Table Of ContentBonn 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
, , ,
'PrincessMaha ChakriSirindhorn NaturalHistoiyMuseum, PrinceofSongkla University>, Hat Yai. Songkhla Province, Thai-
land90112. E-mail: [email protected]:
departmentofBiology, FacultyofScience, PrinceofSongkla University\ Hat Yai, Songkhla Province, Thailand 90112:
sHarrisonInstitute, Bower-woodHouse, 15StBotolph’sRoad, Sevenoaks, Kent, TNI3 3AQ, UK: E-mail:
[email protected]
Abstract. Basedon field surveys undertaken between 2010 and 2013, museum studies inThailand and theUK, and an
extensive literaturereview, thispaperprovides informationonthe28 speciesand 12 generaofmurinerodentscurrently
known from peninsularMyanmar,Thailand and Malaysiaand Singapore. It incoiporatesadetailedsummaryofpastre-
search, 1851-2013,oftheMurinae inthestudyareaand includesdescriptivecharactersoftheexternal,cranial andden-
talmorphologyandmeasurementsforeachoftherodentspecies. It listsandmapsthe93 murinetaxadescribedfromthe
peninsula, 84 ofwhich arecurrentlyconsideredto be synonymsat species level. Each ofthe 389 different localities on
the28distributionmapsisnumberedand linkedto itssource,eitherliteratureormuseumspecimen,and listed intheon-
line gazetteer. The global conservation status ofeach species is obtained from the IUCN Red List. Remarks are made,
wheredataareavailable, on theecology, karyology, fossil history, sperm morphology, phylogeny, andtaxonomic histo-
ryandambiguities. Recommendationsaremadeforfurtherresearch.Aseriesofillustratedmatrixkeysisprovidedtoas-
sistwith the identificationofall themurinegeneraand species within thestudyarea.
Key words.Taxonomy,distribution, identificationkeys, karyology, ecology, conservation status.
INTRODUCTION
Witli2,277 species,theorderRodentia representsapprox- Witmer 2004). Some ofthe mostvirulentdiseases borne
imately42 %ofall mammaldiversity(Musser& Carleton byrodentsin SoutheastAsia include: hantavirus, leishma-
2005). Geographically widespread and highly adaptable, nia infection, leptospirosis, scrub typhus, toxoplasmosis
rodents occupy a vast array ofdiverse ecological niches. and viral haemorrhagic fevers (Chaval et al. 2010; Her-
They impact on the composition, structure, and succes- breteauetal. 2012,http://www.ceropath.org/research/ro-
sion ofvegetation and fulfil many important ecosystem dent bomediseases).
services, includingassistingwithnutrientcyclingandthe Recognisingthe importance ofrodents, both as biolog-
dispersal ofseedand spores.Throughtheirburrowingac- icalentitiesandbecauseoftheirassociationwithman,the
tivities, theymixand aerate soils andwiththeirhigh bio- currentpaper,which is basedontheunpublishedworkof
mass, theyprovide an essential prey base formany pred- Pimsai (2012), seeks to provide a baseline for future re-
ator species (Witrner 2004). search of murines in peninsular Myanmar, Thailand,
A minority of species causes significant problems to Malaysia and Singapore. The study area is illustrated in
man. Forexample, inAsia, it isconsideredthat inanyone dark grey in Fig. 1 and all subsequent maps. It should be
%
particularareabetween 5 and 10 ofrodenttaxaare ma- notedthatalthoughthenorthernboundary isclearlyanar-
joragricultural pests(Aplin etal. 2003).As such, theyeat tifactofgeography, correspondingtothenorthernlimitof
crops in the field, typically reducing yields ofrice by 5 peninsular Thailand, it also has zoogeographical signifi-
to 10%(Aplin etal. 2003),6% inpineapples(Joomwong canceapproximatingasitdoestothe Indochinese-Sunda-
%
2007) and 5 in oil palms (Aplin et al. 2003). They eat, ic transition zone, although it is recognisedthat the exact
spoil and contaminate stored food and post-harvest loss- locationofthiszonevariesamongstdifferenttaxaandhas
%
es of20 are not unusual. They also carry diseases that been interpreted in different ways by different authors
can be transmitted eitherdirectly or indirectly to humans (Woodruff2003, Woodruff&Turner 2009, Hughes et al.
and livestock (Aplin et al. 2003; Chaval et al. 2010; 2011 ).
Received: 18.02.2014 Correspondingeditor: R. Hutterer
Accepted: 18.04.2014
-
16 Uraipom Pimsai et al.
44wereoriginallydescribedasnewspeciesand49assub-
species. Forty-two were described from peninsular
Malaysia(as understoodtoday), 30 frompeninsularThai-
land, 19 frompeninsularMyanmarand 2 from Singapore.
Most were named at the start ofthe 20"’ Century, 65 be-
tween 900and 1919and 19between 1931 and 1941 with
1
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, Rattusannandalei, andR. tiomanicus—highlight-
ed in bold in Table 1) are considered today as synonyms,
at species level. To further aid understanding ofthe tax-
onomy, the type locality ofeach taxon is plotted on the
relevant distribution map.
Thedistributionmapsarethefirstforall murinerodents
from the study region that seek to give a higher level of
geographical resolutionby featuringspot localitiesrather
than the more familiar shaded maps as available in Mar-
shall (1988), Corbet & Hill (1992), and Francis (2008).
Previously, somespot mapswereavailable,aspartoflarg-
er taxonomic studies, for certain species such as Ha-
palomys longicaudatus (Musser 1972), Chiropodomys
Fig. 1. Study area: peninsular Myanmar, Thailand and gliroides (Musser 1979), Sundamys muelleri (Musser &
Malaysia, and Singapore. Localitiesfromwhichspecimendata Newcomb 1983) and Bandicota (Musser & Brothers
were obtained are mapped and listed in the Gazetteer. 1994).
Literaturesources forthemapsweredrawn froma large
The paper focuses on the Murinae as this is the most numberofreferences datingbackto Blyth (1851). These
speciose subfamilyworldwide, with 561 species and 126 references, togetherwith many others directly relating to
genera,witlunthelargestRodentfamily, theMuridae, 730 murine rodents in the study area, are reviewed and sum-
speciesand 150 genera (Musser& Carleton 2005). With- marised below. It is hopedthat this review will provide a
in the studyarea,the Murinae,theratsandmice, havethe valuable starting point for all who wish to conduct fur-
highest population sizes, the greatest biomass, the great- ther research on murine rodents in the study area.
est impact on man ofall the rodents, and the richest di- The majority, approximately 60 % ofthe 115 publica-
versity: 28 species and 12 genera (sensu Musser & Car- tions included in the review is concerned with aspects of
leton 2005) - Bandicota bengalensis (Gray). B. indica taxonomy.Thisisespeciallythecaseforpapersand books
(Bechstein), B. savilei Thomas, Berylmys berdmorei publishedinthefirstonehundredyears(1851-1950). Dur-
(Blyth), B. bowersii(Anderson), Chiropodomysgliroides ing this time, all 40 publications were eitherexclusively,
(Blyth), Hapalomyslongicaudatus Blyth, Lenothrixcanus or primarily taxonomic and 30 included descriptions of
Miller, Leopoldamys ciliatus (Bonhote), L. sabanus new murine taxa (Table 1). Post 1950, researchers in the
(Jentink), Maxomys inas (Bonhote), M. rajah (Thomas), peninsulabeganto investigateadditionalaspectsofmurine
M. surifer (Miller), M. whiteheadi (Thomas), Mas mus- rodents, including reproduction, behaviour, ecology,
culus Linnaeus, M. caroli Bonhote, Niviventercameroni karyology, palaeontology, phylogeny, and the transfer of
(Chasen), N. cremoriventer(Miller), N.fulvescens(Gray), diseases between rodents and man.
PithecheirparvusKloss, Rattusandamanensis(Blyth), R. Aparticularaim ofthis paper(interalia) is to comple-
annandalei (Bonhote), R. argentiventer (Robinson & ment and support exciting new research looking at the
Kloss), R. exulans (Peale), R. norvegicus (Berkenhout), phylogenyofrodentsintheregionbasedoncombinedmi-
R. tanezumi(Temminck), R. tinmanicus(Miller),andSun- tochondrial and nuclear markers, for example Latinne et
damys muelleri (Jentink). al. (2012, 2013b). The paper seeks to support the idea of
As a baseline, the paper seeks to put the complex tax- a multi-approach to rodent identification as evinced by
onomyoftheMurinae fromthestudyregion intocontext. Chaval et al. (2010) by providing a detailed summary of
This subfamily includes taxa from two tribes, the Murini the alpha taxonomy ofthe Murinae. It is hoped that the
and Rattini, and one ill-defined group, the ‘Murinae in paper will facilitate future research ofrodent taxonomy
certissedis’as definedby Lecompte et al. (2008). All 93 in SoutheastAsia, andespeciallyamongstin-country stu-
taxadescribedfromtheareaare listedinTable 1. Ofthese, dents and zoologists based in Thailand, Malaysia and
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
.
Murinerodents ofthe Myanmar-Thai-Malaysianpeninsulaand Singapore 17
Myanmar,andwill helppromotestudiesthatcombine both onomists and foremost amongst these are the studies of
morphometric and molecular approaches. Musser (including Musser 1972, 1973a, 1973b, 1979,
It isanticipatedthatthepaperwill assistthosewhowish 1981; Musser et al. 1979; Musser & Brothers 1994; and
to investigate further the taxonomy ofspecies, the status Musser & Newcomb 1983).
ofwhichhasbeenquestionedinaseriesofpapersandon- Similarly, the ecological summaries in the text draw
line publications, for exampleBerylmys berdmorei, Sun- heavily onthe studies ofarange ofprevious authors, no-
damysmuelleri, andRattustiomanicus(inMusser& New- tably those ofMedway (1969) and a series ofpapers by
comb 1983),Leopoldamyssabanus(in Musser& Carleton Harrison (dating from 1950 to 1966) and Lim (1966 to
2005 and Balakirev et al. 2013)andNiviventerfulvescens 1975). Conservation status is drawn from the on-line
(in Musser & Carleton 2005 and Balakirev et al. 2011), IUCN Red List (http://www.iucnredlist.org/). Ofthe 28
Maxomyssurifer(inGorogetal. 2004,Aplinetal. 2008c species, all but six are listed as of ‘Least Concern’. The
and Latinne et al. 2013b), and Rattus tanezumi (Heaney exceptionsareone ‘Endangered’species,Hapalomyslong-
& Molur2008 andAplinetal. 2011). It isalsoinresponse icaudatus\ four ‘Vulnerable’ species,Maxomys rajah M.
topaperssuchasFrancisetal. (2010),whichalthoughpri- whiteheadi, Niviventer cameroni, and N. cremoriventer;
marily concerned with bats, suggest that the number of andone“Datadeficient’species,Pithecheirparvus How-
mammal species currently recognised in Southeast Asia ever, each ofthe28 species plays an important role in the
mayrepresentonly 50%ofthereal diversity. Inaddition, ecosystems ofthe peninsula and each in its own way is
it supports Waengsothom et al. (2009) who highlight the an important subject for future research.
need for further rodent research in Thailand. There are great opportunities for a whole range offu-
Thispaperseekstocomplementthetaxonomicworkof turestudies,notjust in thepeninsula but in SoutheastAsia
close colleagues workingon othersmall mammal taxa in generally, including taxonomy (incorporating both mor-
the region. This team ofyoung taxonomists, based in the phometries and genetic studies), phylogeny, phylogeog-
PrinceofSongklaUniversity(Thailand),National Univer- raphy and palaeontology (looking at evolutionary histo-
sity ofLaos, the Royal University ofPhnom Penh (Cam- ries), ecologyandbehaviour(especiallylinkedtoecosys-
bodia), and the Institute ofEcology and Biological Re- tem services), as well as those concentrating on the role
sources (Vietnam), have studied extensively a range of ofrodents in disease transmission. It is our intention that
taxa in mainland SoutheastAsia. Togetherwith members the current study will provide a baseline to promote and
ofthe Harrison Institute, UK and other collaborating in- facilitate this work in the Myanmar, Thailand and
stitutions, they have described several new species ofbat Malaysian peninsular and Singapore.
(Douangboubpha et al. 2011, Soisook et al. 2013a and
2013b; Thong et al. 2012) and a new Dying squirrel
(Sanamxayetal. 2013).Additionally,theyhaveundertak- LITERATURE REVIEW OF MURINE RODENT
en many studies ofsmall mammal diversity and distribu- RESEARCH 1851-2013
tion (extantandfossil)inthesamearea(Fureyetal. 2012;
Ith et al. 201 1; Kingsada et al. 2011; Pearch et al. 2013; Thefirstpublication relevanttothecurrent study is Blyth
Thomas et al. 2013). (1851) who described Mus berdmorei (= Berylmys berd-
Correct identification ofspecies is of primary impor- morei) from the Mergui (= Myeik)Archipelago (noexact
tance to many studies (Chaval et al. 2010). Some species locality). Subsequently,hepublishedontwomurinesfrom
areconsideredagricultural pestswhilstothers, evenwith- Pinang(=Penang), Malaysia(Blyth 1865); onewasanew
in the same genus, are harmless oreven beneficial, feed- taxon, Mus rama (Cantor in Blyth, 1865) (=Musmuscu-
ing on harmful invertebrates (molluscs and insects) and lus) andthe otherwasMussetifer{setiferis currentlyre-
invasiveweeds. Somespeciesarethereservoirofzoonot- ferredto Bandicota indica butthese specimens are more
,
ic diseasesdeleterioustoman,whilstothers, closely sim- probably referable to B. bengalensis). Three years later,
ilarinexternal morphologyarenot. To meet thechallenges Peters (1868) described a new taxon ofpencil-tailed tree
ofmurineidentificationwithinthestudyarea,we havede- mouse, Chiropodomys penicillatus (= C. gliroides); the
vised a series ofkeys that together, help with the identi- type localityisthoughttobetheMalaysianpeninsula, al-
ficationofall 28murinespecies. Thoseforthespeciesare though no details were given (Corbet & Hill 1992).
included in a series ofTables in the relevant sections of The first synthesis ofmammal records from the region
the text. Those for the genera are in Tables 25A, B & C. was provided by Flower (1900) who reported on 160
Thediagnosticcharactersincluded in thematrix keyshave mammal species collected from Thailand and Malaysia.
been extracted from the generic and species descriptions. He included seven murine taxa from the Thai-Malay
These matrices include a range of external, cranial and peninsula. Miller (1900a) described Mus tiomanicus (=
dental characters. Muchofthe information hasbeendrawn Rattus tiomanicus) and Mus obscurus (renamed M. pul-
fromourownobservationsbutnonewouldhavebeenpos- lus Miller, 1901, = Rattus exulans) from Tioman Island.
sible without the detailed and inspiring work ofpast tax- In the same year. Miller (1900b) published the first nra-
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
Uraiporn Pimsai et al.
jor paper dedicated to the peninsular murines. He de- currently referred to Rattus tiomanicus. In the following
scribed seven new rat taxa, which had been collected in year, Bonhote (1906) provided additional data on ten
Trong(=TrangProvince,Thailand). Ofthese,twoarestill murid taxa from southern peninsular Malaysia. This pa-
recognised as valid species today: Mus cremoriventer(= per included the description oftwo new taxa, Mus inas
Niviventercremoriventer)andM. surifer(=Maxomyssu- (=Maxomys inas) andMusklossi(=Maxomyswhitehea-
rifer).Theotherfivearetreatedassynonyms(sensuMuss- di). Thomas (1907) described two new species from
er&Carleton2005): Musvociferous(=Leopoldamyssa- Pinang(Penang),Malaysia, GunomysvarillusandG. var-
banus),M.ferreocanus(=Berylmysbowersii),M. validus ius, both are currentlyreferredtoBandicota bengalensis.
(=Sundamys muelleri), M. asper(=Maxomys whitehea- Kloss(1908a)describedtwonewmurinetaxa,Mus vil-
di), andM. pellax (=Maxomys rajah).All the type spec- losus (=Rattusannandalei) from Singapore andMussu-
imens are deposited in the United States National Muse- rifermicrodon (=Maxomyssurifer) fromTioman Island,
um. Malaysia. Hissubsequentpaperonaprovisionallistofthe
Miller (1900c) reported on fourteen mammal species mammals from the peninsular region was an update of
collectedon the Batang Islands(todayin Satun Province, Flower(1900)and included31 muridtaxa(Kloss 1908b).
Thailand) and Pulo Lankawi (today in Kedah Province, Specimenscollected fromTarutao Island(today in Sat-
Malaysia). He described four new taxa, three geograph- un Province, Thailand) and remitted to the Natural His-
icalracesandonespecies,ofmurines,Mus vociferouslan- tory Museum, London, included one new subspecies of
cavensis (= Leopoldamys sabanus), Mus suriferflavidu- murine,describedbyThomas&Wroughton(1909)asMus
lus(=Maxomyssurifer),Mussuriferbutangensis(=Max- vociferans tersus (=Leopoldamyssabanus). Meanwhile,
omyssurifer) and Muspannosus (= Rattus tanezumi). basedonacollectionfromtheTrengganuArchipelago,off
Whilst Miller was publishing in America, in Britain, thenorth-eastern coastoftheMalaysianpeninsula, Kloss
Bonhote (1900) described an isolated, endemic species, (1911a)describedtwonewsubspecies,Mussurifergran-
Mus ciliata (= Leopoldamys ciliatus), from Perak in dis andM. s.flavigrandis, both ofwhich are today treat-
Malaysia. He also recorded a numberofadditional local- ed as synonyms ofMaxomys surifer, more detailed de-
ities for a range ofmurine taxa, including Mus mettada scriptionsofthesetwotaxa, includingmeasurements,were
[,«'c] (= Millardia meltada) from Bukit, Jalor, Malaysia. subsequently included in Kloss (1911b). Robinson &
This latter record is probably mistaken as the species is Kloss(1911a)describedtwonewtaxa,Musrattusrumpia
currentlyconsideredtoberestrictedtothe Indian Subcon- (=Rattustiomanicus)andMussuriferleonis(=Maxomys
tinent (Musser & Carleton 2005). surifer) from Malaysiaand Singaporerespectivelywhilst
In 1900-1901,anextensivecollectionofmammalswas Robinson & Kloss (1911b) included an additional new
made in the islands ofthe MerguiArchipelago (= Myeik Malaysian murine taxon, Mus muellerifoederis (= Sun-
Archipelago) off the west coast of Tennasserim (= damysmuelleri).Thefollowingyear, Robinson(1912)de-
Tanintharyi Province), Burma (= Myanmar). The speci- scribedafurthertwonewtaxa,Epimyssuriferpemangilis
mens were presented to the United States National Mu- andE. s. aoris, bothofwhich arealsotreatedtodayassyn-
seum. They included many murines, of which Miller onymsofMaxomyssurifer. Thespecimensweredeposit-
(1903a) subsequently described nine as new species. To- ed in the Selangor Museum.
day, all are considered to be synonyms. Three, Mus Miller(1913)publishedafurtherreviewof‘Malaysian’
matthaeus, M. stridulus and M. Incas are referable to mammals. It included fifteennew taxaofmurine rodents
Leopoldamyssabanus MusgilbiventerisreferabletoNi- from peninsular Myanmar, Thailand and Malaysia. Sev-
;
viventer cremoriventer, and Mus luteolus, M. umbridor- en new taxa were published from the Mergui (= Myeik)
sum, M. bentincanus M. casensis and M. domelicus are Archipelago in Myanmar. All are now considered to be
, ,
referable to Maxomys surifer. In the same paper, on the synonyms with three, Epimys vociferans clarae, E. v. in-
basis ofnine specimens from Tioman Island, offthe east sularum. andE. stentor, currentlyreferredtoLeopoldamys
coast ofMalaysia, Miller described one new murid tax- sabanus and four, Rattus rattusfortunatus, R. r. exsul, R.
on, Mus stridens (=Leopoldamyssabanus). r. insulanus, and R. r. dentatus referred to R. tanezumi. A
Meanwhile, Bonhote (1903b) named a new species of further two taxa, Epimys gracilis (marginally extralimi-
rat from Bukit Besar, Jalor, Malaysia, Mus bukit (= Ni- tal to the current study region) and E. lepidus were col-
viventerfulvescens). Inafurtherpaper,basedonacollec- lected from the Tennasserim mainland (= part of
tion from northern peninsularMalaysia, Bonhote 903a) Tanintharyi Division, Myanmar) both are currently con-
(1
recorded an additional fourrat species and described an- sideredtobesynonymsofNiviventerfulvescens. Twotaxa
other three, ofwhich one Mus annandalei (= Rattus an- were described from what istoday Satun Province, Thai-
nandalei) is still recognised as valid. Both Musjaloren- land. One, Epimys solus (= N. cremoriventer), is from
sis (= Rattus tiomanicus) andMusgriseiventer(= Rattus Tarutao Island and the other, E. pannellus (= R. tanezu-
tanezumi) are today treated as synonyms. A new taxon, mi) from Butang Island. Ofthe four taxa from peninsu-
Musjarak, named by Bonhote (1905) from Malaysia, is larMalaysia, three,Epimyrattus vicalana, E. roa, andE.
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
.
Murine rodents ofthe Myanmar-Thai-Malaysian peninsulaand Singapore 19
tingius arenow referred toRattustiomanicus and one, R. AsiatoNewGuinea. Chasen (1937)providedinformation
victor to Sundamys muelleri. on a range ofmurines from the islands offthe west coast
,
Robinson & Kloss (1914) reported on a collection of ofpeninsular Thailand and described six subspecies of
specimens obtained from Thailand and Malaysia. Three Rattus tanezumi, namely: panjius, alangensis, lontaris,
newtaxaofmuridsweredescribedfromKohSamui,Thai- kadanus, moheius, andpipidonis.
land. All are today considered to be synonyms, namely, In Chasen (1940), an extensive work on ‘Malaysian’
Epimysjerdonipan (= Niviventerfulvescens), E. surifer mammals(inreality itincludedtaxafrom Malaysia,Thai-
spurcus (= Maxomys surifer) and E. remotus (= R. an- land and Indonesia), 35 pages were devoted to the murid
damanensis).Thetwonewtaxa frompeninsularMalaysia rodents. Heincludedlistsofspecies,subspecies,andsyn-
wereEpimys orbus (=N.fulvescens) and E. suriferman- onyms. In addition, he named a number ofnew taxa, in-
icalis (=M. surifer). The specimens were donated to the cludingRattusrapitcameroni,whichistodayrecognised
Federated Malay States Museum. asa distinct, endemic species,Niviventercameroni, from
Kloss 1916a) providedadditional data on the distribu- the Cameron Highlands, peninsular Malaysia. Othernew
(
tionoffourrodent taxa,whichtodayarereferredtoMax- taxa included: Rattusrattusrobinsoni(=R. tanezumi);R.
omys surifer Rattus tanezumi, Sundamys muelleri and r. perhentianus and R. r. pemanggis (both = R. tiomani-
Berylmvs bowersii. In two subsequent papers, Kloss cus); R. sabanus dictatorius and R. s. salanga (both =
(1916b) included the description of a new subspecies, Leopoldamyssabanus); andR. suriferpuket, R. s. telibon,
Pithecheirusmelanurusparvus, which iscurrentlyrecog- R. s. muntiaandR. s.pidonis(all=Maxomyssurifer). Sub-
nised as a distinct species, Pithecheirparvus, and Kloss sequently a supplement was published by Ellerman &
(1916c)describedEpimyssurifereclipsis(=Maxomyssu- Morrison-Scott (1955).
rifer). Robinson & Kloss (1916a) reported on six murine Sody (1941) reported on a collection ofrats from the
taxa from Kedah Peak. Indo-Malayan-Australian region, with particular empha-
Meanwhile, inpeninsularMyanmar,Wroughton(1915) sis on murines from Indonesia. The paper included one
provided distribution andecological data fornine murine new taxon from the current study region, Rattus rattus
taxainsouthernTenasserim(=Tanintharyi Division). Hin- chaseni(=R. argentiventer) from Krian, Perak, Malaysia.
ton(1919), in hisextensivepaperon the houserats ofthe Meanwhile Ellerman (1941 )publishedhis monographon
IndianSubcontinentandMyanmar,describedonenewtax- the families and genera ofliving rodents with the murids
on ofmurid rodent, Rattus rattus tikos (= Rattus tanezu- included in volume 2. The exhaustive information con-
mi), also fromTenasserim. Gyldenstolpe (1917) reported tained inthistreatise includedasummaryofgeneric char-
onthemammalscollectedontwo Swedishexpeditionsto acters and a list of subspecies and synonyms, many of
Thailand, including the peninsula and Gyldenstolpe whichoriginated from peninsularMyanmar,Thailandand
(1919)summarisedthemammalswhichhadbeenrecord- Malaysia.
ed from Thailand, including many rodent taxa from the In the 1950s, there was considerable interest in rodent
peninsula. Kloss (1919) included additional localities of behaviour, reproduction, and links to human disease.
Rattus rajah surifer (= Maxomys surifer) from peninsu- Working mostly in peninsular Malaysia, Harrison pub-
lar Myanmar-Thailand and R. r. neglectus (= R. rattus) lished prolifically (Harrison & Traub 1950; Harrison &
from peninsular Thailand. Lindsay (1926) reported on a Lim 1950 and Harrison 1952a, 1952b, 1954a, 1954b,
collectingtrip to the islands ofthe Mergui (= Myeik)Ar- 1955). Harrison (1956a) examined seasonality in rodent
chipelago,which includedrecordsoffivemurinespecies. breeding rhythms and looked at age-weight ratios for
Chasen& Kloss(1931)discussedaspectsofRattustax- eleven murine taxa from Malaysia to determine survival
onomyofspecimensoriginatingfromislands intheStraits rates. He also recorded Bandicota indica from peninsu-
ofMalacca. The paper included the description ofa new lar Malaysia and for the first time reviewed the discrim-
taxon, Rattus rattuspayanus (= R. tiomanicus) from Pu- inating characters between this species and B. bengalen-
lauPaya,offthewestcoastofMalaysia. Kloss(1931)de- sis (Harrison 1956b). Harrison (1957a) lookedathabitats
scribed a new taxon, Rattus canus malaisa (= Lenothrix andmicro-habitatsofvariousmurinerodentsinMalaysia.
canus) from near Kuala Lumpur. Additional papers by Harrison (1957b, 1957c, 1957d,
Chasen (1933) published on Malay Rattus. He includ- 1958, 1961 and 1962) lookedatvariousaspectsofthebe-
eddescriptions,ecological informationanddistributionda- haviour, ecology and parasites ofmurine rodents.
ta from Malaysia and Thailand for three taxa which are Hill (1960) undertook a detailed review ofRobinson’s
today referred to Rattus tiomanicus. R. tanezumi and R. collectionsof‘Malaysian’mammals, which hadbeende-
argentiventer Chasen(1936)reviewedsomenewandex- positedintheNatural History Museum, London’s collec-
isting specimens ofGunomys (= Bandicota bengalensis) tion. Despite the title, these specimens, including many
known from Penang Island, Malaysia. In the same year, rodents, originatednot only from Malaysia but also from
Tate (1936) undertook a taxonomic review ofthe Muri- Thailand, Myanmar and Indonesia. Descriptions and
dae,with astudyareaextendingfrom mainlandSoutheast measurementsofthedifferenttaxawereincluded,numer-
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
-
20 Uraipom Pimsai etal.
ous localities were listed, and four new taxa were de- (specimens subsequently referred to Leopoldamys cilia-
scribed, Rattus rattuspharus, R. r. sribuatensis, R. r. ka tus) and R. sabanus (L. sabanus) from peninsular
banicus, and R. r. terutavensis, all ofwhich are now re- Malaysia.Yong& Dhaliwal(1970)discussedvariationin
ferred to R. tiomanicus. pelage colour in Rattus bowersii (= Beiylmys bowersii)
Dhaliwal (1961), based on Searle & Dhaliwal (1961), from Malaysia. Marshall & Nongngork (1970) studied
reviewed the distribution and habitats offive species of seven rat species on Koh Samui, Thailand with informa-
Rattus (today referred to R. annandalei, R. exulans, R. tion on the discriminating characters and their ecology.
norvegicus, R. tanezumi, andR. tiomanicus)inSingapore Lim (1970) provided information on the ecology, diet,
Island. Subsequently, he discriminated between two Rat- food preferences, and endoparasites of four species of
tus taxa, diardii (= R. tanezumi and jalorensis (= R. murid rodent, Rattus sabanus (=Leopoldamyssabanus),
)
tiomanicus) (Dhaliwal 1962 and 1963). R. muelleri (= Sundamys muelleri), R. bowersii (=
Medway (1964a) undertook areview ofRattus inas (= Beiylmys bowersii) and R. edwardsi (here considered to
Maxomys inas) with informationon its ecology, distribu- be Leopoldamys ciliatus) based on specimens collected
tion and external and cranial characters. He compared throughout much ofpeninsular Malaysia.
specimens from peninsular Malaysia with those from Muul & Lim 1971 provided some valuable informa-
( )
Sabah. In the same year, he published on the marmoset tion on the distribution and ecology offour lesserknown
rat, Hapalomys longicaudatus, from Malaysia (Medway murid rodents, Rattus edwarsi (here considered to be
1964b). Rudd ( 1965) compared the weights of 10 murid Leopoldamys ciliatus), R. annandalei, Lenothrix canus,
taxa in the State of Selangor, examining individual and andPithecheirparvusin Malaysia. Lim etal. (1971)pub-
sexual variation. Harrison (1966) summarised nearly 20 lished a review ofthe small mammals ofPenang Island,
years ofresearch undertaken by the Institute of Medical Malaysia which included information on ten murid taxa.
Research, Kuala Lumpur in his rather informal book on Yong (1971) described a new taxon, Rattus tiomanicus
themammalsofMalaysiaand Singapore. Medway & Lim tenggolensis, from Pulau Tenggol, Malaysia.
(1966) reviewed the taxonomy of three Rattus taxa: Musser(1972) provided a detailed review ofthe genus
tiomanicus,jalorensis and diardii. Hapalomys includinginformationontaxonomy, ecology,
Lim et al. ( 1965) looked for the presence ofparasitic and distribution in Myanmar, Thailand and Malaysia.
nematodes in 14 species of murid rodent in the Kuala Yong etal. (1972) looked atthe karology ofdiardii(= R.
Lumpur area. Lim (1966) studied the predation of mol- tanezumi)andtiomanicusandjalorensis(=R. tiomanicus).
luscs by wild rats since it was considered probable that Musser(1973a) undertookadetailedreviewofRattuscre-
land slugs and snails were intermediate hosts in the life- moriventer(=Niviventercremoriventer)and included in-
cycle ofa nematode, which is the cause ofeosinophilic formation on taxonomy, distribution, and ecology.
meningoencephalitisinman. Medway(1967)providedde- Markvong et al. (1973) studied the karyology ofmurids
tailed informationon thebreedinghabitsofChiropodomys from throughout Thailand and included a rare record of
gliroides based on specimens collected from Selangor Rattus berdmorei (= Beiylmys berdmorei) from peninsu-
State, Malaysia. larThailand.
Yong 1968)providedkaryologicaldataonfourspecies Yong (1973) undertooka study ofthe chromosomes of
(
ofmurines collected from peninsular Malaysia, namely: Chiropodomys gliroides from Malaysia. Lim & Muul
Rattus bowersiiferreocanus (= Beiylmys bowersii), R. (1975) studied the rare arboreal murine, Pithecheir
muelleri validus (=Sundamys muelleri), R. edwardsicil- parvus, in peninsular Malaysia and included information
iatus(=Leopoldamysciliatus)andR. sabanusvociferans on the diagnostic characters, size, preferred habitats, dis-
(= L. sabanus). Yong ( 1969a) and Yong (1969b) includ- tribution, habits ofcaptive specimens, diet, andhelminth
ed the karyology ofR. berdmorei (= B. berdmorei) from infection. Langham & Ming (1976) included the first
Thailandbutwithoutexactlocalitydata.Yong(1969c)re- record ofMus caroli from peninsular Malaysia with in-
ported on rats from Kedah Peak, Malaysia and Harrison formationon thediagnostic charactersandecology. Med-
(1969) reported on the abundance and population densi- way & Yong (1976) studied aspects ofrat systematics in
tyofmammals,includingmurinerodents,inMalayanlow- peninsular Malaysia.
land forests. Marshall (1977b) providedasummaryofthemuridro-
Meanwhile, Medway (1969) published his monograph dentsofThailand forLekagul & McNeely 1977). He in-
(
on ‘The wild mammals ofMalaya’, providing informa- cluded illustrated keys, some synonyms, information on
tion on the distribution, identification, subspecies, habits reproductive status, photographs of live animals and
and reproductionofall mammals from Malaysiaand Sin- skulls, some measurements, short descriptions, distribu-
gapore, including the murine rodents. tion maps, and karyology. In the same year, Marshall
Yong (1970) included data on the external morpholo- 1977a) provided a detailed review ofthe genus Mus in
(
gy, skull anatomy, habitat, breeding behaviour, karyolo- Asia, includingthe Myanmar-Thai-Malaysian peninsula,
gy, serology and immunohaematology ofRattus edwarsi with illustrated keys, information on karyology, distribu-
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
Murine rodents oftheMyanmar-Thai-Malaysian peninsula and Singapore 21
tion, diagnostic characters and ecology. Chan ( 1977) and sular Thailand. Gorog et al. (2004) looked at the phylo-
Chan et al. (1978 and 1979) looked at protein variation geography of three murine species, Maxomys surifer,
and systematics in Malaysian rats. Tweedie (1978) pro- Leopoldamys sabanus, and M. whiteheadi in the Sunda
videdabriefintroductionto therodents ofMalaysiawith shelf. Francis (2008) published a guide to the mammals
a few measurements and some information on habits. ofSouth-eastAsia. Maps and illustrations wereprovided
Musseretal. (1979)undertookadetailedrevisionofthe for 26 ofthe murine species from the study region and
genus Maxomys, with much data relevant to peninsular werefurthersupportedwithbriefinformation ondiagnos-
Myanmar,ThailandandMalaysia. Inthesameyear, Muss- tic characters, measurements, taxonomic notes, distribu-
er (1979) also undertook a detailed study ofthe arboreal tion,andecology(twoadditionalmurinespeciesweredis-
rat genus Chiropodomys, with information on the taxon- cussed but not illustrated).
omy, ecology and distribution. Fain et al. (1980) studied Waengsothorn et al. (2009) included data about the ro-
theparasitic mite faunaofMalaysian rodents. Yong etal. dent specimensheld in the ‘Centre forThai National Ref-
(1982) included information on the karyology of Ha- erence Collections (CTNRC)’. They identified and
palomys and Pithecheir. Musser (1981) published a de- mapped geographical areas ofThailand that have been
tailed taxonomic paperwhich focused on two generaNi- well studied in the past and suggested priority areas for
viventer and Leopoldamys with further comments on fieldresearchinthefuture. Mostrecently, therehavebeen
Lenothrix and Maxomys. a series ofpapers on rodent-borne diseases in Thailand
Abe (1983) reviewedthe taxonomyofNiviventerfrom (Jittapalopong et al. 2008, 201 1; Herbreteau et al. 2012).
Thailand and included bukit in the synonymy ofN.fiul- MeanwhilePagesetal. (2010) lookedatRattiniphyloge-
vescens. In the same year, Musser & Newcomb (1983) ny, using DNA sequence information, to determine
publishedamonographontwogenera,BerylmysandSun- speciesboundaries.Theresultsofthis latterpaperarede-
damys, andalsoprovideddetailedinformationon 12 oth- signed to assist with medical health projects relating to
ergenera known fromthe region, including 5 thatare en- murine rodents. Most recently, Pimsai (2012) undertook
demic to the Sunda Shelf. Langham (1983) studied the a studyofthemurinerodentfaunaofpeninsularThailand
small mammalsofpeninsular Malaysia, includingKedah and Malaysia,which isthe basisforcurrentpaper.Achma-
Peak. di etal. (2013) lookedatthephylogeny,diversityandbio-
Breed & Yong (1986) looked at the spermmorphology geography ofSoutheastAsianMaxomys, including spec-
of 19 taxa of murid rodents from Malaysia. Marshall imens frompeninsularMalaysiaandLatinneetal. (2013b)
(1988) in the second edition of Lekagul & McNeely lookedatthe diversityandphylogenyofMurinaeassoci-
(1988) once again provided taxonomic, ecological, and atedwith limestone karstthroughoutThailand, including
distribution dataon36speciesand 8 generaofmurinero- the peninsula. Pearch et al. (2013) undertook a review of
dents. Boonsong & Felten (1989) described a new small mammal fossil faunas from Thailand including ro-
species ofbandicoot rat, Bandicota bangchakensis (= B. dents from peninsularThailand.
savilei) fromNakhon Si Thammarat, Thailand. (It should
benotedthatthe publicationsoftheThai naturalist Boon-
song Lekagul are sometimes referred to Boonsong and METHODS
sometimes to Lekagul).
Corbet& Hill (1992)publishedataxonomic monograph Species. The taxonomy ofthe 28 species and 12 genera
on the mammal fauna ofSoutheast Asia. They provided followsMusser&Carleton(2005). Differencestothistax-
dichotomousandcharactermatrix keys,acomprehensive onomic opinion are noted in the text and particularly in
list ofsynonyms and subspecies, and distribution maps. the ‘Taxonomic notes’ for each species.
The book includes 12 genera and 23 species ofMurinae Material. Much ofthe data are based on the literature.
from peninsular Myanmar, Thailand and Malaysia and However,additional specimenswerecollectedinThailand
Singapore. Musser& Brothers(1994) undertookadetailed usingcagetrapsandaluminiumlivetraps,whichwere set
review ofthe genus Bandicota in mainland SE Asia, in- both on the ground and in trees. The methodology fol-
cludingtaxonomy,ecologyanddistribution.Threespecies lowedAplin etal. (2003). Thisnewmaterial wasdeposit-
were recognised as occurring in the study area, B. ben- edasvoucherspecimens (dryskins andskulls) in the zo-
galensis (with a very restricted range), B. savilei and B. ological collection ofthe Princess Maha Chakri Sirind-
indica. homNatural HistoryMuseum, PrinceofSongklaUniver-
Chaimanee (1998) produced an invaluable, detailed sity, Thailand (PSUNHM).
monograph ofthe Plio-Pleistocene rodents ofThailand, An additional 165 specimens of 18 species were stud-
which also includedmuch informationrelevant totheex- ied from 65 localities in Thailand and Malaysia (Pimsai
tant species, especially their dentition. Chaimanee & 2012, Figure 18). The vast majority (61) ofthese locali-
Jaeger(2001)subsequentlyreviewedtheevolutionofRat- tieswereinThailand, including 18inpeninsularThailand,
tus and incorporated fossil data originating from penin- with only four localities in peninsular Malaysia. These
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
22 Uraipom Pimsai etal.
muzzle supraorbital
vibrissae crownofhead
baseof
vibrissae/ pinna(ear)
whiskers napeofneck
dorsalsurface/upperparts
cheek
/ midlineofback
' -j
throat
/
lineof
demarcation tail,bicoloured ' tip/terminal/
base/proximal distalendoftail
ventralsurfaci '
underside/ endoftail
underparts hind hindli
A
Fig. 2. (A): External characters ofa murine rodent (Maxomyssurifer), as referred to in the text; (B): arrangementofmammae
inanadultfemalemurinerodent(afterAplinetal. 2003,Figure4.4);(C):thestructureoftheundersideofalefthindfoot(Leopoldamys
sabanus).
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
Murine rodents ofthe Myanmar-Thai-Malaysian peninsula and Singapore 23
specimenswereheldin theCentre forThai National Ref- tion, the dentition ofa youngLenothrix canus is includ-
erence Collections (CTNRC), Thailand Institute of Sci- ed in Fig. 4B, which can be related to the older individ-
entific and Technological Research (TISTR); the Harri- ual in Fig. 18B. It isimportantthatage-relateddifferences
son Institute, UK (HZM); and theNatural History Muse- in tooth morphology are borne in mind when using den-
um, London (BMNH). tal characters for identification purposes.
Descriptions.Genericandspeciesdescriptionsarebased All drawings ofthe skulls and dentition were made by
onpersonal observation(where specimenswereavailable UP with a camera lucida and a stereo-microscope. One
totheauthors)andonthe literature. Literature sources in- specimen ofone species for each genus is illustrated.
cludedoriginal descriptionsofthedifferenttaxaand sub- Measurements. In theoriginal study, Pimsai (2012)took
sequent taxonomic studies and monographs. The princi- a series ofmeasurements from all specimens available to
pal literature sources are listed at the beginning ofeach her in three Thai collections (see Material above). How-
description. ever, this original study was primarily focused on differ-
Generic descriptions are restricted to the characters of ences between genera and was less concerned with spe-
species found within the study area. They do not include cific differences. Forthecurrent study, itwasconsidered
any additional characters ofspecies which are found ex- thatthenumberofspecimensmeasuredwastoosmall and
clusively extralimitally. Ifonly one species in a genus is theirgeographical origin toorestricted forthesemeasure-
found in the study area, the generic description is omit- ments to be considered representative for many of the
ted. species. In addition, for some taxa, such as Niviventer
Common terms used in the description ofthe external cameroni, Maxomysinas, andPithecheirparvus, nomeas-
charactersare illustratedinFig. 2A. Wheredataareavail- urements were available in the initial study. Therefore,
able, the arrangement ofthe mammae ofeach species is measurements included here are from a wide variety of
given, the terminology ofwhich is given in Fig. 2B and literaturesources, includingmanytypedescriptions. Usu-
followsAplin et al. (2003). The structure ofthe hind feet ally more than one set ofmeasurements is included for
foreach genus is illustrated where specimenswereavail- each species. These have been selected to represent dif-
able to the authors with sufficient resolution ofstructure ferentpopulationsfromdifferentgeographical areasand/or
to permit an accurate drawing. The nomenclature ofthe becausethey includeadditionalmeasurementsthatarenot
pads is illustrated in Fig. 2C. The different hairtypes, in- included in the primary sources. All sources are listed in
cluding ‘spines’ and guard hairs are illustrated in Fig. the caption for each Table.
3A-C. The section oftail ofBandicota indica shows the Itisappreciatedthatdifferentresearchersusedifferent
arrangementoftherowsofscales,typical ofmanymurine measuringtechniques. In consequence, it is impossibleto
species(Fig. 3D).AphotographofMaxomyssuriferisin- define measurements so that they exactly correspond to
cludedtoshowaclearlydefineddemarcationontheflanks the method used by all the authors. However, the defini-
between the dorsal and ventral pelage (Fig. 3E); the pho- tionsgiven below followwhatweconsidertobebestprac-
tograph of the Bandicota indica specimen illustrates a tice. For the cranial measurements (Fig. 6), we have pri-
pelage type that has no clear definition between the up- marily followed Musser (1979) and Harrison & Bates
per and lower parts (Fig. 3F). (1991). More measurements are included than those re-
Descriptionsofthecranialanddental charactersforeach ferred todirectly in the currenttext. It is believedthatthe
genusaresupported inthetextbyaseriesof12 drawings. additional definitions will assistthose workingwith sup-
Technicaltermsoffeaturesusedinthegeneric/specificde- portingliteraturethatdoes not includecleardefinitionsof
scriptions are lettered and numbered and linked to the measurements. For the external characters, we have fol-
drawings. Thismethodhasbeenadoptedtoassistall those lowedAplin et al. (2003).
usingthepaperwhoarenotfamiliarwith cranial andden- Headandbodylength (HB): fromthetip ofnoseto the
tal morphology. Toassistfurther, thenomenclatureofthe distal end ofthe anus;
cranial structuresisalso illustrated in Fig. 4Aandtheden- Tail length (T): from the middle ofthe anus to the tip
tal structures in Fig. 4B. ofthe tail (not including hairs);
In general,theskull andteeth ofyoungadultspecimens Ear length (E): taken inside the ear from the bottom of
(where available) were used for illustrative purposes, as the notch tothefurthestpointalongthe rim/margin ofthe
itwas considered thatthese were themostuseful in help- pinna;
ingto identify specimens caught in the field. However, it Hind foot length (HF): fromthe back ofthe heel to the
is appreciated that the morphology ofthe cusp patterns tip ofthe longest toe (not including the claws);
changes through time with young individuals often hav- Mass: whole body mass ofthe specimen (in grams);
ing much more clearly defined cusps than older individ- Greatest skull length (GSL): the greatest antero-poste-
uals. Examples ofthese changes are illustrated in Fig. 5 rior length ofthe skull, taken from the most projecting
forfourdifferentspecies(Berylmvsbowersi, Maxomyssu- point at each extremity, irrespective of what structures
rifer, Rattus tanezumi, and Sundamys muelleri). In addi- form these points;
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
24 Uraipom Pimsai etal.
Fig. 3. Hair and tail ofMurine rodents. (A): short (underfur) and long (overfur) hairs ofRattus tanezumi; (B): spines ofMax-
oofmydsemsaurrciafteri;on(Co):ngtuhearfdlahnakirbseotfweBeannduipcpoetraainndclilcoaw;e(rD)p:elraogwes; Fo:fBsacanldeiscootnatihnedtiacial o-fnBoanldiniecootfadeimndaircccra,tiEo:nMo.nstuhreiffelra-nkclbeeatrwelienne
upperand lowerpelage. Notto scale.
Occipital nasal length (ONL): the distance from the tip Condylobasal length (CBL): from an exoccipital
ofthe nasals to theposteriormargin ofthe occiput(often condyle to the anterior alveolar margin ofthe most for-
butnotalwaysthesameasGLS,especiallyinrodentswith wardly projecting upper incisor tooth;
procumbent incisors);
Bonn zoological Bulletin 63 (1): 15-114 ©ZFMK
