 103; Andaman Is-

lands.

Epimys remotus Robinson and Kloss 1914: 231; Samui Island,

East peninsular Thailand.

Description (based on Robinson & Kdoss 1914, Aplin et

al. 2003 = R. sikkimensis ): This is a moderately large, ar-

boreal rat (Table 20) with a mean mass of 221 g (Mar-

shall 1988). The pelage on the upper parts is somewhat

shaggy and includes thin, flexible, grooved, spiny-hairs.

The colour is an admixture of yellow-orange and sooty

brown, darkest on the back, where the long black guard

hairs are most numerous; the flanks are browner, with the

dark grey hair roots more prominent. The under parts are

creamy white, sharply demarcated from the flanks. The

hind feet are large and have silky, whitish hairs, darker

brown in the mid-part; they have large well-developed in-

terdigital and metatarsal pads, which are covered with fine

lamellae/ridges (to assist with climbing). The ears are large

for Rattus. The tail is long and unicoloured, brownish-

black; on average, it considerably exceeds head and body

length. The vibrissae are characteristically long (up to 60

mm in length), thick and black. There are six pairs of

mammae: one pair pectoral, two pairs postaxillary, three

pairs inguinal (Marshall 1988). The skull is strongly ridged

and broad; the sides of the braincase are slanted, not ver-

tical as in R. tanezumi (Corbet & Hill 1992). The rostrum

is robust and relatively short for its width. The incisive

foramina are long, extending to the anterior border of the

first (M1) molar or between the anterior roots of the M1.

Photographs of the skull are available in Musser & New-

comb 1983, Figures 110 and 111, = R. sikkimensis ) and

Marshall 1988, page 470, = R. remotus).

Karyology. 2N= 42 (based on specimens from Samui Is-

land, Thailand); autosomal pairs I, 4, 9, and 13 are sub-

telocentric; autosomal pair number 1 1 is heteromorphic;

the karyotype is illustrated in Markvong et al. ( 1973: Fig.

17).

Fossil history. Specimens referred to R. koratensis and

R. sikkimensis (see Taxonomic notes, below) and dating

back to the early Pleistocene are known from nine local-

ities in Thailand, including three in the peninsula

(Chaimanee 1998, Pearch et al. 2013).

Taxonomic notes. The taxonomy of this species is com-

plex. Specimens from Thailand have been variously re-

ferred to remotus by Marshall ( 1988) and Corbet & Hill

(1992), koratensis Kloss by Chaimanee ( 1998) and sikki-

mensis by Chaimanee (1998) and Musser & Newcomb

( 1983). Aplin et al. (2003) include both names. Howev-

er, following Musser & Carleton (2005), specimens from

the study area are here referred to R. andamanensis.

Fig. 38. Distribution of Rattus andamanensis in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: 1 . Ban Pa

Lau; 2. Koh Kra; 3. Koh Tau; 4. Koh Pha-ngan; 5. Koh Samui.

For full locality details, see Gazetteer and Appendix I.

Distribution and conservation status. Rattus andama-

nensis has an extensive range from eastern Nepal to north-

east India, Bhutan, southern China, Myanmar, northern

Thailand, and Indochina; it is also known from the An-

daman and Nicobar Islands. In the study area, it occurs

in one locality in northern peninsular Thailand and four

islands in the Gulf of Thailand (Fig. 38).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, presumed large pop-

ulation, occurrence in several protected areas, tolerance

of habitat modification, and because it is unlikely to be

declining fast enough to qualify for listing in a more threat-

ened category” (Aplin et al. 2008i).

Ecology and reproduction. In peninsular Thailand, the

holotype was collected in the hills of Koh Samui (Robin-

son & Kloss 1914), where it is also reported from ever-

green forest (Markvong et al. 1973). According to Mar-

shall (1988), it is found in orchards and secondary forest

in Koh Samui; it shares this habitat with Niviventer ful-

vescens and Rattus tanezumi (= Rattus bukit and Rattus

rattus robinsoni respectively in Marshall 1 988). It is con-

sidered to be arboreal (Aplin et al. 2003).

Rattus annandalei

Annandale’s rat, Singapore rat

Mus annandalei Bonhote 1903a: 30; Sungkai, South Perak,

Malaysia.

Mus villosus Kloss 1908a: 146; Singapore Island.

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Uraipom Pimsai et al.

Description (based on Bonhote 1903a; Kloss 1908a). This

is a medium-large rat (Table 20) with a mass of 145-250

g (Medway 1969), 120-260 g (Muul & Lim 1971). The

fur is moderately soft and without spines. The upper body

is grizzled yellowish-brown (fulvous) to buff-brown. It is

darker posteriorly owing to the prevalence of long black

guard hairs, which attain 40 mm on the rump. It is also

darker down the middle of the back. The flanks are paler,

owing to the grey basal hair showing through and the ab-

sence of dark annulations to the hairs. In some individu-

als there is a narrow band (4 mm) of pure buff on the

flanks. The under parts are white, tinged with yellow; the

line of demarcation on the flanks is well marked in some

individuals, less so in others. The buffy white of the chest

and belly extends to the wrists and to the upper lip behind

and below the whiskers. In males, there is a brown pre-

scrotal patch. The forearms and thighs are greyish-brown;

the inner sides of the fore limbs are white and of the hind

limbs, including the ankles, are dark brown. The feet are

dark brown. In general, the tail is considerably longer than

the head and body and is a uniform black in colour, cov-

ered in short stiff hairs. The ears are somewhat elongat-

ed and naked. There are two or three pairs of pectoral and

two or three pairs of inguinal mammae (Medway 1969).

The skull is elongated and has noticeably large tympan-

ic bullae. The incisive foramina tend to be short and ex-

tend to the anterior border of the first (M1) molar. Photo-

graphs of the skull are included in Musser & Newcomb

(1983, Figures 110 and 111).

Fossil history. There are no records of fossil R. annan-

dalei from Thailand (Pearch et al. 2013).

Taxonomic notes. According to Medway ( 1969), speci-

mens from Perak are referable to R. a. anncindalei where-

as those from Selangor and to the south are comparable

to R. a. bullatus Lyon 1908, which was described from

Rupat Island, east of Sumatra.

Distribution and conservation status. Rattus anncindalei

has a relatively restricted range; it is found in eastern

Sumatra; Padang and Rupat Islands off the east coast of

Sumatra; peninsular Malaysia and Singapore. Its distribu-

tion in the study area is illustrated in Fig. 39.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “as although it is probably undergoing decline due

to the conversion of forested habitat to agricultural lands

in parts of its range, it is unlikely to be declining fast

enough to warrant listing in a higher category of threat”

(Aplin & Lunde 2008c).

Ecology and reproduction. In Perak in peninsular

Malaysia, it was apparently uncommon (Chasen 1940).

Medway (1969) suggested that it was localised, confined

to secondary woodland and scrub. However, in Selangor,

it was common in lowland secondary forests and in kam-

pong rubber estates; here it was trapped in the lower

branches of trees and also on the ground. None was tak-

en in deep forest (Muul & Lim 1971). In Singapore, it was

Fig. 39. Distribution of Rattus annandalei in the Myanmar-

Thai-Malaysian peninsula and Singapore. Malaysia: 1. Taip-

ing; 2. Bruas; 3. Tanjong Tuan (= Tanjong Hantu); 4. Sungkai;

5. Tanjong Malim; 6. Kuala Selangor; 7. Bukit Cherakah; 8. Buk-

it Lagong; 9. Bukit Lanjang; 10. Meru; 11. Subang Forest Re-

serve; 12. Cheras; 13. Kuala Lumpur; 14. Klang; 15. Bukit Man-

dol. Singapore: 16. Botanical Gardens. For full locality de-

tails, see Gazetteer and Appendix I.

widespread (Harrison 1966). According to Corbet & Hill

( 1992), it is not an agricultural pest.

Rattus argentiventer

Ricefield rat

Epimys rattus argentiventer Robinson & Kloss 1916b: 274; Pasir

Ganting, West coast of Sumatra, 2°7’S.

R. r. chaseni Sody 1941: 269; Krian, Perak, Malaya.

Description (based on Musser 1973b, Aplin et al. 2003).

This is a medium-sized rat with a robust body and a rel-

atively short tail, which usually averages shorter than head

and body length (Table 20). The pelage feels soft and sleek

to the touch; it includes thin, pale, flexible, grooved, spiny-

hairs. The guard hairs are black and relatively short and

inconspicuous. The upper parts are yellowish brown

speckled with black, giving a characteristic ‘salt and pep-

per’ effect. Although some specimens are more greyish

and others have more yellowish and brown hues, the ‘salt

and pepper’ effect is always distinctive. The underparts

are silvery grey (never creamy), with white hair tips and

grey bases; the midventral region is often a buffy-brown;

the throat is white. The flanks and sides of the head are

paler than the head and back; this colour extends to the

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67

upper surface of the fore and hind feet. The hind feet are

long and narrow and have six pads, four interdigital and

two metatarsal; these pads are small and smooth relative

to those in R. tiomanicus and R. tanezumi , with the lamel-

lae/grooves scarcely indicated (hind foot illustrated in

Musser 1973b, Figure 2). The cheeks are grey. In front of

each ear is a tuft of orange hairs. These tufts are clearly

visible in young individuals; they may be less conspicu-

ous in some adults that have longer pelage (Guy Musser

pers. comm.). The tail is unicoloured, dark above and be-

low; it is usually but not always shorter than the head and

body. Females have six pairs of mammae: one pectoral,

two postaxillary, one abdominal, and two inguinal. The

skull has a short, broad and deep rostrum, which curves

conspicuously downwards. The interorbital region is nar-

row; the braincase is oval. The incisive foramina are long

and wide; posteriorly they terminate between the first up-

per molars (M1), a point much further back than in R.

tanezumi. The tympanic bullae are relatively large. The

dentition is robust. Photographs of the skull are includ-

ed in Musser & Newcomb ( 1983, Figures 1 10 and 111)

and Marshall ( 1988, page 468).

Karyology. 2N= 42 (based on specimens collected in rice

fields in the Bangkok area); autosomal pairs 1 to 9 are sub-

telocentric; the karyotype is illustrated in Markvong et al.

(1973, Figure 16).

Sperm morphology. Information on the sperm morphol-

ogy of two specimens from peninsular Malaysia is includ-

ed in Breed & Yong (1986).

Fossil history. Rattus argentiventer is known from a sin-

gle late middle Pleistocene site in north-eastern Thailand

(Pearch et al. 2013).

Taxonomic notes. The taxon argentiventer was described

originally as a subspecies of Rattus rattus (= R. tanezu-

mi). However, this view was rejected by Musser ( 1973b)

and was not followed by subsequent authors, including

Corbet & Hill (1992), Musser & Carleton (2005) and

Maryanto (2003). The taxon chaseni is included as a syn-

onym (Maryanto 2003), although the tail is uncharacter-

istically long (Table 20).

Distribution and conservation status. Rattus argentiven-

ter has an extensive range from Thailand and Indochina

to Malaysia, Indonesia, and the Philippines. Within the

study area, it is known from peninsular Myanmar, Thai-

land and Malaysia (Fig. 40).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, adaptability to dis-

turbance, large population, its occurrence in a number of

protected areas, the absence of threats, and because the

population is considered to be stable” (Ruedas et al.

2008a).

Ecology and reproduction. In Thailand, R. argentiven-

ter was collected in secondary evergreen forest in Surat

Thani Province at approximately 400 m. a.s.l. (UP unpub-

lished data). In Malaysia, it is restricted to ricefields, scrub.

Fig. 40. Distribution of Rattus argentiventer in the Myanmar-

Thai-Malaysian peninsula and Singapore. Myanmar: l.Tavoy

Island; 2. King’s Island; 3. Ross Island; 4. Tenasserim Town; 5.

Maliwun; 6. Victoria Point. Thailand: 7. Ban Kam Phuan; 8.

Khao Phlu (base of), Koh Samui; 9. Rajjaprabha Dam; 10. Klong

Phraya Wildlife Sanctuary; 11. Bangnara; 12. Rangae.

Malaysia: 13. Kedah Peak; 14. Kampong Pulau Betong; 15.

Bagan Triang; 16. Parit Buntar; 17. Tanjong Piandang; 18. Bri-

ah; 19. Krian; 20. Kuala Kurau; 21. Telok Anson; 22. Bagan

Serai; 23. Gunong Semanggul; 24. Maxwell’s Hill; 25. Selens-

ing; 26. Taiping; 27. Bagan Datoh; 28. Pahang (State); 29. Sungei

Tua; 30. Kuala Lumpur; 31. Cheras. For full locality details,

see Gazetteer and Appendix I.

grassland and young plantations; it is absent from the for-

est interior (Medway 1969). Aplin et al. (2003) suggest

that it avoids village habitats except as a vagrant but is

found in gardens and orchards; it favours areas that are

regularly flooded. Both sexes dig burrows in well drained

soil, in bunds between flooded paddies, or in and around

raised vegetable gardens and orchards. The burrows of fe-

males tend to be more extensive than those of males. One

burrow can house two or more litters from the same fe-

male (Medway 1969, Aplin et al. 2003). Its diet includes

a high proportion of insects, including termites in grass-

land and grasshoppers in rice fields; it also feed on young

rice plants, grasses, paddy weeds, grass seeds and cereal

grains. It may feed on some vertebrates and snails. It is a

pest of young oil palms (Harrison 1 954a, 1962, Lim 1966,

Medway 1969). Typically crop losses in rice growing ar-

eas to R. argentiventer may be 10-20 % but chronic loss-

es of 30-50 % may occur in areas that are particularly

favourable to this rodent or where the multiple cropping

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Uraipom Pimsai et al.

of rice provides a continuous supply of food. On average

10% of females are pregnant without seasonal variation.

Mean litter size is 6.0 and ranges from 5 to 7 (Harrison

1961, Medway 1969). However Aplin et al. (2003) sug-

gest that breeding patterns are closely linked to rice pro-

duction, with the first litter as the rice reaches booting

stage, the second litter during the ripening stage, and the

third litter shortly after harvest. At certain times nearly 1 00

% of females may be pregnant. Females usually become

pregnant at a point when body weight reaches 60-120 g.

Males usually reach sexual maturity at 90+ g (estimated

at about 59 days). The gestation period is 20-26 days. In

the wild, on average females give birth to about 6 infants.

Rattus exulans

Polynesian rat

Rattus exulans Peale 1848: 47; Tahiti, Pacific Ocean.

Mus obscurus Miller 1900a: 213; Tioman Island, Malaysia. Pre-

occupied by M. obscurus Waterhouse 1837 (= Akodon obscu-

rus) = M. pul /us Miller 1901: 178.

Description (based on personal observation. Miller

1900a, Medway 1969, Corbet & Hill 1992, Aplin et al.

2003). This is a small rat, which is superficially similar

to a large mouse (Table 20). The fur is rather short, about

8 mm in the middle of the back, and there are numerous

well-developed, slender, grooved, pale, hair-like spines;

these bristles are best developed on the upper surface,

whilst on the ventral surface they are scarcely evident. The

dorsal surface is grey-brown to brown intermixed with dull

yellowish-red buff; the head and lower back tend to be

browner than the rump, flanks, shoulders and legs. The

ventral surface of the body and the inner side of the legs

are pale, irregularly washed with brownish yellow. The

bases to the hairs are slate-grey throughout the body. The

feet are grey. The tail is uniformly dark brown. It is an-

nulated with about 16 rings/cm in the midpart of the tail;

these rings are obscurely divided into scales. The tail has

very short hairs, which do not obscure the rings nor form

a pencil (tuft) of hairs at the tip. The hind feet have six

well-developed pads; the proportionately larger interdig-

ital and metarsal pads clearly distinguish this species from

a large Mus (Fig. 28C). The whiskers (vibrissae) on the

head are long. According to Medway ( 1969), there are four

pairs of mammae; two pairs pectoral and two pairs in-

guinal; Aplin et al. (2003) state that there are one pair pec-

toral, one pair postaxillary and two pairs inguinal. The

skull is the smallest of all the Rattus in the study region

(Table 20). Unlike Mus (Fig. 28B), the length of the an-

terior upper molar (M1) in R. exulans is less than the com-

bined lengths of the second and third molars (M: and M').

Photographs of the skull are included in Musser & New-

comb ( 1983, Figures 110 and 1 1 1 ) and Marshall ( 1988,

page 474).

Karyology. 2N= 42 (based on specimens collected in Chi-

ang Mai, northern Thailand); autosomal pairs 1 to 11 are

subtelocentric; the karyotype is illustrated in Markvong

et al. (1973, Figure 19). According to Badenhorst et al.

(2009), the diploid number is 2N= 42 and and the auto-

somal fundamental number (NFa) = 54, 58 or 59.

Sperm morphology. Information on the sperm morphol-

ogy of two specimens from peninsular Malaysia is includ-

ed in Breed & Yong (1986).

Fossil history. There are no recorded fossil specimens of

R. exulans from Thailand (Pearch et al. 2013).

Taxonomic notes. According to Corbet & Hill (1992), ge-

ographical variation in this widely distributed species is

not significant, mainly affecting the ventral pelage,

which ranges from light to dark grey with a variable

amount of brown in the hair tips. Extralimitally, montane

forms tend to be slightly darker with longer, softer pelage

and flattened spines absent or sparse. In the study of Pages

et al. (2010), R. exulans formed a discrete monophyletic

group.

Distribution and conservation status. Rattus exulans has

an extensive range from Bangladesh through Southeast

Asia to Taiwan, Indonesia, the Philippines, New Guinea,

the Southwest Pacific islands, and islands off northern

Australia. Its distribution is probably caused by a combi-

nation of inadvertent human introductions and possibly

some natural rafting (Musser & Carleton 2005). Within

the study area, it is known from peninsular Thailand and

Malaysia and Singapore (Fig. 41).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, tolerance of a broad

range of habitats, presumed large population, and because

it is unlikely to be declining fast enough to qualify for list-

ing in a more threatened category” (Ruedas et al. 2008c).

Ecology and reproduction. Rattus exulans is variably de-

scribed as being a ‘ground rat’ (Medway 1969) and as

‘highly arboreal' (Aplin et al. 2003). According to Cor-

bet & Hill ( 1992), it is a common pest species found in

houses, granaries, cultivated areas, scrub and forest, usu-

ally on the ground. In Thailand, it is reported from hous-

es, markets, the vicinity of houses and also rice paddies

close to buildings (Marshall 1988). In field surveys for the

current study, it was found in a forest near buildings and

in secondary evergreen forest in Surat Thani Province (ap-

proximately 400 metres, a.s.l.). Specimens were also col-

lected in rain forest in Narathiwat Province (approximate-

ly 200 metres a.s.l.) and in a building in Songkhla Province

(UP unpublished data). Extralimitally, in mainland South-

east Asia and Bangladesh, it has been found to coexist with

R. tanezumi in village houses. It climbs around in tall

grasses or low trees and on the walls and roofs of hous-

es. It usually constructs a nest of leaves and grass in dense

grass and situated some 200 mm or more above the

ground. Inside buildings, nests are usually located in roof

thatch and less often in piles of straw or other material on

the ground (Aplin et al. 2003). In Malaysia, 23-29 % of

females were pregnant, with little seasonal variation. Mean

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

69

Fig. 41. Distribution of Rattus exulans in the Myanmar-Thai-

Malaysian peninsula and Singapore. Thailand: 1. Koh Tao; 2.

Ban Tai; 3. Ban Bang Khla, Kho Samui; 4. Rajjaprabha Dam;

5. Khao Lak; 6. Koh Yao Yai; 7. Prince of Songkla University;

8. Muang Yala; 9. Ban Thon; 10. Biserat; 11. Hala Bala Wildlife

Research Station. Malaysia: 12. Langkawi Island; 13. Penang

Island; 14. Maxwell’s Hill: 15. Bukit Jong; 16. Aring; 17. Tan-

jong Hantu; 18. Pahang (State); 19. Semangko Pass; 20. Klang

Gates; 21. Kuala Lumpur; 22. Ulu Langat; 23. Tulai Island; 24.

Tioman Island; 25. Malacca; 26. Segamat; 27. Aur Island; 28.

Tinggi Island; 29. Tcbraun; 30. Pelepak; 31. Johore Bahru; 32.

Kangka; 33. Pasir Gudang. Not located: Gomok (Johore State).

Singapore: 34. Tanglin. For full locality details, see Gazetteer

and Appendix I.

litter size is 4.3 (1 to 8) (Harrison 1955, Medway 1969).

Mean length of life in the wild is thought to be 3.2 months

(Harrison 1956a).

Rattus norvegicus

Norway rat

Mus norvegicus Berkenhout 1769: 5; Great Britain.

Description (based on personal observation, Medway

1969, Marshall 1988, Corbet & Hill 1992, Aplin et al.

2003, Francis 2008). This is a large rat (Table 20) with a

mass of between 150-400 g (Medway 1969). It has short

fur, which is slightly stiff but is without bristles; it has long

guard hairs. It is grey-brown to brown on the dorsal sur-

face; pale-brown or grey on the ventral surface; there is

often a white patch on the chest. There is no clear line of

demarcation between the colour on the flanks and the bel-

ly. Black (melanistic) individuals are quite common in

some areas. The nose is long and broad and the eyes and

ears are relatively small. The tail is almost always short-

er than the head and body. It is usually weakly bicoloured,

dark above and slightly paler below, sometimes mottled

in appearance. The front and hind feet are large, with poor-

ly developed foot pads and inconspicuous ridges. In con-

trast to the dark feet of Bandicota, those of R. norvegicus

are mostly white (dark hairs in melanistic individuals).

There are six pairs of mammae with three pairs pectoral

and three pairs inguinal. The skull is robust, medium-large

to very large in some individuals with long incisive foram-

ina. The skull ridges are well-defined in older individu-

als and characteristic in shape; they are relatively straight

and situated close to one another, which gives the impres-

sion of a relatively narrow braincase. The toothrows are

long. Photographs of the skull are included in Marshall

( 1988, page 462) and Musser& Newcomb (1983, Figures

110 and 111).

Karvology. 2N= 42, FN= 62 (based on specimens from

Thailand); seven pairs of metacentric chromosomes; four

pairs of subtelocentrics; nine pairs of telocentrics; the X

and Y sex chromosomes are telocentrics (Musser & New-

comb 1983).

Fossil history. There are no recorded fossil specimens of

R. norvegicus from Thailand (Pearch et al. 2013).

Taxonomic notes. Wild populations of this species are

thought to have inhabited Southeast Siberia, northern Chi-

na and parts of Japan. However, inadvertent human intro-

ductions worldwide have obscured wild geographical pat-

terns.

Distribution and conservation status. Rattus norvegicus

has a worldwide distribution but is more common in the

colder latitudes of the Northern and Southern Hemi-

spheres. In the wanner climates of the tropics it is restrict-

ed to habitats that have been highly modified by humans

(Musser & Carleton 2005). Within the study area, it is

recorded from Thailand and Malaysia and Singapore (Fig.

42) but is probably considerably more widespread in ur-

ban areas than this map suggests.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ as it is “a common species with no major threats”

(Ruedas 2008).

Ecology and reproduction. According to Medway

(1969), R. nor\>egicus is confined to ships, harbours and

neighbouring built-up areas. It is apparently unable to

compete with native rats in rural habitats. Aplin et al.

(2003) describe it as a terrestrial, burrowing species with

poor climbing skills, which is often found close to water,

such as along rivers and major irrigation canals. It occurs

in urban areas, where it lives in and around buildings and

animal yards, feeding on refuse and stored food. Occasion-

ally, it is found in cultivated areas but the level of dam-

age to crops is not known in Southeast Asia. It is a com-

munal species that constructs large and complex burrow

systems that may be occupied for many years. Its nests

are lined with leaves and other soft material and are con-

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Uraipom Pimsai et al.

Fig. 42. Distribution of Rattus norvegicus in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: 1. Koh

Samui. Malaysia: 2. Kota Bharu; 3. Gurun; 4. Georgetown,

Penang Island; 5. Gunong Ijau; 6. Kuala Lumpur; 7. Johore

Bahru. Singapore: 8. Singapore Island. For full locality de-

tails, see Gazetteer and Appendix 1.

structed within the burrow. No data are available on lit-

ter size in Southeast Asia. In India, mean litter size is 8. 1 .

The gestation period is 22 to 24 days.

Rattus tanezumi

Oriental roof rat. Oriental house rat. Oriental ship rat

Rattus tanezumi Temminck 1 844: 5 1 ; Japan (possibly from near

Nagasaki on Kyushu Island).

Mus griseiventer Bonhote 1903a: 30; Perak, Malaysia.

Mus pannosus Miller 1900c: 190; Adang Island, Butang Islands,

Thailand.

Epimys panellus Miller 1913: 8; Rawi Island, Butang Islands,

Thailand.

Epimys rattus dentatus Miller 1913: 14; Hastings Island, Mer-

gui Islands, Myanmar.

Epimys rattus insulanus Miller 1913: 14; Heifer Island, Mergui

Islands, Myanmar.

Epimys rattus exsul Miller 1913: 15; James Island, Mergui Is-

lands, Myanmar.

Epimys rattus fortunatus Miller 1913: 15, Chance Island, Mer-

gui Islands, Myanmar.

Rattus rattus tikos Hinton 1919: 400; Tenasserim Town, Myan-

mar.

Rattus rattus panjius Chasen 1937: 85; N Panjang Island, west-

ern peninsular Thailand.

Rattus rattus alangensis Chasen 1937: 87; Alang Yai Island,

western peninsular Thailand.

Rattus rattus lontaris Chasen 1937: 88; Lontar Island, western

peninsular Thailand.

Bonn zoological Bulletin 63 (1): 15-114

Rattus rattus kadanus Chasen 1937: 89; Kadan Island, western

peninsular Thailand.

Rattus rattus moheius Chasen 1937: 91; Pipidon Island, west-

ern peninsular Thailand.

Rattus rattus pipidonis Chasen 1937: 92; Pipidon Island, west-

ern peninsular Thailand.

Rattus rattus robinsoni Chasen 1940: 154; Samui Island, east-

ern peninsular Thailand.

Description (based on personal observation and Bonhote

1903a, Chasen 1937, Medway & Lim 1966, Medway

1969, Musser & Califia 1982, Marshall 1988, Corbet &

Hill 1992, Aplin et al. 2003, Francis 2008). This is a medi-

um-large rat (Tables 20 and 22, Fig. 5 1C) with a mass of

85-240 g (Medway 1969). The dorsal pelage includes thin,

flexible, grooved, spine-like hairs, these are pale basally

with black tips and are more evident in older individuals.

The extent of the spines varies between populations - for

example, they are scarcely evident in the island form lon-

taris. There are long guard hairs that extend well beyond

the overhairs and are conspicuous over the lower part of

the back and rump; these are mostly black, although some

are also paler for part of their length. In general, the up-

perparts are grizzled olive brown, sometimes with grey-

ish or reddish-orange tones, sometimes there is darken-

ing on the midline of the back and the crown of the head;

some individuals are paler, others are darker. The under-

parts vary in colour from grey to brown-grey to creamy-

white, sometimes with a yellowish tinge. Medway & Lim

( 1966) suggest that the colour of the ventral pelage may

be related to habit, with those living in urban areas in

mainland Malaysia having greyish-brown hairs on the bel-

ly and those on Singapore Island, which occupy rural as

well as urban areas having variably coloured belly hairs

from greyish-brown to pale grey. The feet are brown, al-

though the hairs on the toes are white. The hind feet are

moderately broad with prominent plantar (interdigital and

metatarsal) pads (but not as prominent as in R. andama-

ttensis) and usually with obvious ridges/lamellae (in con-

trast to R. argentiventer, the pads of which are character-

istically smooth). The ears are thinly furred, rounded and

relatively large. The tail is shorter, equal to or rather longer

than head and body (Table 22); it is uniformly dark/black

throughout, covered with numerous short stiff black hairs;

very occasionally there is a short, white tip. There are five

or six pairs of mammae; one pectoral pair, one or two

postaxillary pairs and three inguinal pairs. The skull is

medium-sized. Medway & Lim (1966) and Musser &

Califia (1982) suggest that R. tanezumi (= R. r. diardii)

has longer incisive formina, longer maxillary toothrows

and more strongly developed and more angular supraor-

bital ridges in comparison to R. tiomanicus (= tiomani-

cus and jalorensis in Medway & Lim 1966). This latter

character is illustrated in Fig. 43. Additional photographs

of the skull are included in Musser & Califia (1982, Fig-

ure 2), Marshall (1988, page 477) and Musser & New-

comb ( 1983, Figure 110 = /?. rattus from Thailand).

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

71

m

Fig. 43. Crania of adult Rattus: (A): R. tanezumi , Sabah (US-

NM 292689, male); (B): R. tiomanicus, Sabah (USNM 292672,

male) modified from Musser & Califia ( 1982, Figure 2). i: the

angular supraorbital ridge of R. tanezumi ; ii: the smooth, curved

supraorbital ridge of R. tiomanicus.

Karyology (based on specimens from Thailand, Musser

and Newcomb 1983 = R. rattus). 2N= 42, FN= 54; there

are seven pairs of metacentric chromosomes; 1 3 pairs of

telocentrics; the X and Y sex chromosomes are telocentrics

(Musser & Newcomb 1983).

Fossil history. There are four localities in Thailand includ-

ing two in the peninsular, from which fossil specimens dat-

ing back to the middle Pleistocene have been recovered

(Pearch et al. 2013). These were referred to R. rattus by

Chaimanee (1998) but are included here in R. tanezumi.

Taxonomic notes. Previously specimens here referred to

tanezumi from Southeast Asia were included in R. rattus

(see Corbet & Hill 1992) with those from Singapore,

Malaysia and peninsular Thailand frequently referred to

R. rattus diardii Jentink, which was described from west-

ern Java (see Musser & Califia 1982). However, Baver-

stock et al. (1983) showed differences in the karyology

between true R. rattus (2N= 38/40) and the Southeast

Asian form, R. tanezumi (2N= 42) and Musser & Carleton

(2005) treated the two taxa as distinct species and includ-

ed all ‘ R . rattus ’from Southeast Asia in R. tanezumi. Sub-

sequently, Aplin et al. (2011) have cautioned that this ‘bi-

partite separation’ over-simplifies a more complex evo-

lutionary history, the details of which have yet to be re-

solved. Meanwhile, Latinne et al. (2013b) suggest that the

taxon tanezumi may itself include a complex of discrete

genetic lingeages and possibly species.

Distribution and conservation status. Rattus tanezumi

has a range that extends from Afghanistan through north

India, Nepal, Bhutan to southern China, and Korea to Tai-

wan and Japan (although it may be introduced to the lat-

ter two countries) (Musser & Carleton 2005). Within the

study area, it is recorded from Myanmar, Thailand and

Malaysia and Singapore (Fig. 44).

Fig. 44. Distribution of Rattus tanezumi in the Myanmar-Thai-

Malaysian peninsula and Singapore. Myanmar: l.NatheMine;

2. King’s Island; 3. Tagoot; 4. Thaget; 5. Banlaw; 6. Tenasser-

im Town; 7. Ross Island; 8. Griddles Island; 9. Sir John Hayes

Island; 10. Malcolm Island; 1 1 . Heifer Island (exact location not

determined); 12. Sullivan Island (Lampi Island); 13. James Is-

land; 14. Loughborough Island; 15. Maliwun; 16. Bankachon;

17. Victoria Point; 18. Hastings Island; 1 9. Victoria Island. Thai-

land: 20. Kaeng Kra Chan National Park; 2 1 . Pa La U; 22. Bang

Saphan; 23. Pak Chan; 24. Ban Bang Bane; 25. Ban Bang Non;

26. Koh Tao; 27. Pa Toh Watershed Management; 28. De Lisle

Island; 29. Ban Tai; 30. Ban Kam Phuan; 31. Koh Surin Nua;

32. Chance Island; 33. Koh Yam Yai; 34. Phato Watershed Con-

servation and Management Unit; 35. Koh Rah; 36. Koh Prah

Tung; 37. Koh Samui; 38. Ban Bang Khla, Koh Sarnui; 39. Khao

Plilu (base of), Koh Samui; 40. Ao Ko; 41. Khao Sai; 42. Ban

Plai Num; 43. Khao Lak; 44. Klong Phraya Wildlife Sanctuary;

45. Koh Boi Yah; 46. North Panjang Island (Koh Yao Noi); 47.

Koh Naka Yai; 48. Koh Maprau; 49. Koh Alan Yai; 50. Panjang

Island (Koh Yao Yai); 51. Pipidon Island; 52. BanTha-Phae; 53.

Banna; 54. Tham Nam; 55. Khao Phu Khao Ya National Park;

56. Ban Khuan Dang; 57. Pak Jam; 58. Khuan Khi Sian; 59.

Trang; 60. Lontar Island; 61. Pulau Muntia (Koh Muk); 62.

Kadan Island; 63. Pulau Mohea; 64. Bulon Island; 65. Tarutao

Island; 66. Koh Rawi; 67. Adang Island; 68. Wang Bla Chan; 69.

Khao Rup Chang; 70. Kuan Khao Wang Forest Park; 71 . Prince

of Songkla University; 72. Pak Num; 73. Ban Nai Tan Yong; 74.

Nam Tok Sai Khao; 75. Ban Lam Mai; 76. Sam Yak A-Sen; 77.

Ban Chok; 78. Ban Thon; 79. Ban Ya Kan; 80. Pa Phm; 81 . Ha-

la Bala Wildlife Research Station; 82. Biserat. Malaysia: 83.

Pelarit; 84. Penang (Island); 85. Krian Road; 86. Kledang Hill;

87. Larut Hills; 88. Taiping; 89. Perak (State); 90. Aring; 91.

Bagan Datoh; 92. Semangko Pass; 93. Tanjong Karang; 94.

Kuala Selangor; 95. Rawang; 96. Gombak Forest Reserve; 97.

Pulau Angsa; 98. Klang Gates; 99. Arnpang; 100. Kuala Lumpur;

101. Pelepak; 102. Pulai; 103. Johore Bahru; 104. Kangka Seni-

bong; 105. Pasir Gudang; 106. Tanjong Surat. Singapore: 107.

Singapore Island. For full locality details, see Gazetteer and Ap-

pendix I.

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Uraiporn Pimsai et al.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “due to its wide distribution and tolerance of a wide

variety of habitat types” (Heaney & Molur 2008).

Ecology and reproduction. In Thailand. R. tanezumi is

found in a wide-range of habitats including buildings and

human habitation, where it is common. However, it has

been collected in primary forest in Phatthalung and Phang

Nga Provinces, tropical rain forest in Naratiwat Province,

evergreen forest in Songkhla and Chumphon Provinces,

and secondary forest in Trang, Surat Thani and Phetch-

aburi Provinces. In Tarutao National Park, Satun Province,

it was found in primary forest on the islands of Tarutao,

Adang, and Rawi, and in secondary forest on Bulon Is-

land (UP unpublished data). In mainland Malaysia, R.

tanezumi is generally associated with houses, outhouses

and stores in urban areas, where it is the dominant species

(Medway 1969, = R. rattus) and can be a significant pest,

responsible for major post-harvest losses ( Aplin et al.

2003, = R. rattus Complex). However in Singapore, it oc-

cupies not only houses but also field and fringe habitats

(Medway & Lim 1966, = diardii ) and in Lao PDR it is

known from field and forest habitats. Where it also oc-

curs in rural habitats, R. tanezumi may cause extensive

damage to cereal, vegetable and fruit crops, including co-

conuts (Aplin et al. 2003). Its diet also includes refuse,

slugs and snails (Medway 1969). It builds loose spheri-

cal nests of shredded vegetation, cloth or other material.

These are placed in a confined space, including in a bur-

row, in roof thatch, in a wall cavity or inside a mud-brick

wall, in a pile of cut wood, in a straw pile, among stored

sacks of grain, among rocks, in a tree hollow or a fallen

log or other natural sites, such as the fork of a tree (Aplin

et al. 2003). In Kuala Lumpur, pregnant females were col-

lected in all months, without significant seasonal varia-

tion (Medway 1969). However, in rural areas pregnancy

is synchronised with cycles of crop maturation (Aplin et

al. 2003). Average gestation period is 21 days and mean

litter size is 5.7 (one to 11). Young reach sexual maturi-

ty at about 80 days (Medway 1969).

Rattus tiomanicus

Malaysian wood rat, Malaysian field rat

Mus tiomanicus Miller 1900a: 209; Tioman Island, Malaysia.

Mus jalorensis Bonhote 1903a: 29; Nong Chik, Pattani, penin-

sular Thailand.

Mus jarak Bonhote 1905: 69; Jarak Island, Straits of Malacca.

Mus rattus rumpia Robinson & Kloss 191 la: 169; Rembia Island,

Sembilan Islands, west of Malaysia.

Epimys tingius Miller 1913: 9; Tinggi Island, eastern Malaysia.

Epimys roa Miller 1913: 10; Maratua Island, eastern Malaysia.

Epimys rattus viclana Miller 1913: 13; Langkawi Island, west-

ern Malaysia.

Rattus rattus payanus Chasen & Kloss 1931: 79: Paya Island,

Straits of Malacca, Malaysia.

Rattus rattus perhentianus Chasen 1940: 155; East Perhentian

Island, eastern Malaysia.

Rattus rattus pemanggis Chasen 1940: 156; Pemanggil Island,

eastern Malaysia.

Bonn zoological Bulletin 63 ( I ): 15 1 14

Rattus rattus pharus Hill 1960: 75; Pisang Island, Straits of

Malacca, Malaysia.

Rattus rattus sribuatensis Hill 1960: 76; Sribuat Island, Pahang,

Malaysia.

Rattus rattus kabanicus Hill 1960: 77; Kaban Island, Johore Is-

lands, Malaysia.

Rattus rattus terutavensis Hill, 1960: 79; Telok Udang, Terutau

Island, Southwest Thailand.

Rattus tiomanicus tenggolensis Yong 1971 : 89; Tenggol Island,

off eastern Malaysia.

Description (based on personal observation and Miller

1900a, Bonhote 1903a, Chasen 1940, Medway & Lim

1966, Medway 1969, Musser & Califia 1982, Marshall

1 988, Corbet & Hill 1 992, Aplin et al. 2003, Francis 2008).

This is a medium-large rat (Tables 20 and 23 ) with a mass

of 55-152 g (Medway 1969). The dorsal pelage is sleek,

moderately long and soft; the proportion of slender spine-

like hairs seems to vary geographically and between in-

dividuals; the guard hairs are black, short and inconspic-

uous, barely extending beyond the overhairs. The gener-

al colour is a warm yellowish-brown, grizzled with black-

ish-brown. The flanks and cheeks are slightly paler than

the back and much less profusely sprinkled with dark

hairs. Each hair is ashy-grey at the base with a broad,

brownish tip. The spine-like hairs are whitish with a black

tip. The underparts tend to be white in mainland popula-

tions in Malaysia but are variably coloured in island pop-

ulations, including white suffused with grey to buffy grey

(Musser & Califia 1982). Throughout the ventral pelage,

except on the chin and throat, the hairs are a light slate-

grey at the base. The line of demarcation between the up-

per and lower surfaces is usually well defined. The tail is

unicoloured, dark above and below. The plantar (interdig-

ital and metatarsal) pads on the hind feet are less well-de-

veloped than those of R. andamanensis and are more

ridged than those of R. argentiventer. According to Med-

way & Lim ( 1966), where R. tiomanicus and R. tanezu-

mi co-exist, the upper toothrow is shorter in R. tiomani-

cus (= tiomanicus and jalorensis in Medway & Lim 1966)

than in R. tanezumi (= diardii in Medway & Lim 1966).

In addition, the supraorbital ridges of R. tiomanicus flow

backwards on either side of the skull in smooth, vase-

shaped curves (Fig. 43). This is in contrast to R. tanezu-

mi, where the supraorbital ridges are high and wide and

form a conspicuous angular shelf just behind the interor-

bital area (Musser & Califia 1982). Photographs of the

skull of R. tiomanicus are included in Musser & Newcomb

( 1 983, Figures 1 1 0 and 111).

Sperm morphology. Information on the sperm morphol-

ogy of two specimens from peninsular Malaysia is includ-

ed in Breed & Yong (1986).

Fossil history. There are no recorded fossil specimens of

R. tiomanicus from Thailand (Pearch et al. 2013).

Taxonomic notes. Previously many specimens here re-

ferred to tiomanicus were included in R. rattus jaloren-

sis. However, Medway & Lim (1966) explored the rela-

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

73

Fig. 45. Distribution of Rattus tiomanicus in the Myanmar-

Thai-Malaysian peninsula and Singapore. Thailand: 1. Raj-

japrabha Dam; 2. Klong Phraya Wildlife Sanctuary; 3. Ghirbi;

4. Junk Seylon (= Phuket); 5. Pulau Telibon; 6. Khuan Kalong;

7. Talok Udang, Terutau Island; 8. Nong Chik; 9. Biserat.

Malaysia: 10. Langkawi Island; 1 1 . Pulau Dayang Bunting; 12.

Paya Island; 13. Kedah Peak; 14. Penang Island; 15. Temengoh;

16. E. Perhentian Island; 17. Great Redang Island; 18. Bukit

Jong; 19. Tenggol Island; 20. Telom; 21 . Taiping Hill (near); 22.

Tanjong Huntu; 23. Pangkor Besar Island; 24. Pulau Rumpia;

25. Jarak Island; 26. Bukit Fraser; 27. Bukit Lanjang Forest Re-

serve; 28. Subang Forest Reserve; 29. Kuala Lumpur; 30. Cheras;

31. Gunong Tampin; 32. Kaban Island; 33. Sribuat Island; 34.

Tioman Island; 35. Pemanggil Island; 36. Pulau Babi; 37. Aur

Island; 38. Tinggi Island; 39. Pisang Island; 40. Pelepak; 41.

Kangka; 42. Pasir Gudang. Singapore: 43. “Various localities”.

For full locality details, see Gazetteer and Appendix I.

tionships between tiomanicus, jalorensis, and diardii

Jentink (which was described from western Java). They

concluded that tiomanicus was the prior name for jaloren-

sis and that these taxa were separate from diardii. This

view was supported by the chromosomal characters ( Yong

et al. 1972) and also by an analysis of red cell proteins

(Chan 1977). Musser & Califta ( 1 982) followed this view,

referring diardii to R. rattus (now considered to be R.

tanezumi in Southeast Asia, Musser & Carleton 2005) and

treating jalorensis as a synonym of R. tiomanicus. They

included a list of synonyms of tiomanicus from through-

out its range and outlined its geographically variable char-

acters. Previously, Hill (1960) provided descriptions of the

pelage colour and measurements (see also Table 23) for

most of the taxa currently referred to R. tiomanicus from

peninsular Thailand and Malaysia.

Distribution and conservation status. Rattus tiomanicus

is a Sundaic species with a range extending from penin-

sular Thailand to Sumatra, Java, Bali, Borneo and

Palawan (Musser & Carleton 2005). Within the study area,

it is recorded from southern peninsular Thailand and

Malaysia and Singapore (Fig. 45).

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, presumed large pop-

ulation, tolerance of a broad range of habitats, and because

its population is likely to be increasing. It is considered a

pest in parts of its range” (Aplin & Frost 2008).

Ecology and reproduction. In southern peninsular Thai-

land, R. tiomanicus was collected in secondary evergreen

forest and in an oil palm plantation in Surat Thani Province

at approximately 400 metres a.s.l. (UP unpublished data).

In mainland Malaysia, this species was found in gardens,

plantations, scrub, secondary forest, mangrove and dis-

turbed or regenerating woodland. It tends to avoid primary

forest, although some island populations are found in high

forest. It enters houses freely (Medway 1969). Although

frequently trapped on the ground, it is also an accom-

plished climber. In oil palm plantations, where it can be

a major pest species (reducing yields by up to 5 %), it of-

ten shelters in piles of cut fronds and less frequently in

cut stumps or fallen logs. Very occasionally, it is found in

terrestrial burrows. Its nests are thought to be off the

ground and presumably in crowns of palms and in hol-

low stumps and logs. Litter size ranges from 2 to 7 with

a mean size of 4.4. Average female pregnancy rates were

17.6 % (27.9 % if restricted to sexually mature individu-

als) on an annual basis (Aplin et al. 2003). Oestrus occurs

every 5-8 days and the gestation period is 21-22 days.

Young reach sexual maturity after about 85 days (Med-

way 1969).

Genus Sundamys

Sundamys rats

Sundamys Musser & Newcomb 1983: 401; type species Mus

muelleri Jentink.

Remarks. This genus is endemic to the Malaysian penin-

sula and islands on the Sunda Shelf. There are three

species. Two are extralimital. Sundamys infraluteus

(Thomas) is restricted to northern Borneo and western

Sumatra. Sundamys maxi ( Sody) is only known from west-

ern Java. Sundamys muelleri is the most widespread. De-

scribed from western Sumatra, its range extends to penin-

sular Myanmar, Thailand and Malaysia, Borneo to the

Philippines and numerous Sundaic islands (Musser &

Newcomb 1983).

Description. The generic description is omitted since there

is only one species of this genus in the study area.

Sundamys muelleri

Muller’s Sundamys

Mus mulleri Jentink 1880: 16; Batang Singgalang, Pada High-

lands, western Sumatra.

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Uraipom Pimsai et al.

Fig. 46. Skull and dentition of Sundamys muel-

leri (54M392 CTNCR), Mueang District, Ra-

nong Province, peninsular Thailand. (A): dorsal,

ventral and lateral view of skull and mandible

( Scale = 5 mm ); ( B ) : occlusal view of upper (left)

and lower (right) left molars (Scale = 2 mm); (C):

left hind foot of S. muelleri

(PSUZC-MM20 12.205), Na Thawi District,

Songkhla Province, peninsular Thailand (Scale =

5 mm). Indicative numbers (i-xxii) and ( 1-3) and

letters (a-ac) are explained in the text ‘Descrip-

tion’ for S. muelleri. M: mouth; lab: labial.

M. validus Miller 1900d: 141; Khow Sai Dow, Trang, peninsu-

lar Thailand.

Mus muelleri foederis Robinson & Kloss 1911b: 245; Ulu

Temengor, Perak, Malaysia.

Rattus victor Miller 1913: 16; near mouth of Rompin River, Pa-

hang, Malaysia.

Description (based on personal observation and Musser

& Newcomb 1983). The body size ranges from large to

very large (Table 24) with a body weight according to

Rudd ( 1 965 ) for males ranging from 2 1 0-4 1 2 g and for

females from 206^441 g and according to Lim (1970) a

mean for 30 males of 335 g and 30 females of 292 g. The

tail is longer than the head and body and unicoloured, dark

brown above and below. Tail scales are large with 9-12

rows of scales/cm. The fur on the upper parts of the body

is thick, shaggy and slightly ‘harsh’ to touch but is with-

out spines (Fig. 51G). There are over-hairs on the rump

(12-18 mm in length) and black guard hairs on the back

and rump (8-15 mm). The colour is dark tawny brown,

darkest in the midline of the back and paler on the flanks.

The under parts have soft, dense and short (6-8 mm) fur.

The colour varies between individuals and geographical-

ly. According to Musser & Newcomb (1983), specimens

from peninsular Malaysia have under parts that are white,

cream, grey or buffy-grey; sometimes the chin and throat

are paler than the rest of the under parts. There is usual-

ly a clear line of demarcation between the dorsal and ven-

tral pelage along the flanks. The ears are small, round, and

dark brown. The hind feet (Fig. 46C) are broad, moder-

ately long, and brown, above and below; they have large

interdigital pads ( 1 ), a conspicuous outer metarsal pad (2)

and an inner metarsal pad (3), which is elongated and kid-

ney-shaped; the claws are unpigmented. Females have

eight mammae: one pair pectoral, one postaxillary and two

inguinal.

The skull is medium to large and robust. The rostrum

(Fig. 46Aiii) is long and wide, sufficiently broad almost

to enclose the nasolacrimal capsules (iv). The nasals (ii)

slightly exceed the length of the rostrum and incisors (xix);

their tips (i) are pointed or rounded. The braincase is long,

broadest between the squamosal roots (xvii), which are set

high on the side of the braincase; it narrows towards the

interorbital area (viii). The supraorbital ridges (vi) are

well-developed; they continue on the lateral margins of

the braincase and terminate at the supraoccipital (xvii).

The zygomata (v) are robust, parallel with each other or

slightly narrowed anteriorly. The incisive foramina (ix) are

wide but short relative to the length of the diastema; their

posterior margins (x) are variable in position, being in

front of, in line with, or posterior to the anterior surfaces

of the first molars ( M 1 ) (xi). The palate (xvi) is long and

broad; its posterior border (xiii) extends beyond the last

upper molars (M3) (xii). The distance from the back edge

of the palate to the ventral margin of the foramen mag-

num (xv) (postpalatal length) is shorter than the palatal

length. The tympanic bullae (xiv) are very small relative

to the size of the skull. In each half mandible, the coro-

noid process (xx) is prominent; the angular process (xxi)

is well-developed but thin; the incisor root (xxii) forms a

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

75

moderately defined process. Numerous photographs of the

skull are included in Musser & Newcomb ( 1983, includ-

ing Figures 33, 42 and 60) and in Marshall (1988, page

480).

The incisors are large and sturdy; the enamel layers are

smooth and deep orange. The upper incisors are ortho-

dont/opisthodont (xix). The molars are slightly cuspidate

in young individuals but become laminate with age (Fig.

5G/H and O/P). The first upper molar (M1) is long and

wide and slightly overlaps the second (M2), which in turn

overlaps the third (M3). In M1, cusps tl, t2, and t3 (Fig.

46Ba) are fused in older individuals (tl is separate in

younger individuals. Fig. 5G); cusps t4, t5, t6 (b) are al-

so fused; the third row comprises one large medial cusp,

which is representative of both t8 and t9 (c); cusp t7 is

absent; in some individuals there is a small, wedge-shaped

posterior cingulum (illustrated in Figure 40 in Musser &

Newcomb 1983). In the second upper molar (M2), there

is a large tl (d) and a smaller t3 (e); in the second row,

cusps t4 (f), t5 (g) and t6 (h) appear fused in some indi-

viduals and less so in others (Fig. 5G and H); in the third

row, cusp t7 is absent; cusps t8 and t9 (i) are fused. In the

third upper molar (M3), cusp tl (j ) is large and prominent;

cusp t3 is usually present but smaller than tl (absent in

Figs 5G and 46B); cusps t4, t5, t6 are fused; the last row

comprises cusp t8 and possibly cusp t9; sometimes cusps

t4, t5, t6, t8 and t9 (k) are fused (as illustrated in Fig. 46B)

as a result of tooth wear.

In the mandibular toothrow, the first lower molar ( M , )

has an anterocentral cusp in approximately half of indi-

viduals (according to Musser & Newcomb 1983), it is ab-

sent in the specimens illustrated in Figs. 50, 5P and 46B;

the anterolabial (1) and anterolingual (m) cusps are fused;

the protoconid ( n ) is fused with the metaconid cusp (o);

a small posterior labial cusplet (p) may or may not be pres-

ent, it is fused with the hypoconid (q) and entoconid cusps

(r) in Fig. 46B but is separate in the younger individual

in Fig. 50; a well-developed posterior cingulum (s) is

present. In the second lower molar (M2), a large anterior

labial cusp (t) is fused with the protoconid (u) and meta-

conid cusps (v) (the anterior labial cusp is not seen in Fig.

50); a posterior labial cusplet (w) is always present, it is

fused, in older individuals, with the hyconid (x) and en-

toconid cusps (y); the posterior cingulum (z) is well-de-

veloped. In the third lower molar (M3), the protoconid (aa)

is fused with the metaconid cusp (ab); the posterior lam-

ina is constituted only by the entoconid cusp (ac).

Karyology. 2N= 42, FN= 59 (male), FN= 58 (female);

there are six pairs of small metacentric chromosomes; one

pair of large subtelocentric chromosomes, one pair of

small subtelocentric chromosomes, 1 2 pairs of telocentric

chromosomes; the X sex chromosome is a large submeta-

centric; the Y sex chromosome is a small telocentric

(Musser & Newcomb 1983).

Sperm morphology. Information on the sperm morphol-

ogy of a specimen from peninsular Malaysia is included

in Breed & Yong (1986).

Fossil history. There are no recorded fossil specimens of

S. muelleri from Thailand (Pearch et al. 2013).

Taxonomic notes. Specimens from peninsular Myanmar-

Thailand-Malaysia are referred to S. m. validus. The ex-

ternal, cranial and dental dimensions are much larger than

those from elsewhere in the species’ range. Musser &

Newcomb ( 1983) suggested that the differences were so

great as to indicate possibly a specific difference. It should

be noted that the smaller size of the holotype of foederis

(BM.21. 1 1 .8.41) (Table 24) reflects the fact that it is a very

young adult-juvenile (Guy Musser pers. comm.). Latinne

et al. (2013b) suggested that there were two genetic lin-

eages of Sundamys in Thailand but that the genetic dis-

tance was low (less than 2.5 %).

Distribution and conservation status. Sundamys muel-

leri has a range that extends from peninsular Myanmar,

Thailand and Malaysia to Sumatra, Borneo, the Philip-

pines and numerous Sundaic islands (Musser & Newcomb

1983). Its distribution is illustrated in Fig. 47.

Its conservation status is listed by IUCN as ‘Least Con-

cern’ “in view of its wide distribution, presumed large pop-

ulation, it occurs in a number of protected areas, and be-

cause it is unlikely to be declining at nearly the rate re-

quired to qualify for listing in a threatened category”

(Ruedas et al. 2008d).

Ecology and reproduction. Sundamys muelleri is essen-

tially a nocturnal, terrestrial rodent although it can climb

the branches of trees. It nests on the ground, usually near

streams and is a good swimmer. In Malaysia, it favours

low-lying land and valley bottoms in primary and tall sec-

ondary forest. It normally spends the day in one or more

regularly used dens, under logs, in holes in the ground or

under the roots of trees. It emerges in late afternoon or ear-

ly evening, although occasionally it may be active in the

middle of the day (Harrison 1957a, Medway 1969, Muss-

er & Newcomb 1983). According Lim 1970, it is rarely

found in lowland primary forest but is abundant in low-

land disturbed primary and secondary forests and man-

grove forests.

In peninsular Thailand, it was collected in primary for-

est at approximately 95 metres a.s.l. in Phatthalung

Province; in evergreen forest at approximately 190 m. a.s.l.

in Chumphon Province and at approximately 50 m. a.s.l.

in Songkhla Provinc; and in secondary evergreen forest

in Phetchaburi Province (UP unpublished data).

In general, it is a species of lowland forest that has rarely

been encountered above 925 metres (3000 feet), although

Yong (1969c) reports it from 985 metres (3200 feet) on

Kedah Peak and Musser & Newcomb (1983) from 1075

metres (3500 feet) from Maxwell’s Hill and Telom in Per-

ak, Malaysia. Its diet consists of insects, fruits, leaves,

shoots and other vegetable matter; also crabs, land snails.

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Uraiporn Pimsai et al.

Fig. 47. Distribution of Sundamys muelleri in the Myanmar-

Thai-Malaysian peninsula and Singapore. Myanmar: 1.

Thagyet; 2. Maliwun. Thailand: 3. Pa La U; 4. Klong Bang

Jai; 5. Ban Tha San; 6. Tasan; 7. Ban Bang Non; 8. Ranong; 9.

Pa Toh Watershed Management; 10. Khao Nong; 11. Ban Plai

Num; 12. Tang Pran; 13. Khao Luang; 14. Nam Tok Tha-Phae;

15. Nakhon Si Thammarat; 16. Khao Phu Khao Ya National

Park; 17. Trang; 18. Ban Khao Chong; 19. Khow Sai Dow; 20.

Nga Chang Waterfall; 21. Kuan Khao Wang Forest Park; 22.

Khao Num Kang National Park; 23. Na Pra Du; 24. Ban Ya Kan;

25. Narathiwat; 26. Tonto Waterfall. Malaysia: 27. Temengoh;

28. Kuala Longnai; 29. Maxwell’s Hill; 30. Ulu Temengor; 31.

Telom; 32. Changkat Mentri; 33. Bukit Besi; 34. Semangko Pass;

35. Ulu Cemperoh; 36. Ginting Bidei; 37. Ulu Gombak; 38.

Kampong Janda Baik; 39. Bukit Lanjang Forest Reserve; 40.

Bukit Lagong; 41. Kepong; 42. Subang; 43. Kuala Lumpur; 44.

Ulu Langat; 45. Rumpin River (near mouth of); 46. Malacca;

47. Sembrong / Sembrong River; 48. Kangka Kuli; 49. Tamok;

50. Kudong. Not located: Ka Kuli (poss. = 48). For full local-

ity details, see Gazetteer and Appendix I.

and even a partially digested caecilian was found in the

stomach of one specimen (Musser & Newcomb 1983).

The young are born in nests built in sheltered positions.

In Malaysia, pregnant females were recorded in all months

but most frequently in the period July to September. Mean

litter size was 3.8, with a range of 1 to 9) (Medway 1969).

GENERIC KEYS

In order to facilitate the correct identification of specimens

to generic level, a range of characters, external, cranial,

and dental has been incorporated into a matrix key (Ta-

bles 25A, B & C). Each character, where possible has been

linked to a drawing or a photograph to help explain the

feature being described. For external characters, a sum-

mary is provided in Figs 48, 49 and 50, with an addition-

al set of colour photographs of the most common murine

genera in the region included in Fig. 51. For the cranial

and dental characters, reference is made to particular fea-

tures illustrated in figures included in the main body of

the paper. For measurements, the reader is guided to the

relevant Tables.

The grouping of the genera corresponds to a group of

four genera that are arboreal specialists, with complex den-

tition, which includes cusp t7 on the first upper molar (M1)

and feet adapted to climbing, most especially large plan-

tar and digitial pads (Table 25A). The second group in-

cludes one mouse genus, Mus, and three of the generally

smaller rats, although it is appreciated that some species

(and/or individuals) of these ‘smaller’ rats, especially in

the genus Rattas and to a lesser extent Maxomys can be

large. The final grouping includes the large rats,

Leopoldamys, Berylmys, Bandicota and Sundamys.

RECOMMENDATIONS

One of the primary objectives of the current study is to

provide a summary of existing knowledge of murine ro-

dents in the Myanmar-Thai-Malaysian peninsula and to

highlight priorities, which would particularly benefit from

further studies.

As is apparent from the paper, much previous taxonom-

ic research of rodents in the study area was morphomet-

ric, whereas that conducted most recently is primarily ge-

netics-based. As noted by a number of previous authors

(Musser & Carleton 2005, Pages et al. 2010, Chaval et al.

2010, Latinne et al. 2013b), it would be of considerable

benefit to combine the two approaches to maximise the

potential to determine phylogenies and resolve difficult

taxonomic problems. In the broadest terms, priority gen-

era for further taxonomic research include Rattus , Niviven-

ter and Maxomys. Particular priorities within these taxa

include:

Maxomys surifer, which as currently understood is con-

sidered to include 4 highly divergent genetic lineages

(Latinne et al. 2013b) and is most probably a complex

of cryptic species (Gorog et al. 2004, Aplin et al. 2008c,

Achmadi et al. 2013);

Niviventer fulvescens, the taxonomy of which is still open

to discussion (Musser & Carleton 2005, Pages et al.

20 1 0). Some authors recognise the peninsular form buk-

it as a discrete species (Corbet & Hill 1992, Balakirev

et al. 2011) whilst others include this taxon in fulvescens

(Abe 1983, Musser & Carleton 2005);

Rattus tanezumi and R. tiomanicus, which are probably

complexes of morphologically cryptic species. Latinne

et al. (2013b) recognised at least four distinct genetic

lineages of Rattus in peninsular Thailand, the identifi-

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Murine rodents of the Myannrar-Thai-Malaysian peninsula and Singapore

77

A MB: 69-102 mm (Table 6)

clear demarcation on flanks

thick, short, soft fur without

spines, with short guard

hairs; reddish brown

J;

HF; 15-22 mm,

white or brown,

enlarged plantar

and digital pads

(Fig. 14C) /

tail averages considerably N-. .

longer than HB,

unicoloured, brown

/

under parts white

2 pairs of mammae

tail tip with well-developed

tuft of hairs

thick, soft and woolly

fur, without spines, grey

y to greyish brown

orange-brown flank

stripe, demarcation

fairly distinct

B

under parts white sometimes • x v

with yellowish tinge; 4 pairs ,

of mammae

HF: 28-33 mm;

brown, enlarged

plantar and digital

pads tail averages considerably

HB: 140-165 mm(Table7)

longer than HB,

unicoloured, brown

C HB: 156-175* mm (Table 19)

tail averages longer than 11B

unicoloured, brown, naked

except at base,

prehensile distally

tail tip without tuft of hairs

very long, soft fur

without spines,

brownish red

tail tip with tuft of hairs

clear demarcation on flanks

under parts

creamy white

2 pairs of mammae

F1F: 26-30 mm; brown,

enlarged plantar and

digital pads

D

HB: 161-220 mm (Table 8)

Fig. 48. External characters of four genera of murine rodents (A): Chiropodomys\ (B): Hapalomys\ (C): Pithecheir ; (D): Lenothrix.

Drawings based on Francis (2008). Size approximately proportional to all genera illustrated in Figs 48, 49, and 50. To be used in

conjunction with the character matrix in Table 25A.

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78

Uraipom Pimsai et al.

HB: 55-90 mm (Table 15)

B HB: 100-280 mm (Table 12)

demarcation in

M. caroli.

indistinct in

A/. muscidus

grey to dark brown

without spines

under parts N ITT: 14-19 mm. 6 small

white to plantar pads (Fig. 28C)

greyish brown

' tail, about equal or slightly

longer than HB

bicoloured or unicoloured

tail tip without tuft pencil of

hairs

clear demarcation on flanks in

A /. rajah and A 1. surfer ; not demarcated

in M. inas and M. whileheadi

light brown to reddish brown,

grizzled with grey or black,

numerous flattened spines

under parts white,

buff, or grey;

4 pairs of mammae

tail tip without tutVpencil

of hairs;

white tip in M. sur fer

HI- : 22-46 mm: 5 or 6

smooth plantar pads

(Fig. 23C)

C HB: 110-170 mm (Table 17)

tail about equal or slightly

shorter/longer than HB

bicoloured (but less

distinctly in M. rajah)

greyish to yellowish brown to

reddish brown, numerous

flattened spines and black / /

guard hairs

A

FIF: 20-38 mm; 6

(Fig. 3 1C)

tail long, always longer

(110-180%) than HB

bicoloured (but unicoloured in

N. cremoriventer)

clear demarcation on flanks

under parts white, buff,

yellowish white;

4 pairs of mammae

tail tip with tuft, pencil

of hairs (except for

N. cameroni)

Fig. 49. External characters of four genera of murine rodents (A): Mus; (B); Maxomys; (C): Niviventer; (D): Rattus. Drawings

based on Francis (2008). Size approximately proportional to all genera illustrated in Figs 48, 49, and 50. Flowever, it should be

noted that there is considerable variation in size between species, especially in Rattus and Maxomys. To be used in conjunction

with the character matrix in Table 25B.

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Murine rodents of the Myanmar-Thai-Malaysian peninsula and Singapore

79

A MB: 180-290 mm (Table 9)

greyish, dark brown, golden-orange.

Fig. 50. External characters of four genera of murine rodents (A): Leopoldamys; (B): Berylmys; (C): Bandicota', (D): Sundamys.

Drawings based on Francis (2008). Size approximately proportional to all genera illustrated in Figs 48, 49, and 50. However, it

should be noted that there is considerable variation in size between species, especially in Bandicota and Berylmys. To be used in

conjunction with the character matrix in Table 25C.

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Uraipom Pimsai et al.

cation of which were not determined. Pages et al. (2010)

suggested further research incorporating molecular da-

ta from the holotypes of the different Rattus taxa, es-

pecially the numerous synonyms, would be the most

suitable way to determine accurately the taxonomy of

this genus in Southeast Asia. Aplin et al. (2011 ) noted

that more research was required to determine the true

phylogeny of the R. rattus-R. tanezumi species complex.

Outside these ‘problem genera’, there are also other in-

teresting, unresolved taxonomic ambiguities:

Berylmys berdmorei and especially its relationship to the

taxon mullulus. In general, the geographical variation

of B. berdmorei from throughout its range is little un-

derstood (Musser & Newcomb 1983);

Berylmys bowersi; Musser & Newcomb ( 1 983 ) noted dif-

ferences in cranial morphology between Indochinese

and Sundaic populations. This observation was support-

ed by both Pages et al. (2010) and Latinne et al. (2013b)

who recognised two distinct genetic lineages, one from

northern Thailand and one from Kanchanaburi Province

and peninsular Thailand;

Leopoldamys sabanus, which may be a species complex.

Musser (1981) and Musser & Carleton (2005) remarked

on the considerable morphological variation between

populations of L. sabanus from the Indochinese and

Sundaic subregions and among insular populations from

the Sundaic subregion. Subsequently, Latinne et al.

(2013b) suggested that L. sabanus (sensu stricto) is es-

sentially confined to the Sundaic subregion whilst Bal-

akirev et al. (2013) restricted L. sabanus to Borneo and

referred specimens from peninsular Thailand-Malaysia

to L. vociferans.

In terms of conservation, there are a number of taxa that

require extensive further research. Much of this is ecolog-

ical, determining the reliance of a particular species on a

particular habitat type. Other studies need to focus on pop-

ulation status and determine existing and potential threats.

Species of special interest include:

Hapalomys longicaudatus, which is listed as ‘Endangered’

by IUCN and is thought to be possibly extinct in Thai-

land and Myanmar (Aplin & Lunde 2008a). A high pri-

ority is to determine its distribution and population sta-

tus in the remaining areas of undisturbed bamboo habi-

tat within evergreen lowland forest;

Maxomys rajah and M. whiteheadi , both of which are list-

ed as ‘Vulnerable’ by IUCN. Both are thought to be at

risk because of the extensive loss and degradation of

lowland forest, which is believed to be their favoured

habitat (Aplin et al. 2008e). Currently, data on their ecol-

ogy and behaviour are very limited and much further

study is required;

Niviventer cameroni is a montane species, endemic to the

Cameron Highlands of peninsular Malaysia. Listed as

‘Vulnerable’ by IUCN, it is thought to be threatened by

habitat loss but currently very little known about its ecol-

ogy (Musser & Ruedas 2008);

Niviventer cremoriventer is another ‘Vulnerable’ species,

whose ecology, population status and behaviour has

been little researched (Ruedas et al. 2008e);

Pithecheir pan’us is listed as ‘Data Deficient’ by IUCN

(Aplin et al. 2008h). Further study is needed to deter-

mine the habitat preferences of this species in order to

support the findings of Muul & Lim (1971) that this

species is found in both secondary and pristine forest.

In terms of ecology and reproductive biology, little is

known about Bandicota savilei, Berylmys berdmorei, or

Lenothrix canus in the study area. In addition, studies fo-

cusing on zoogeography and phylogeography would be

of considerable interest.

Finally, rodents are frequently regarded by man in a neg-

ative context. It would therefore be of great value to un-

dertake some detailed research of the ecosystem services

provided by this diverse, adaptable and often abundant

group of small mammals.

Acknowledgements. In Thailand, UP is grateful to the Depart-

ment of Biology and the Research Assistant Scholarship of the

Faculty of Science and Graduate School of the Prince of Songk-

la University (PSU) for academic and financial support. We are

pleased to thank Phannae Sa-ardit, Pipat Soisook, Yingyod Lap-

wong, Amphom Plapplueng, and all the staff of the Princess Ma-

lta Chakri Sirindhorn Natural History Museum for access to the

Museum’s specimen collection.

UP is especially grateful to Christopher Imakando for donat-

ing photographs of some live rodents as well as all members of

the Small Mammals and Birds Research Unit at PSU for their

help with fieldwork, specimen preparation, and for their encour-

agement, including Bounsavane Douangboubpha, Ith Saveng,

Ariya Dejtaradol, Sunate Karapan, Tuanjit Srithongchuay,

Daosavanh Sanamxay, and Ngagyel Tenzin. UP is grateful also

to Dr. Surachit Waengsothom of the Thailand Institute of Sci-

entific and Technological Research (TISTR), who permitted the

examination and loan of specimens in his collection. Finally, UP

would like to thank her father, mother, sisters, brothers and all

of her family members for their support and encouragement.

In the United Kingdom, the authors are grateful to David Har-

rison, Nikky Thomas, and Beatrix Lanzinger of the Harrison In-

stitute for their advice and for their helpful provision of litera-

ture. We are also grateful to the staff of the Mammal Section

and General Library of the Natural History Museum, London,

especially Paula Jenkins, for granting access to the collections

and for locating publications. Elsewhere, we thank Bandana Aul

of the Bombay Natural History Society and Alice Latinne of the

Universite de Liege for forwarding valuable literature.

We are pleased to acknowledge the Darwin Initiative (Project

No. 18-002), who have generously supported the training of a

new generation of young, in-country taxonomists (including UP)

in Southeast Asia and the Systematics Association, UK for their

financial support of UP for visits to the research collections of

the Thailand Institute of Scientific and Technological Research.

Finally, we thank Guy Musser and one other anonymous review-

er for their most valuable comments on the manuscript.

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Fig. 51. Seven common murine genera in peninsular Myanmar-Thai-Malaysia. (A): Maxomys (M. surifer); (B): Niviventer ( N .

cremoriventer)\ (C): Rattus ( R . tanezumi)\ (D): Leopoldamys (L. sabanus)', (E): Berylmys (B. bowersi)\ (F): Bandicota (B. indica);

(G): Sundamys (S. muelleri). Not to scale.

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Table 1. A list, in chronological order, of the 95 taxa of murine rodent described from the Myanmar-Thai-Malaysian peninsula

and Singapore. Taxa in bold are currently recognised as distinct species (Musser & Carleton 2005).

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Table 2. External, cranial and dental measurements of Bandicota. Measurements sourced from: [i] Musser & Brothers (1994); | iij

Chasen (1936: Chasen selected the ‘five largest specimens’ available to him) and [iii] Pimsai (2012).

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Table 3. Matrix key to the three species of Bandicota currently known from the Myanmar-Thai-Malaysian peninsula.

Table 4. External, cranial and dental measurements of Berylmys. Measurements sourced from: [i] Musser & Newcomb (1983),

[ii] Pimsai (2012) and [hi] Miller ( 1900b).

Table 5. Matrix key to the two species of Berylmys currently known from the Myanmar-Thai-Malaysian peninsula.

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Table 6. External, cranial and dental measurements of Chiropodomys gliroides. Measurements sourced from: [i] Musser (1979)

and [ii] Pimsai (2012).

Table 7. External, cranial and dental measurements of Hapalomys longicaudatus. Measurements sourced from: [i] Musser ( 1972)

and [ii] Musser & Newcomb (1983).

Table 8. External, cranial and dental measurements of Lenothrix canus. Measurements sourced from: [i] Musser & Newcomb (1983),

[ii] Kloss (1931) and [iii] Muul & Lim (1971).

Table 9. External, cranial and dental measurements of Leopoldamys. Measurements sourced from: [i] Yong ( 1970), [ii] Medway

(1969), [iii] Bonhote (1900), [iv] Pimsai (2012) and [v] Miller (1900b).

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Table 10. Matrix key to the two species of Leopoldamys (sensu Musser & Carleton 2005) currently known from the Myanmar-

Thai-Malaysian peninsula.

Table 11. External, cranial and dental measurements of Leopoldamys sabanus (sensu Musser & Carleton 2005) from a range of

localities (see Fig. 22) in Myanmar, Thailand and Malaysia. Measurements sourced from [i] Miller (1903a), [ii] Hill (1960), [iii]

Miller ( 1913), and [iv] Chasen (1940).

Table 13. Matrix key to the four species of Maxomys (sensu Musser & Carleton 2005) currently known from the Myanmar-Thai-

Malaysian peninsula.

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Tabic* 12. External, cranial and dental measurements of Maxomvs. Measurements sourced from: [i] Musser et al. ( 1 979), [ii] Hill

(1960), [iii] Medway (1969) and [iv] Pimsai (2012).

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Table 14. External, cranial and dental measurements of Mcixomys surifer (sensu Musser & Carleton 2005) from a range of lo-

calities (see Fig. 25) in Myanmar. Thailand and Malaysia. Measurements sourced from [i] Miller (1903a), [ii] Hill (1960), [iii]

Kloss (1916c) and [iv] Kloss (1911a).

Table 15. External, cranial and dental measurements of Mus. Measurements sourced from: [i] Aplin et al. (2003), [iij Marshall

1977a, [iii] Francis (2008), [iv] Bonhote (1902), [v] Kloss (1921), [vi] Pimsai (2012), and [vii] Medway (1969).

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Table 16. Matrix key to the two species of Mus currently known from the Myanmar-Thai-Malaysian peninsula.

Table 17. External, cranial and dental measurements of Niviventer. Measurements sourced from: [i] Chasen (1940), [iia] Med-

way ( 1969, = R. fulvescens), [iib] Medway (1969), [iii] Francis (2008), [iv] Musser (1973a); [v] Miller (1903a), [vi] Miller (1913),

[vii] Pimsai (2012), [viii] Abe ( 1983), [ix] Robinson & Kloss ( 1914) and [x] Bonhote (1903b).

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Table 18. Matrix key to the three species of Niviventer (sensu Musser & Carleton 2005) currently known from the Myanmar-

Thai-Malaysian peninsula.

Table 19. External, cranial and dental measurements of Pithecheir. Measurements sourced from: [i] Musser & Newcomb (1983)

and [iij Kloss (1916b).

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Table 20. External, cranial and dental measurements of Rattus. Measurements sourced from: [i] Robinson & Kloss 1914, [ii] Hill

( 1960), [iii] Marshall (1988), [iv] Corbet & Hill 1992, [v] Harrison Institute collection, [vi] Medway (1969); [vii] : Bonhote (1903a),

[viii] Kloss 1908a, [ix] Pirnsai 2012, [x] Sody ( 1941), [xi] Miller ( 1900a) and [xii] Medway & Lim (1966).

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Table 21. Matrix key to the seven species of Rattus currently known from the Myanmar-Thai-Malaysian peninsula.

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Table 22. External, cranial and dental measurements of Rattus tanezumi from a range of localities (see Fig. 42) in Myanmar,

Thailand and Malaysia. Sourced from [i] Hinton (1919), [ii] Miller ( 1 913), [iii] Hill (1960), [iv] Chasen (1937), [v] Miller ( 1900c)

and [vi] Bonhote ( 1903a).

1): 15-1

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Table 23. External, cranial and dental measurements of Rattus tiomanicus from a range of localities (see Fig. 43) in Myanmar,

Thailand and Malaysia. Based on [i] Hill (1960), [li] Chasen & Kloss ( 1931 ) and [iii] Yong (1971).

Table 24. External, cranial and dental measurements of Sundamys. Measurements sourced from: [i] Pimsai (2012), [ii] Miller

(1900d), [iii] Musser & Newcomb (1983), [iv] Medway (1969), [v] Miller (1913) and [vi] Robinson & Kloss (1911b).

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Table 25A. Character matrix of four murine genera: Chiropodomys, Hapalomys, Pithecheir and Lenothrix. *: measurements of

the holotype of Pithecheir parvus are omitted (see Table 19).

Table 25 B. Character matrix of four murine genera: Mus, Maxomys, Niviventer and Rattus.

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Table 25C. Character matrix of four murine genera: Leopoldamys, Berylmys, Bandicota and Sundamys.

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June 2014

First record of the parasitoid wasp

Tachinaephagus zealandicus Ashmead, 1904

(Hymenoptera: Chalcidoidea: Encyrtidae) in Germany

Ralph S. Peters

Zoologisches Forschungsmuseum A. Koenig , Adenauerallee 160, 53113 Bonn, Germany; E-mail: R.Peters@zfmk.de.

Abstract. The first record of the parasitoid wasp Tachinaephagus zealandicus from Gennany is presented. A total of twelve

specimens were reared from a puparium of Lucilia sp. (Diptera: Calliphoridae), collected from compost in Ingelheim am

Rhein (Rhineland-Palatinate). While probably originating from Australia, the species is now cosmopolitan with numer-

ous records from all continents. Its hosts are mainly synanthropic flies, and it has been repeatedly used as biological con-

trol agent. Given its synanthropy and its uniqueness as one of very few encyrtid species that parasitize fly puparia, it is

unlikely that it has been overlooked in the past. It is, however, unknown where it comes from and why it appears now

for the first time. This perplexity demonstrates the need for more thorough studies on parasitoid wasps and other noto-

riously understudied taxa. Currently, monitoring appearance of invasive species and other potentially influencing faunal

changes still relies mostly on chance findings. A combination of large DNA barcoding and collecting initiatives such as

GBOL and digital accessibility of zoological collection data could help improving this problematic situation in the fu-

ture.

Keywords. Parasitoids, Germany, faunistics.

INTRODUCTION

Tachinaephagus zealandicus (Hymenoptera: Chalci-

doidea: Encyrtidae) was first described from Australia

(Ashmead 1904) and probably originates from the South-

ern Hemisphere (Olten 1971), but has been distributed by

man over many parts of the world. Distribution records

include New Zealand, Argentina, Brazil, Georgia, South

Africa, Java, Carribian Islands, Israel, Mauritius, New

Caledonia, and USA (see Noyes 2012 and references

therein). It is a larval-pupal parasitoid of cyclorrhaphous

Diptera, including many synanthropic species such as

Musca domestica (house fly), Stomoxvs calcitrans (stable

fly), Fannia canicularis (lesser house fly), and Lucilia