Records of the Western Australian Museum 18: 179-192 (1996). 
Morphological variation among eastern Indonesian island populations of 
Hipposideros bicolor (Chiroptera: Hipposideridae), 
with descriptions of three new subspecies 
D.J. Kitchener 1 , Y. Konishi 1 and A. Suyanto 2 
1 Western Australian Museum, Francis Street, Perth, Western Australia 6000, Australia 
2 Balitbang Zoologi, LIPI, Jin Ir. H. Juanda 9, Bogor 16002, Indonesia 
Abstract - Hipposideros bicolor (Temminck, 1834) is recorded for the first time 
in the Indonesian Lesser Sunda islands of Sumbawa, Selaru, Sumba, Savu, 
Roti and Timor. Morphological comparisons of cranial, dentary, dental and 
external body characters and univariate and multivariate statistical analyses 
of measurements of these characters were carried out on 76 adult specimens 
from six islands in the Lesser Sundas and, for comparison, with specimens 
from Java and Peninsula Malaysia. 
This study reveals that H. bicolor differentiates morphologically into three 
subspecies, which are described herein. These subspecies, and an 
indeterminate subspecies from Sumbawa island, are closer phenetically to H. 
b. bicolor from Java than to H. b. atrox from Peninsula Malaysia. 
INTRODUCTION 
Hill (1963) characterised the Hipposideros bicolor 
group by the following features: small to moderate 
body size; large broadly rounded ears, normally 
with an internal fold or thickening of the 
membrane of the ear at the antitragal lobe; elongate 
and narrow skull; unspecialised auditory region; 
upper incisors weak with outer lobe obsolescent or 
obsolete. The taxonomy of H. bicolor and closely 
related forms has been confused since the work of 
Andersen (1918). Hill in Corbett and Hill (1992) 
considered the bicolor group in the Indo-Malay 
region comprised 22 species. 
Hill (1963), in his classical review of Hipposideros, 
recognised the following forms of H. bicolor: 
H. bicolor bicolor 
(Temminck, 1834) 
H. b. javanicus Sody, 1937 
H. b. pomona Andersen, 1918 
H. b. gentilis Andersen, 1918 
H. b. sinensis Andersen, 1918 
H. b. atrox Andersen, 1918 
H. b. major Andersen, 1918 
North West Java, 
Banka island 
Central Java 
Southern India 
North India, 
Sikkim, Burma 
South China, 
Hainan, Thailand, 
Indochina 
Malay Peninsula, 
Terutau island, 
Tioman island, 
Sumatra 
Enggano island 
H. b. erigens Lawrence, 1939 Mindoro island 
Philippines 
H. b. macrobullatus Tate, 1941 South Sulawesi 
Hipposideros javanicus was first placed in 
synonymy with H. b. bicolor by Tate (1941) and this 
was later supported by Hill (1983) following his 
examination of an additional Javanese specimen of 
this subspecies. Hill el al. (1986) stated that "the 
forms pomona, gentilis and sinensis were probably 
conspecific". Hill in Corbet and Hill (1992) 
confirmed this and placed gentilis and sinensis as 
subspecies of H. pomona. Hill et al. (1986) agreed 
with Strien (1986) and Bergmans and van Bree 
(1986) that H. macrobullatus was a species. Hill in 
Corbet and Hill (1992) stated that H. b. major may 
be synonymous with H. b. atrox; he considered that 
in Indonesia the forms of H. bicolor were H. b. atrox 
(Nias and Enggano islands, Sumatra) and H. b. 
bicolor (Bangka island, Java, Flores island and 
possibly Timor island and Kangean island (Hill 
and Rozendaal 1989)). With regard to their 
presence on Timor island Goodwin (1979) 
examined three specimens from Tutuala, on the 
northeastern tip of East Timor, which he was 
unable to assign to subspecies. 
Kitchener and Maharadatunkamsi (1995) 
described a unique form of Hipposideros of the H. 
bicolor group from Sumbawa island, based on a 
single skull and two carcasses. They could not 
confidently ascribe it to a species on the material 
available but it was closest to H. bicolor in overall 
