Morphological variation in Hipposideros bicolor 
with H. b. atrox of Peninsula Malaysia. It may be 
that the form of H. bicolor on Sumbawa island is a 
further subspecies. 
The morphological differentiation of a number of 
species of micro- and megachiroptera into two or 
three subspecies in the Lesser Sunda islands has 
been documented (Kitchener and Suyanto 1996). 
This morphological differentiation shows some 
patterns, and as summarised by Kitchener and 
Suyanto (1996), "reflect the last major glacial 
(18,000 yr BP) island arrangement (and) suggests 
that many of these morphological changes, 
presumably reflecting evolutionary processes, are 
of relatively recent origin". These authors further 
state that such differentiation to subspecies tends 
to occur in the easternmost islands or on islands in 
the Outer Banda Arc (Sumba, Roti, Savu and 
Timor). Hipposideros bicolor has followed these 
general patterns and differentiated into subspecies 
on the easternmost island of Selaru and on islands 
in the Outer Banda Arc (Timor and Roti/Savu). 
Interestingly, the single specimen from the island 
of Sumba in the Outer Banda Arc, appeared to be 
phenetically closer to the population of H. bicolor 
on Selaru island than to the populations on the 
isolated islands in the Outer Banda Arc of Roti/ 
Savu and Timor. 
Only two studies have examined the association 
between geographic variables and morphology 
among bats in the region (Kitchener et al. 1992, 
Kitchener and Maharadatunkamsi 1996). These 
studies showed that longitude was the most 
important associate in determining overall skull 
and body size. In the former study, Hipposideros 
diadema tended to be smaller towards the east 
contrasting with the latter study on Cynopterus 
nusatenggara which tended to be larger to the east. 
Cynopterus nusatenggara also became larger on more 
isolated islands. The form of H. bicolor on the 
western island of Sumbawa was much larger than 
those forms on islands to the east and south - 
suggesting that this species followed the overall 
size trend of its congener. 
Other specimens examined 
Hipposideros bicolor bicolor 
Kiskenda, Java (7°06'S, 110°16'E); 29 2; WAM 
M(39314, 39360). 
Hipposideros bicolor atrox 
Ampang Impounding, Selangor Peninsula 
Malaysia (3°08'N, 101°46’E); 22 2, IS S', WAM 
M(21158-61). 
ACKNOWLEDGEMENTS 
We gratefully acknowledge the support of the 
191 
Executive Director, Western Australian Museum, 
Mr Andrew Reeves; the Director, Puslitbang 
Biologi (LIP1), Dr Soetikno Wirjoatmodjo, and the 
Director, Balitbang Zoologi (LIPI), Drs M. Amir. 
To our many colleagues who assisted greatly in 
the field, particularly Dr R.A. How and Mr R. 
Johnstone, Western Australian Museum; Ir I. 
Maryanto, Ir Maharadatunkamsi and Bapak 
Boeadi, Balitbang Zoologi, we extend our thanks 
for their efforts and companionship. 
Expedition costs were defrayed by grants to D. 
Kitchener from: National Geographic Society, 
Washington and Australian Nature Conservation 
Agency, Canberra. Some expenses were borne 
personally. Garuda Indonesia kindly defrayed 
freight costs of the expeditions. Mrs N. Cooper, 
Western Australian Museum, ran the computer 
analyses. Mrs S. Dalton, Western Australian 
Museum, typed the manuscript. 
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