Records of the Western Australian Museum 17: 125-142 (1995). 
Systematic review of Nyctimene cephalotes and N. 
Pteropodidae) in the Maluku and Sulawesi regions. 
D.J. Kitchener 1 , W.C. Packer' and A. Suyanto 2 
1 Western Australian Museum, Francis Street, Perth, Western Australia 6000 
2 Museum Zoologicum Bogoriense, LIP1, Jalan Ir. H. Juanda 9, Bogor, Indonesia, 16122 
Abstract - A univariate and multivariate statistical study of the morphology 
of island populations of Nyctimene in the Maluku region, Indonesia, 
distinguishes Nyctimene keasti Kitchener, 1993, from the Tanimbar and Kai 
Islands, as a species. The form from the Tanimbar islands is described as a 
new subspecies of N. keasti; it is not associated with N. cephalotes as 
previously considered in lilt. Nyctimene cephalotes is restricted to Sulawesi 
and the Maluku region. The Sulawesi population of N. cephalotes is described 
as a distinct new subspecies. The Aru population of N. albiventer is 
somewhat differentiated from the Papua New Guinea population of N. a. 
papuanus; its subspecific status is not determined. 
Close island populations of N. c. cephalotes, N. k. keasti and N. k. subsp. 
nov. show considerable morphological differentiation. 
INTRODUCTION 
Kitchener et al. (1993) examined morphological 
variation among populations of Nyctimene 
albiventer (Gray, 1863) in the Maluku region. They 
showed that the population on Aru was 
morphologically similar to the Papua New Guinea 
N. a. papuanus K. Andersen, 1910; while those on 
the Kai and Banda Neira Islands ( N. a. keasti 
Kitchener, 1993) were distinct from both the Aru 
form and the nominate subspecies on Halmahera 
and Temate Islands. 
The discovery of N. a. keasti confuses somewhat 
the previous diagnostic distinction between N. 
albiventer and the closely allied N. cephalotes (Pallas, 
1767). This distinction had been based on the larger 
size of N. cephalotes and the fact that the females 
had a dorsum that was much paler than the males, 
whereas in N. albiventer there was no such colour 
distinction between the sexes (Anderson 1912). A 
number of Nyctimene a. keasti specimens had 
forearm lengths overlapping those of both N. a. 
albiventer and N. cephalotes. Further, female N. a. 
keasti were much paler dorsally than the males. 
In the study region, N. cephalotes is reported from 
the following islands: Ambon (fixed as the type 
locality by Andersen 1912); Seram; Buru, Larat 
Island (Tanimbar group); Timor and Sulawesi 
(Andersen 1912; Hill in Corbet and Hill 1992). The 
records from Larat Island and Timor Island appear 
to devolve on single specimens collected late last 
century (Andersen 1912, Goodwin 1979). 
Between 1987 and 1993 expeditions by staff from 
the Western Australian Museum and Museum 
Zoologicum Bogoriense carried out a survey of the 
terrestrial vertebrate fauna of South Sulawesi, 
Maluku and Nusa Tenggara, Indonesia. On these 
expeditions, series of Nyctimene that would be 
attributed to N. cephalotes by earlier authors, were 
collected from the following islands: Seram; 
Ambon; Buru; Yamdena and Selaru, Tanimbar 
group; and South Sulawesi. This paper reports on 
an examination of these extensive modern 
collections which allow both a re-appraisal of 
morphological variation among island populations 
of N. cephalotes and N. albiventer and a 
reassessment of their taxonomy in the study 
region. 
MATERIALS AND METHODS 
A total of 142 adult specimens (listed in 
specimens examined section) was examined. These 
were from Banda Neira Neira I. (2); Dullah I., Kai 
Kecil (7); Kai Besar I. (5); Wokam I., Aru islands 
(27); the Halmahera group (6); Papua New Guinea 
(13); Ambon (10); Seram (16); Buru (2); Sulawesi 
(9); Selaru (13) and Yamdena (32). The localities of 
these specimens are shown in Figure 1. Apart from 
the Halmahera group specimens (Australian 
Museum, Sydney), all specimens are currently 
lodged in the Western Australian Museum. 
Seventeen measurements of skull, dentary and 
dental characters and six of external body 
characters (all in mm) were recorded from adult 
specimens. 
The measurements recorded were (all 
measurement involving teeth were to alveoli): GSL, 
