THE GUNONG BENOM EXPEDITION 1967 
4. NEW RECORDS OF MALAYAN BATS, WITxH TAXONOMIC 
NOTES AND THE DESCRIPTION OF A NEW PIPISTRELLUS 
By J. E. HILL 
The 1967 Expedition to Gunong Benom, Pahang, 1965 resulted in the accession 
of a number of specimens of bats of especial taxonomic interest, including one 
example that has proved to represent an undescribed species of Pipistrellus. Addi- 
tionally, during recent years, specimens of similar interest or in some cases represent- 
ing species new to Malaya have been received from Lord Medway, formerly of the 
Unit of Zoology, School of Biological Sciences, at the University of Malaya, Kuala 
Lumpur. 
All measurements in this paper are in milhmetres : where these refer to a series 
of specimens the minimum and maximum values are given, followed by the mean 
in parentheses. 
Rhinolophus malayanus Bonhote, 1903 
? B.M. 68.812 (in alcohol, skull extracted). Wang Tangga, Kaki Bukit, Perils, 
6° 39' N, 100° 12' E. 2 January 1968. 
i B.M. 68.816, $9 B.M. 68.813-815 (all in alcohol, skulls extracted). Kisap 
Forest Reserve, Pulau Langkawi, Kedah, 6° 23' N, 99° 52' E. 31 December 1967. 
Rhinolophus malayanus is not listed from the States of Malaya by Chasen (1940 : 
36) and does not appear to have been subsequently recorded. Described originally 
from Biserat, Patani, southern Thailand, its presence in Perils and on Pulau Lang- 
kawi is not unexpected. The species is recorded from Koh Lak, on the east coast 
of southern Thailand rather to the north of Patani by Gyldenstolpe (1916 : 13), 
from the Laos Mountains by Andersen (1905a : 89) and from Laos by Tate and 
Archbold (1939 : 6) and by Phillips (1967 : 634), while Osgood (1932 : 218) records 
it provisionally from Muong Moun, Tonkin (North Vietnam) . 
These specimens from Perils and Langkawi agree closely with the type specimen, 
having the lancet with slightly concave lateral margins and an elongate tip and 
with the lateral margins of the sella almost parallel. The connecting process is 
rounded and rises slightly above the tip of the sella: its point of insertion on the 
24 J- E. HILL 
posterior leaf is a little above the corresponding insertion on the back of the sella. 
The small anterior premolar (pm^) is in the toothrow but much reduced : the second 
lower premolar (pnia) is shghtly extruded or is external to the toothrow, with pma 
and pm4 sometimes almost in contact. 
Measurements (B.M. 68.812, 68.816, 68.813-815, in that order) : length of forearm 
40-3, 39-6, 42-2, 41-2, 40-7; width of sella, 2-2, 2-o, 2-i, 2-o, i-g; greatest length of 
skull i8-2, 18-5, i8-o, 17-9, i8-o; condylocanine length 15-3, 15-5, 15-2, 15-2, 15-1; 
supraorbital length (distance from junction of supraorbital ridges to median anterior 
point of nasal swellings) — , 4-9, 4-9, — , 4-8; width of nasal swellings — , 4-9, 5-1, 
5-1, 5-0; zygomatic width 8-8, 8-9, 8-8, 8-5, 8-8; least interorbital width — , 2-2, 
2 -3, 2 -3, 2 "4, width of braincase 7 -4, 7 -4, 7 -4, 7 -4, 7 -4 ; mastoid width 8 -4, 8 -4, 8 -2, 8 -3, 
8-3; qI-cI 4-4, 4-5, 4-4, 4-2, 43; m^-m^ 67, 6-5, 65, 6-4, 63; c-m^ 6-8, 6-8, 6-6, 
6-5, 6-6; length of mandible 11-3, — , — , — , 11-4; c — nis 6-9, 7-2, 6-9, 6-8, 6-8. 
Rhinolophus malayamis is very similar to R. borneensis, differing chiefly in the 
size and shape of the median anterior nasal swellings, which in malayanus are more 
inflated, higher, wider and more clearly demarcated than in borneensis, extending 
laterally down the sides of the rostrum and thereby reducing the area of the lateral 
sweUings as compared with that species. Robinson and Kloss (1915 : 116) recorded 
as R. borneensis (?) a specimen from Khao Nawng, Bandon, southern Thailand but 
this subsequently became the type specimen of R. robinsoni Andersen (igi8 : 375). 
An examination of the type specimen of R. chaseni Sanborn (1939 : 38) from Con 
Son Island (Pulo Condore), off the southeast coast of Indochina indicates that 
chaseni should be considered a subspecies of R. borneensis rather than related to 
R. malayanus as was suggested by its describer or conspecific with it as thought by 
Phillips (1967 : 634) from the published description. 
Rhinolophus rofeinsoni Andersen, 1918 
A single specimen from Eraser's Hill, Pahang (see below) resembles the nominal 
species robinsoni and klossi in its major features and although in some respects it 
differs slightly from these, in others it connects them. They have never been 
described in detail and rest on brief diagnoses by Andersen in a key pubUshed (on 
behalf of Andersen by Oldfield Thomas) in 1918. The examination of a Umited 
number of specimens suggests that they are conspecific: robinsoni has line priority 
and becomes the specific name. 
The species is relatively small with large, bluntly pointed ears; prominent anti- 
tragal lobe ; large anterior leaf covering the muzzle ; lancet cuneate ; sella wide, its 
central part (excluding the basal lobes) expanded laterally so that its width at this 
point nearly equals or is greater than the width at its base, but sharply constricted 
a little above its central point; upper margin of sella slightly convex; connecting 
process low and rounded, its insertion on the posterior leaf level with or beneath its 
MALAYAN BATS AND A NEW PIPISTRELLUS 25 
insertion on the back of the sella; third, fourth and fifth metacarpals subequal; 
second phalanx of third digit short, its length less than one and one half times the 
length of the first phalanx ; braincase elongate with low sagittal crest ; supraorbital 
region relatively long, the supraorbital length (from junction of supraorbital crests 
to median anterior point of nasal swellings) exceeding the width across the lateral 
anterior nasal swelhngs; a clearly defined frontal depression; prominent nasal 
swellings, the median anterior pair high and globular; mesopterygoid fossa wide with 
deep mesopterygoid pit; anterior upper premolar (pm2) in the toothrow, usually 
in contact with canine and posterior premolar (pm^) ; second lower premolar (pms) 
small, partially or whoUy extruded, with pm2 and pm4 sometimes in contact or 
nearly so. Measurements appear in Tables i and 2. 
The external and cranial features of R. rohinsoni refer it to the ferrumequinmn 
group of Tate and Archbold (1939 : 2) (the simplex group of Andersen (1905a : 75, 
1905b : 648) or inegapkyllus group of that author (1918 : 374). Its wing structure 
places it in either the simplex (more properly the megaphyllus) or horneensis sub- 
groups of Tate and Archboldi and not in the affmis subgroup as these authors 
defined it and in which they listed both rohinsoni and klossi. The long supraorbital 
region of rohinsoni is characteristic of the simplex subgroup but its rounded sella, 
short palatal bridge and extruded second lower premolar ally it more closely to the 
horneensis subgroup and the species is evidently closely related to R. horneensis 
which in many respects it resembles. However, in R. horneensis the sella is narrower 
with its lateral margins parallel or nearly so and the connecting process is inserted 
on the posterior leaf at a point above its insertion on the back of the sella. The 
supraorbital region is shorter and the median anterior nasal swellings in horneensis 
less globular and wider, extending laterally to the sides of the rostrum. 
Rhinolophus rohinsoni rohinsoni Andersen, 1918 
Rli[inolophus] rohinsoni Andersen, 1918 : 375. Khao Nawng. Bandon, Lower Thailand. 
Type specimen B.M. 18.8.2. i (in alcohol (in bad condition), skull extracted). 
Larger than R.r. klossi with slightly narrower lancet; width of sella as in klossi, 
not greater as stated by Andersen; palate wider; mesopterygoid fossa wider anter- 
iorly; sphenoidal bridge very shghtly wider; second lower premolar (pms) completely 
extruded with pm2 and pm4 in contact. 
Rhinolophus rohinsoni klossi Andersen, 1918 
Rh\inolophus] klossi Andersen, 1918 : 375. Pulau Pemanggil, Johore Archipelago, off east 
coast of Malaya. Type specimen B.M. 18.8.5.5 ('ii alcohol, skull extracted). June 1915. 
(?) (J B.M. 67.214 (in alcohol, skuU extracted). Pine Tree Hill, Eraser's Hill, 
Pahang, 3° 43' N, 101° 42' E, at 4500 ft. 10 October 1966. 
1 Tate and Archbold mistakenly noted (p. 5) that the second phalanx of the third digit is unshortened 
in the simplex and horneensis subgroups and shortened in the affinis and feyrutnequinum subgroups: in 
fact, the reverse is true as indicated by Andersen (igo5a). 
26 
J. E. HILL 
Smaller than R.r. robinsoni with lancet slightly wider at its base ; palate narrower ; 
mesopterygoid fossa narrower anteriorly; sphenoidal bridge narrower; second 
premolar (pms) partially or wholly extruded, pm2 and pm4 separated by a narrow 
space or in contact. One example (B.M. 55.1479, from Pulau Pemanggil) has the 
right pms completely extruded but the left pm2 has been lost and pnis in this ramus 
remains wholly in the toothrow. Specimens from Pulau Pemanggil and Pulau 
Aor have relatively small skulls: one specimen from Pulau Tioman has a slightly 
longer forearm and a larger skull approaching the skull of robinsoni in size but 
nevertheless this example retains the narrow palate, narrow mesopterygoid fossa 
and narrow sphenoidal bridge of klossi. 
The specimen from Pahang is referred to klossi with some reservation. Externally, 
it almost exactly resembles this subspecies, but although its skull is similar to that 
of klossi, in size it approaches or in some respects exceeds the skull of robinsoni and 
the supraorbital region is longer and the mesopterygoid fossa narrower than in 
either of these. The material available, however, is totally inadequate for any 
attempt at evaluating the status of the northern and southern populations of 
R. robinsoni on the Malay Peninsula. 
Table i 
External measurements of Rhinolophus robinsoni 
R.r. klossi 
MALAYAN BATS AND A NEW PIPISTRELLUS 
27 
Rhinolophus macrotis dohrni Andersen, 1907 
(J B.M. 67.1595 (in alcohol, skull extracted), $ B.M. 67.1596 (skin and skull). 
Base Camp, Gunong Benom, Pahang, 3° 51' N, 102° 11' E, 700 ft. 20-23 February 
1967. 
(J B.M. 67.1597 (in alcohol). Above Camp 2, Gunong Benom, 2500 ft. 7 April 
1967. 
(JcJ B.M. 67.1598-1598A (in alcohol). Camp 3, Gunong Benom, 3500 ft. 7-8 
March 1967. 
cJcJ PM 45, PM 47-48 (skeletons), B.M. 67.1599 (in alcohol), PM 43 (in alcohol, 
returned to the University of Malaya). Camp 3, Gunong Benom, 3600 ft. lo-ii 
March 1967. 
Rhinolophus macrotis has not previously been recorded from the States of Malaya 
although known from Sumatra {R.m. dohrni) and, following Tate (1943 : 2), from 
Thailand {R.m. siamensis), Fukien [R.m. caldwelli) , Szechuan [R.m. episcopus), Nepal 
{R.m. macrotis) to the Philippines. However, according to Osgood (1932 : 219), 
28 J. E. HILL 
caldivelli and siamensis are apparently sympatric in Tonkin (North Vietnam) but the 
specimens he refers to siamensis are intermediate in size between caldwelli and the 
very much smaller siamensis. Specimens from Gunong Benom agree very closely 
with R.m. dohnii but are very slightly larger: among Malayan Rhinolophus the species 
may be readily recognized by its cuneate lancet, very broad, tongue-shaped sella 
which has a light covering of hairs on its anterior face, and low connecting process 
which originates below the apex of the sella. Cranially it is distinguished by its long 
palatal bridge and by the anterior upper premolar (pm2), which has a prominent, 
inwardly deflected cusp and which stands in the toothrow and is divided from the 
canine and posterior premolar (pm*) by small interspaces. 
Measurements of seven specimens in alcohol and five skuUs (except where stated) : 
length of forearm 44 -5-45 -3 (44-9) ; greatest length of skull ig-i-ig-S (19-4) ; greatest 
length of skull to front of canine 18-5-187 (i8-6); condylocanine length i6-3-i67 
(16-5); basilar length to front of canine I4-6-I4-9 (14-8); width of nasal swellings 
5-0-5 -2 (5-1); zygomatic width 8-3-8-6 (8-5); least interorbital width (four skulls 
2-4-2-6 (2-5); width of braincase 7-5-7-8 (7-6); mastoid width 8-8 — 9-2 (8-9); ci-ci 
4-0-4-3 (4-1); m3-m3 5-7-5-9 (5-8); c-m3 6-6-6-8 (6-7); length of mandible (two 
skulls) II -8; c-m3 6-8-7-0 (6-9). 
Hipposideros lylei Thomas, 1913 
(J9 B.M. 64.831-832 (in alcohol, skulls extracted). Wang Tangga, Kaki Bukit, 
Perils, 6° 39' N, 100° 12' E. 7 March 1964. 
Chasen (1940 : 42) listed H. lylei from the States of Malay, apparently on the basis 
of two skins in the collection of the National Museum, Singapore from Bukit Chint- 
amani, Pahang (Medway, in lift.). Described originally from Thailand, its occurrence 
in the northern part of the States of Malaya is not unexpected. Measurements 
(B.M. 64.831-832): length of forearm 78-2, 75-0 greatest length of skull 30-2, 
29-0; condylobasal length 26-8, 25-5; condylocanine length 26-3, 25-3; rostral 
width 9-4, 9-3; zygomatic width 15-9, 15-9; least interorbital width 4-4, 4-2; width 
of braincase 12-0, 11-9; mastoid width 14-2, 14-1; ci-c^ 7-3, 7-1; m^-m^ ii-o, 10-7; 
c-m3 11-3, ii-o; length of mandible 19-9, 18-9; c-ma 12-4, 12-0. 
Coelops Jrithii Blyih, 1848 
(J B.M. 67.215 (in alcohol, skull extracted but fragmentary). i6th. mile, Ulu 
Gombak, Selangor, 3° 18' N, 101° 43' E. 27 October 1966. 
This specimen constitutes the first definitive record of the species from the States 
of Malaya. Bats of the genus Coelops are rare in collections and its taxonomy seems 
uncertain: Tate (1941a : 5) reviewed it briefly and summarized its history but no 
subsequent work has appeared. The discovery of C. frithii in Malaya has led to an 
MALAYAN BATS AND A NEW PIPISTRELLUS 29 
examination of the material of Coelops in the collections of the British Museum 
(Natural History), which, although quite inadequate for any general revisionary 
treatment, has sufficed to show that two distinct species exist in that country and 
which has suggested the hkely allocation of most of the remaining named forms. 
The Malayan specimen agrees closely with C. frithii which may be characterized 
by: elongate, narrow outline of the lappets projecting from the supplementary 
leaflets flanking the anterior leaf; narial part of the anterior leaf not especially 
depressed and not sharply demarcated from the intermediate part of the leaf, the 
intervening ridges extending from the central process or 'sella' toward the lateral 
margins of the leaf low and indefinite ; posterior part of the upper surface of the 
rostrum sloping anteriorly; rostral inflations low, separated by a shallow sulcus, in 
profile forming only a slight convexity ; a definite pocket or recess within the maxillary 
root of the zygoma, the upper edge of which is shallowly curved anteriorly to ter- 
minate on the rostrum above the anterior face of the second molar; upper tooth- 
rows convergent anteriorly; posterior lower premolar (pm4) elongate and narrow; 
lower molars with prominent, horizontal external cingula. Named forms apparently 
conspecific with frithii are bernsteini Peters, inflatus Miller, &inicus G.M. Allen, 
and formosamis Horikawa. The range of the species extends from Bengal {frithii) 
to southern China {sinicus, i?iflatus), Formosa [formosanus) to Java and Bah [bmi- 
steini: Tate (1941a : 7, 11) records specimens from Bali). Osgood (1932 : 226) 
records inflatus from Tonkin and Annam (North Vietnam) and there are specimens 
from Laos and Thailand in the British Museum (Natural History) which must be 
referred to C. frithii, previously recorded from Thailand by Gyldenstolpe (1916 : 15). 
The Malayan example thus fills a distributional hiatus but insufficient specimens are 
available to justify any attempt at elucidating the validity of subspecies or to 
allocate it at this level. Measurements of this specimen : length of forearm 43-8; 
of tibia 17-0; maxillary toothrow (c-m3) 5-7. 
The type specimen of C. robinsoni Bonhote (1908 : 4) from the foot of Gunong 
Tahan, Pahang differs so markedly from C. frithii as to justify specific separation 
although EUerman and Morrison-Scott (195 1 : 132) considered the possibihty that 
these might be conspecific. Coelops robinsoni may be distinguished from C. frithii 
by: rounded, wide outline of the lappets projecting from the supplementary leaflets; 
narial part of the leaf depressed and sharply demarcated from the intermediate 
leaf by prominent ridges extending from the central process or 'sella' toward the 
lateral margins of the leaf; posterior part of the upper surface of the rostrum nearly 
honzontal and only slightly inclined anteriorly; rostral elevations large, separated 
by a prominent sulcus, in profile forming a distinct convexity; a small foramen but 
no definite pocket or recess within the maxillary root of the zygoma, the upper edge 
of which is sharply curved anterioriy to temiinate on the rostrum above the posterior 
face of the second molar; upper toothrows nearly parallel; posterior lower premolar 
(pm4) wide; lower molars wth low, upwardly curving cingula. 
The genus Chilophylla was described by Miller (1910 : 395) for the species Chilo- 
phylla hirsuta, described concurrently from the Alag River, Mindoro, Philippines 
from a skin without skull. Later, this author (1912 : 117) referred a specimeii 
30 J. E. HILL 
(United states National Museum 175000) from Port Swettenham, Malaya to hirsuta 
and described the cranial characters of the species from its skull. Subsequently, 
Miller (1928 : 85) agreed with an opinion originally advanced by Andersen (in litt.) 
that Chilophylla should be referred to the synonymy of Coelops, but the name has 
nevertheless persisted, Taylor (1934 : 247) using it in his account of the Philippine 
land mammals and Chasen (1940 : 48) including it in the Malaysian fauna, admittedly 
with the reservation that it seemed extremely close to Coelops. There can be no 
doubt from the meticulous descriptions of the skull of U.S.N.M. 175000 by Miller 
(1912 : 117) and Tate (1941a : 5) together with the plate in Howell (1929 : pi. 2, 
fig. 2e) which presumably depicts this skull (as C. robinsoni) that this specimen from 
Port Swettenham represents C. robinsoni. The status of C. hirsuta must remain 
uncertain until the features of its skull are known but its small size (length of 
forearm 33-8), the presence of a curved ridge extending laterally above each nostril 
towards the margin of the horseshoe and the statement by Miller (ibid.) that he 
could detect no external peculiarities that seemed of specific importance between 
hirsuta and the Port Swettenham specimen suggest affinity with C. robinsoni. 
Coelops robinsoni is recorded from Teratau Island, Thailand by Chasen (1940 : 47) 
and from Khao Nawng, Bandon, southern Thailand by Robinson and Kloss (1915 : 
116). The two specimens recorded by Robinson and Kloss are now in the British 
Museum (Natural History) (B.M. 68.605-606, formerly Federated Malay States 
Museum 531/12, 532/12) : both are subadult and their skulls cannot now be found. 
From their external features both are referable to C. frithii, the length of the fore- 
arm in B.M. 68.605 being 39-4 and in B.M. 68.606 38-2 while the measurements of 
the skulls as quoted by Robinson and I\loss are in close agreement with those of that 
species. 
Myotis siligorensis (Horsfield, 1855) 
(J B.M. 67.219 (skin and skull). Cheras cave, Panching, near Kuantan, Pahang, 
3° 53' N, 103° 09' E. 23 January 1967. 
This is the first record of Myotis siligorensis from the States of Malaya. Although 
there are few specimens in collections the species has been recorded from Kumaon, 
Nepal and Sikkim {M.s. siligorensis), Fukien (M.s. sowerbyi Howell), Tonkin (North 
Vietnam) (M.s. alticraniatus Osgood) and Thailand [M.s. thainiis Shamel). From 
descriptions this specimen is similar to alticraniatus or thaianus but in the absence 
of comparative material no definitive subspecific determination can be made. 
The dorsal surface of the specimen is dark brown and the ventral surface of similar 
colour but slightly more buffy. Measurements: length of forearm (from collector) 
29-6; greatest length of skull 12-2; condylobasal length ii-i; condylocanine length 
10-4; zygomatic width 7-0; least interorbital width 3-0; width of braincase 6-o; 
depth of braincase 4-5; mastoid width 6-3; m^-m^ 4-6; length of entire upper 
toothrow 5-1; c-m^ 4-4. Bats of this species may be recognized by their small 
MALAYAN BATS AND A NEW PIPISTRELLUS 31 
size, high, domed braincase and much reduced canines, the lower tooth about equal 
in height to the posterior lower premolar {pm4), with the second lower premolar 
(pma) in the axis of the toothrow. 
Myotis horsfieldii horsfieldii (Temminck, 1840) 
(Jc? B.M. 65.320 (skin and skull), B.M. 65.321 (in alcohol, skull extracted). 
Ampang Reservoir, Kuala Lumpur, Selangor 3° 11' N, ioi°49'E. 15 October 1964. 
Vespertilio horsfieldii Temminck was considered a synonym of Myotis adversus 
(Horsfield, 1824) by Dobson (1878 : 291 et seq) and by Tate (1941b : 551, 558). 
However, Thomas (1910 : 385) identified a series of specimens from Java (which he 
(with Wroughton) had formerly (1909a : 381) determnied as M. hasseltii) as 
M. horsfieldii, pointing out that this species could be distinguished iromM.adversushy 
the attachment of the wing membrane to the metatarsus instead of to the end of the 
tibia as in adversus. It is clear from a drawing now preserved in the British Museum 
(Natural History) copy of Monographies de Mammalogie at Temminck's description 
of horsfieldii that Thomas had submitted illustrations of this feature in his "adversus" 
from Java and in the specimens of "hasseltii" which he had from the same collection 
to F. A. Jentink at the Rijksmuseum van Natuurlijke Historic, Leiden, wherein are 
Temminck's type specimens. Jentink indicates on this drawing that the insertion 
of the wing membrane in one of the "cotypes" of horsfieldii ('c' of his catalogues 
(1887 : 280, 1888 : 186)) is identical with the specimens which Thomas considered 
representative of hasseltii and which he subsequently recognised as horsfieldii. 
Myotis horsfieldii is a relatively small species, the dorsal surface being blackish and 
the ventral surface blackish grey, with narrow, uninflated braincase, narrow inter- 
orbital region, interdental palate terminating anteriorly to a line joining the posterior 
faces of m3-3, long post-palatal extension, the median post-palatal spine supported 
by bony laminae. The second upper premolar (pm^) is large, one quarter to one 
third the crown area of pm^, usually but not always completely intruded from the 
toothrow so that pm^ and pm^ are almost in contact, with pm^ not compressed in the 
row ; second lower premolar (pms) one third or a little more the crown area of pmj, 
separating pm2 and pm4 but usually slightly intruded from the toothrow. The 
species may be distinguished from M. adversus and M. hasseltii by its small size, 
blackish coloration and narrow braincase. These specimens (and one other, $ B.M. 
62.2133, also from Ampang Reservoir, obtained originally by the Institute for Medi- 
cal Research, Kuala Lumpur) agree closely with M.h. horsfieldii and are rather larger 
than M. deignani Shamel, 1942, from Chiengmai, Thailand, which seems likely to be 
a subspecies of M. horsfieldii. 
Measurements (B.M. 65.320-321, 62.2133 (forearm only), in that order) : length of 
forearm 387, 37-8, 36-4, 36-9; greatest length of skull 157, 15-9, 15-2; condylobasal 
length 14-3, 14-2, 137; condylocanine length 13-6, 13-6, 13-1; zygomatic width — , 
32 J. E. HILL 
9-3, — ; least interorbital width 3-6, 3-5, 3-4; width of braincase 7-3, 7-3, 6-9; 
mastoid width 7-9, 7-9, 7-5; ci-ci 4-3, 4-2, — ; m3-m3 6-i, 6-i, 57; c-m3 5-7, 5-9, 5-6. 
Myotis hasseltii (Temminck, 1840) 
? B.M. 65.322 (skin and skull). Kuala Gula, Perak, 4° 52' N, 100° 32' E. 
8 November 1964. 
(J B.M. 68.842, ? B.M. 68.843 (both in alcohol, skulls extracted). Kuah, Pulau 
Langkawi, Kedah 6° 16' N, 99° 50' E. i January 1968. 
$ subadult B.M. 68.844 (in alcohol). Kangar, Perils, 6°27'N, 100° 12' E. 
I January 1968. 
These specimens agree closely with those from Lekop, Karimon Island, Riau 
Archipelago, Indonesia recorded by Thomas and Wroughton (1909b : no) as 
'Myotis adversus (Horsf.) (?)' and subsequently identified by Thomas (1916 : 4) as 
Leuconoe hasseltii, and also with two specimens from Java labelled as originating 
from the Ifijksmuseum van Natuurlijke Historie, Leiden prior to 1844 and thereby 
possibly examined by Temminck. They may be distinguished from specimens 
attributed to the otherwise similar species adversus by their short and not woolly 
pelage, short post-palatal extension lacking thin bony laminae supporting the 
post-palatal spine and in having a minute second upper premolar (pm^) with the 
second lower premolar (pma) very small, usually intruded from the toothrow some- 
times to the extent that pm2 and pm4 are in contact. There exist apparently two 
colour variants : two specimens from Java (both old) are brownish above and below as 
is one from Karimon Island but two others from that island and four from Malaya are 
greyish brown dorsally and paler greyish white or greyish buff ventrally. 
Measurements (B.M. 65.322, B.M. 68.842-843); length of forearm 402, 407, 
41-2; length of foot 8-5, 10. 0, 9-5; greatest length of skull — , 16-3, i6-i; condylo- 
basal length 14-2, 15-0, 14-8; condylocanine length 13-5, 14-3, 14-0; zygomatic 
width — , 10-3, — , least interorbital width 3-8, 4-2, 3-9; width of braincase 7-6, 
8-0, 8.0; mastoid width 8-o, 87, 8-6; c^-ci 4-1, 4-6, 4-5; m^-m^ 6-i, 6-6, 6-6; 
c-m3 57, 6-1, 6'0. 
The named forms of large-footed Myotis (sometimes referred to a subgenus, 
Leuconoe) from southeastern Asia need detailed revision but unfortunately the 
majority seem represented by no more than a few isolated specimens or by a series 
from no more than one or a few localities. There seems Uttle doubt that horsfieldii 
is a distinct species (Medway, 1965 : 60) despite the views of Dobson (1876 : 127, 
1878 : 292) and Tate (1941b : 551, 598) that it is a synonym of adversus. The 
relationship between hasseltii and adversus as here understood is obscure and it may 
be that the advent of further Javan specimens and an examination of the type 
material in Leiden might well alter their taxonomy. Meanwhile, hasseltii is used 
for those bats in which pma is very small and usually intruded or fully intruded from 
the toothrow, adversus for those in which this tooth is larger and stands within the 
MALAYAN BATS AND A NEW PIPISTRELLUS 33 
toothrow or is only slightly intruded, and which have rather long, woolly pelage and 
a long post-palatal extension, its median spine supported by thin bony laminae. 
Pipistrellus javanicus javanicus (Gray, 1838) 
(J (skin and skull) B.M. 71.813). Camp 2, Gunong Benom, Pahang 3° 51' N, 
100° 11' E, 1700 ft. 31 January 1968. 
The small anterior upper premolar is sometimes lacking from the toothrows of 
Pipistrellus (Kuzyakin, 1944 : loi, 1950 : 338 et seq., 1965 : 103) ; on the other hand, 
it may sometimes be present in those of Eptesicus (Hayman, 1954 : 289, 290; 
HiU, 1966 : 303). However, this specimen is noteworthy for the presence of a small 
supernumerary premolar in the right toothrow, situated internally in a shallow 
angle formed by the postero-intemal face of the canine and the antero-intemal 
face of the anterior upper premolar normal to this species, which is situated more 
nearly in the line of the toothrow than the corresponding tooth in the other row. 
The supernumerary tooth is small, with a narrow cingulum and a slender conical 
cusp slightly exceeding the normal anterior premolar in height and just visible 
externally. It is tightly in contact with the canine and the normal anterior pre- 
molar, with a basal area of one half that of this tooth. There are no pecularities in 
the mandibular dentition. 
Pipistrellus ridleyi Thomas, 1898I 
(S B.M. 67.1604 (skin and skull). Base Camp, Gunong Benom, Pahang, 3° 51' N, 
102° 11' E, 700 ft. 16 February 1967. 
This species has been represented hitherto in the collections of the British Museum 
(Natural History) only by the type and paratype specimens, B.M. 98.3.15.5-6, from 
Selangor. The teeth of the example from Gunong Benom are less worn than are 
those of the type specimen and agree closely with those of the paratype, the skull 
of which has now been prepared from the specimen in alcohol. The description 
of the incisors by Thomas is based solely on the type specimen and an examination 
of specimens in which wear is less evident enables some additional features to be 
noted. The inner upper incisor (i^) is low, short and wide, narrow anteriorly but 
wider posteriorly, its length at the cingulum only shghtly exceeding its greatest 
width. The principal cusp is projected forward beyond the premaxilla and is 
directed slightly inwardly rather than anteriorly : in profile its anterior surface is 
strongly convex and its posterior face slightly concave so that it is hooked posteriorly. 
There are low, paired posterior accessory cusps; the labial cusp is smaller than the 
lingual cusp and in the type specimen is almost worn away so that the inner cusp 
becomes readily visible from the side. The outer upper incisor (i^) is short and 
wide, its width exceeding its length, and has a principal cusp hooked posteriorly so 
that the tooth has a caniniform appearance. The principal cusp is equal in height 
to the principal cusp of i^ and is flanked by two lateral accessory cingulum cusps: 
1 Further study of this species indicates that it should be transferred to Myotis (HiU & Topal, in press) . 
34 J. E. HILL 
the lingual cusp is well-developed and prominent but the labial cusp is smaller, 
less obvious and in the type specimen has been worn away. The anterior upper 
premolar (pm^) is large, in the toothrow and separated from pm'* by a narrow 
diastema. The lower incisors are tricuspid and stand in the line of the toothrow: 
as remarked by Thomas, the third is considerably more massive than the other two. 
The anterior lower premolar (pmo) is not greatly reduced and is not extruded from 
the toothrow. 
Measurements of B.M. 67.1604, followed by those of the type and paratype 
specimens: length of forearm 28-8, 28-5, 29-1; greatest length of skull 12-3, 127 
12-3; condylobasal length ii-6, ii-8, 11-5; condylocanine length lo-g, ii-2, ii-i; 
width across anteorbital foramina 3-6, 3-5, 37; zygomatic width — , 8-o, — ; least 
interorbital width 2-9, 3-0, 3-3; width of braincase 6-2, 6-3, 6-2; mastoid width 
67, 67, 6-8; ci-ci 3-4, 3-3, 3-4; m3-m3 5-4; 5-3, 5-3; c-m3 4-4, 4-5. 44; length of 
mandible — , 87, 87; c-ms 47, 4-8, 47. 
Pipistrellus societatis sp. nov. 
Type and only specimen: ^ B.M. 67.1605 (in alcohol, skull extracted). Base 
Camp, Gunong Benom, Pahang, Malaya, 3°5i'N, 102° 11' E, 800 ft. Collected 
15 March 1967 by Lim Boo Liat and Yong Hoi Sen for the Gunong Benom Expedi- 
tion, 1967. Original number BM 225. 
Diagnosis: Allied to and externally closely resembhng Pipistrellus circumdatus 
(Temminck), but differing in smaller size; in greater inflation of the anterior part 
of the braincase; in its shorter, narrower rostrum which is more inflated laterally; 
in the absence of well-defined supraorbital ridges; in its more abruptly curved 
supraorbital margin; in the absence of a deeply excavated frontal depression; 
narrower narial emargination which is not constricted posteriorly; in having a 
shorter post-palatal extension with the pterygoid region widened posteriorly; in 
shallow basial pits which are not sharply excised into the basisphenoid ; in shorter 
toothrows, less massive dentition and much reduced third upper molar, in which 
the metacone and third commissure have been all but lost. 
Description: Externally almost exactly like P. circumdatus but with slightly 
shorter forearm. Dorsal pelage predominantly blackish brown, heavily overlaid by 
orange or bronze tipping, the individual hairs basally brown, blackish brown for 
most of their length and profusely but irregularly tipped with orange or bronze. 
Crown pelage orange tipped but with a small area of straw-coloured, dark brown 
tipped hairs just anterior to the junction of the aural anterior margin with the head, 
a feature not so far noted in circumdatus. Ventral pelage dark brown, the hairs 
tipped with greyish white : there is a small wart bearing a few longer, darker hairs 
medianly situated on the throat just anterior to a Hne joining the angles of the mouth. 
Fur extending a little on to the uropatagium but tibia not especially hairy, the digits 
with a sparse cover of moderate hairs. 
Ears large and bluntly rounded, their anterior margin strongly convex in the 
proximal half, with prominent, posteriorly directed basal lobule. Distal half of 
MALAYAN BATS AND A NEW PIPISTRELLUS 35 
anterior margin almost straight, upper part of posterior margin similarly nearly 
straight but becoming convex with an emargination opposite the base of the tragus 
and terminating in a wide, quadrangular antitragal lobe, its anterior edge standing 
squarely to the margin of the ear and to the cheek, inserted a little behind and below 
the angle of the mouth, forming a small downwardly directed pendent lobule similar 
to that of Chalinolobus gouldi. Tragus large, its sharply concave anterior margin 
terminating in a rounded, anteriorly directed point: posterior margin strongly 
convex, its upper part almost horizontal, terminating basally in a large, rounded 
lobule. Ear margins, especially anteriorly, yellowish white, a feature remarked by 
Temminck in the original account of P. circumdatus and evident to some extent in 
dried specimens. 
Skull short, wide, with rounded, rather globular braincase, inflated and elevated 
in the frontal region, the lambdoid crests not especially prominent and the sagittal 
crest lacking. Interorbital region wide, the supraorbital crests only faintly defined, 
terminating in small tubercles. Frontal depression shallow, with weakly defined 
boundaries extending anteriorly as a very shallow median sulcus, bordered by wide, 
low lateral swellings which contrast with the higher, more ridge-like lateral swellings 
of circumdatus. Rostrum short and wide, inflated laterally above anteorbital 
foramen, the rostrum in frontal aspect having a rounded rather than angular profile 
as in that species. Supraorbital ridges curving abruptly from the interorbital 
region to join the anterior margin of the orbit which forms a flange at the side of the 
rostrum. Narial emargination parallel-sided, almost square posteriorly. Palate 
short with a narrow anterior emargination a little wider than the distance between 
the inner faces of i^"^, extending posteriorly to a line joining the back of the canines. 
Post-palatal extension short, with apparently narrow post-palatal spine, the 
pterygoid wings divergent posteriorly. Basial pits shallow, not sharply excised 
into the basisphenoid. 
Teeth of type specimen heavily worn; inner upper incisor (i^) massive, faintly 
bifid at tip, \vith small posterior cingulum cusp, outer upper incisor (i^) approxi- 
mately one third basal area of i^ with large central cusp flanked by smaller lateral 
cusps, its tip reaching slightly above cingulum of i^, separated from canine by a 
narrow diastema and pushed forward so that the upper incisors lie almost in a 
straight line. Anterior upper premolar (pm^) very small, completely intruded from 
toothrow: in the type specimen pm^ is lacking from the right toothrow but a small 
alveolus can be discerned. Third upper molar (m^) reduced, narrow, platelet-like: 
in the type specimen the complete loss of the third commissure is clearly due to 
wear but it is obvious that this and the metacone are much reduced. Inner (ii) and 
second (12) lower incisors incipiently four-cusped; \z shorter and wider than these, 
tricuspid. Anterior lower premolar (pm2) one fourth crown area of pm4: hypoconid 
and entoconid of m^ slightly reduced. Measurements of P. societatis and P. circum- 
datus appear in Table 3. 
Remarks: This new species is clearly very closely related to P. circumdatus, a 
poorly known species apparently so far represented only by the female type specimen 
from Tapos, Java in the Rijksmuseum van Natuurlijke Historic, Leiden (Jentink, 
36 J. E. HILL 
1887:277, 1888:178); a specimen (B.M. 7.1.1. 401) labelled "Java" from the 
Tomes Collection, now in the British Museum (Natural History) and possibly one 
of the original specimens; an example (B.M. 61.12.10.1) without skull labelled 
"India" and also in the British Museum (Natural History); and from a specimen 
from Pyepat, Upper Burma, recorded by Anthony (1941 : 81) and Tate (1942 : 250) 
and now part of the collection of the American Museum of Natural History. 1 
Through the courtesy of the authorities of the Rijksmuseum van Natuurlijke 
Historic I have been able to examine the type specimen and I am indebted to Dr. 
K. F. Koopman who has arranged the loan of the Burmese specimen from the 
American Museum of Natural History. 
Both P. circimidatus and P. societatis may be readily recognized by their striking 
coloration, the blackish dorsal pelage having characteristic orange or bronze tips. 
The skull of circumdatus, however, does not seem to have been described except in so 
far as Tate (1942 : 250) based his group definition on the skull of A.M.N.H. 1 14850 
from Burma. Cranially, circumdatus is more massive than societatis, its braincase 
less inflated frontally and with a low sagittal crest. The supraorbital ridges are 
prominent, terminating in small tubercles, enclosing a deeply excavated frontal 
depression which extends anteriorly as a shallow rostral sulcus, bound laterally by 
low longitudinal inflations which converge to merge anteriorly. Rostrum longer and 
broader than in societatis, the area above the anteorbital foramen uninflated but 
instead sloping more abruptly from the inflations bordering the rostral sulcus to 
give the rostrum in frontal aspect an angular rather than rounded outline. Supraorbital 
ridges curving less abruptly from the interorbital region than in societatis, the anterior 
margin of the orbit forming a flange as in that species. Narial emargination nar- 
rowed posteriorly with a rounded apex, the anterior palatal emargination equal in 
width to the distance between the inner faces of i^'^ or slightly exceeding it and 
extending posteriorly almost to a line joining the posterior faces of the canines. 
Post-palatal extension slightly longer than in societatis, with narrow post-palatal 
spine, the pterygoid wings parallel: basial pits prominent, deep, sharply excised 
into the basisphenoid. Dentition as described for P. societatis but teeth larger and 
more massive, m^ not reduced but with well-developed metacone and clearly evident 
third commissure, while m^ has the hyopconid and entoconid unreduced. The skull 
of the Burmese example differs from the skulls of the type and second Javan 
specimens in slightly smaller size, less prominent cranial crests, its supraorbital 
ridges terminating in shght tubercles which are barely evident in the Javan examples, 
in a shallower frontal depression, shorter post-palatal extension and narrower post- 
palatal spine, and shghtly less massive dentition. It is possible therefore that 
adequate series of specimens may show the mainland and Javan populations to be 
subspecifically distinct. ^ 
The name of the new species is selected in allusion to the combined effort of the 
several members of differing nationalities and institutions who constituted the 
Gunong Benom Expedition, 1967. 
1 Lord Medway has now obtained (1971) a specimen of P. circumdatus from Fraser's Hill. Pahang. 
It agrees closely with the Burmese example. 
malayan bats and a new pipistrellus 
Table 3 
Measurements of Pipisiyelliis societatis and P. circtimdatus 
37 
Hesperoptenus blanfordi (Dobson, 1877) 
? B.M. 65.345 (in alcohol, skull extracted). Jenka, Temerloh, Pahang, 3° 37' N, 
102° 30' E. Collected and presented by the Earl of Cranbrook. 
Described originally from Tenasserim, this species is relatively rare in collections 
but is recorded by Anderson (1881 : 133) from Johore and by Robinson and Kloss 
(1915 : 116) from Khao Nawng, Bandon, Lower Thailand, while Thomas (1916 : 2) 
lists specimens from the Semangkko Pass, below Eraser's Hill, on the Selangor- 
Pahang boundary; Gunong Tampin, Negri Sembilan; Telok Bahang (= Rahang), 
Penang, and from Kuala Lumpur, Selangor. The specimen listed from Khao 
Nawng by this author is doubtless that previously recorded by Robinson and Kloss. 
The species is easily recognized by its small size (length of forearm 26-28), the 
38 J. E. HILL 
presence of a broad, well-developed pad at the base of the thumb extending to the 
base of the second metacarpal, and by the small size and extreme displacement 
of the outer upper incisor (i^) which is intruded from the toothrow to the extent that 
it lies directly behind the large inner upper incisor (i^), in contact with the posterior 
face of that tooth and with the lingual face of the canine. 
Davis (1962 : 42) recorded a specimen from the Sapagaya Forest Reserve, Sanda- 
kan, Sabah (5°37'N, 118° 04' E) as Hesperopteniis doriae. Through the kindness 
of Dr. J. C. Moore of the Field Museum of Natural History, Chicago, I have been able 
to borrow and examine this specimen, F.M. 77025. As Davis points out, it is a 
juvenile male. However, it is much larger than the adult type specimen of Hesper- 
opteniis doriae and agrees very closely with the type specimen of H. fomesi, described 
by Thomas (1905 : 575) from Malacca, to which it must be referred. Hesperopteniis 
tomesi has been known hitherto only from the type specimen and the example from 
Sabah is therefore the first record for Borneo. It may be recognized by its large size 
(length of forearm c. 50) which is similar to that of the Indian H. tickelli, by its rich, 
dark brown colour, bj' its displaced outer upper incisor and by its strongly imbricated 
lower incisors. Hesperopteniis doriae is smaller (length of forearm 38-0) and is 
(from specimen in spirit) pale brown, with the outer upper incisor not displaced from 
the toothrow and the lower incisors not much imbricated. 
Murina huttoni (Peters, 1872) 
^ B.M. 67.1606 (in edcohol, skull extracted). Near Camp 4, Gunong Benom, 
Pahang, 3° 51' N, 102° 11' E, 4800 ft. 17 March 1967. 
This species establishes the presence of M. huttoni in Malaya, whence M. stiilta, 
M. cydotis and the striking M. aenea have already been recorded (Hill, 1964). 
Although very similar to M. cydotis peninsularis in external appearance, this species 
may be readily distinguished by the features of its molar teeth, the mesostyles of 
mi-2 being more developed than in M.c. peninsularis with the paracones and proto- 
cones less reduced, and especially by the unreduced hypoconids and entoconids 
of mi_2, which are separated from the protoconids and metaconids by a wide trough. 
These features indubitably refer the specimen to huttoni: in cydotis the mesostyles, 
paracones and protocones of mi-2 are reduced and in mi_2 the hypoconids and ento- 
conids are much reduced, sometimes barely evident, and are only narrowly separated 
from the protoconids and metaconids. The degree of reduction is less apparent in 
ma. Subspecific determination of the specimen is less certain : in dorsal coloration 
it resembles M.h. rubella from Fukien, the hairs with bright golden brown sub- 
terminal annulations as in that subspecies. 
Measurements: length of forearm 34-1; greatest length of skull 17-5; condylo- 
basal length 15-6; palatal length 8-5; length orbit-gnathion 4-5; rostral width 5-5; 
zygomatic width 9-4; least interorbital width 4-3; width of braincase j-^; height 
of braincase 6-i; mastoid width 8-o; ci-ci 4-3; m^-m^ 5-5, c-m^ 5-8. 
MALAYAN BATS AND A NEW PIPISTEELLUS 39 
Murina cyclotis peninsularis Hill, 1964 
$ B.M. 67.1607 (skin and skull). Base Camp, Gunong Benom, Pahang, 3° 51' N, 
102° 11' E, 700 ft. 20 February 1967. 
(J B.M. 68.845 (in alcohol, skull extracted). Waterworks Road, Batu Pahat, 
Kangar, Perils, c. 6° 27' N, 100° 12' E. 4 January 1968. 
These specimens are the first examples of this subspecies to be recorded since its 
description (Hill, 1964). Although in some dimensions the specimen from Perils is 
similar to M.c. cyclotis, it agrees with M.c. peninsularis in having the rostrum wide 
anteriorly, with toothrows which are not especially convergent, and in massive 
canines, premolars and molars. Measurements (B.M. 67.1607, B.M. 68.845) : length 
of forearm 38-8, 347; greatest length of skull i8-6, 17-4; condylobasal length 
167, I5-4; length orbit-gnathion 4-5, 4-3; palatal length 9-2, 8-2; rostral width 
6-0, 5-8; zygomatic width 10-9, 9-9; least interorbital width4-5, 4-3; width of brain- 
case 8-i; 8-o; height of braincase 67, 67; mastoid width 9-1, 8-5; ci-c^ 5-1, 4-5 
m3-m3 6-0, 5-6; c-m3 6-2, 5-5. 
Shamel (1942 : 327) recorded two specimens from northern Thailand as M.toxopei, 
described by Thomas (1923 : 254) from the island of Bum in the Moluccas. From 
the description and measurements given by Shamel it seems more probable that 
these specimens represent M. cyclotis. 
Miniopterus schreibersii blepotis (Temminck, 1840) 
(?(J B.M. 67.216-217 (skins and skulls). Pine Tree Hill, Eraser's Hill, Pahang, 
3° 43' N, 101° 42' E, 4250 ft. 12 October 1966. 
cJ B.M. 67.218 (in alcohol). Eraser's Hill, 4750 ft. 22 October 1966. 
Although known to occur from mainland localities as far south as Koh Lak in 
southern Thailand (specimen in British Museum (Natural History)) and from 
Sumatra, Java and Borneo, there seems to be no pubhshed record of M. schreibersii 
from Malaya, whence these specimens now establish it. Measurements of B.M. 
67.216-217: length of forearm 44-8, 50-8; condylobasal length 15-0, 16-3; condylo- 
canine length 14-1, 15-5; zygomatic width 9-0, 9-4; least interorbital width 4-0, 
4-2; width of braincase 8-i, 8-5; m^-mS 6-6, 7-4; c-m^ 6-0, 6-5. The length of 
forearm in B.M. 67.218 is 49-3. The large example (B.M. 67.217) agrees almost 
exactly with a similarly large specimen from Kok Lak, further north on the east 
coast of Lower Thailand. 
Small specimens of M. schreibersii are difficult to distinguish from large examples of 
the sympatric species M. viedius. However, apart from its usually longer fore- 
arm, M. schreibersii differs from medius in longer skull and wider palate : specimens 
from Java, Malaya and Indochina have a length of forearm of 43 -6-50 -8, condylo- 
basal length 147-16-3, condylocanine length 13-8-15 -5 and m3-m3 6-5-7-4 while 
specimens of medius from Java and Pulau Kaban (off the east coast of Johore, 
States of Malaya) have a length of forearm of 40-8-43-1, condylobasal length 13-8- 
14-3, condylocanine length 13-1-13-5 and mS-m^ 5-8-6-3. 
40 J. E. HILL 
Miniopterus tnedius Thomas and Wroughton, 1909 
$ B.M. 67.1610 (skin and skull). Bukit Cheras, Panching, near Kuantan, Pahang, 
3° 53' N, 103° 09' E. 17 Febraary 1967. 
This is a further Malayan record for the species, hitherto known on the mainland 
only from Gunong Pondok, Perak (Chasen, 1926 : 156) but also recorded from 
Pulau Kaban, off the east coast of Johore, and from Pulau Terutau, southern 
Thailand by Thomas (1916 : 4). Measurements: length of forearm 42-2; greatest 
length of skuU 14-4; condylobasal length 13-8; condylocanine length 13-0; zygo- 
matic width — ; least interorbital width 37; width of braincase 77; mastoid width 
8-1 ; ci-ci 4-0; m3-m3 5-9; c-m^ 5-4. Measurements of other specimens from 
eastern Pahang (supplied by Lord Medway) (cJcJ? Medway 66.46-48) length of fore- 
arm (Medway 66.47-48 only) 43; condylobasal length i3-8-i4-o; condylocanine 
length 12 -9-13 -2; m3-m3 5 -8-5 -9. 
SUMMARY 
Taxonomic notes on the more unusual species of bats obtained by the 1967 
Expedition to Gunong Benom are presented in this paper, with similar accounts 
of other specimens of taxonomic interest obtained in recent years in Malaya by 
Lord Medway and his associates. The genus Coelops is reviewed so far as available 
material permits, with the recognition of two valid species, C.frithii and C. robinsoni. 
A new species, PipistreUus societatis, is described from Gunong Benom: Rhinolophus 
malayamts, R. robinsoni (?) klossi, R. macrotis dohrni, Coelops frithii, Myotis sili- 
gorensis and Murina huttoni are added to the bat fauna of the States of Malaya 
while additional specimens of Hipposideros lylei, Myotis horsfieldii horsfieldii, 
Myotis hasseltii, PipistreUus ridleyi, Hesperoptenus blanfordi, Murina cyclotis 
peninsularis, Miniopterus schreibersii blepolis and M. mediiis are recorded. Hesper- 
optenus tomesi is recorded for the first time from Borneo from a specimen hitherto 
thought to be H. doriae and an example of anomalous dentition in PipistreUus 
javanicus is described. 
ACKNOWLEDGEMENTS 
I am indebted to the authorities of the Rijksmuseum van Natuurlijke Historie, 
Leiden, the American Museum of Natural History, New York and the Field Museum 
of Natural History, Chicago, for the loan of specimens essential to this study. 
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99-129. 
J. E. Hill 
Department of Zoology 
British Museum (Natural History) 
Cromwell Road 
London. S.W.7