Morphological variation in Pteropus lombocensis 
crown length; M'B, first upper premolar crown 
breadth; FA, forearm length; D2M, digit 2 
metacarpal length; D3M, digit 3 metacarpal length; 
D3P1, digit 3 phalanx 1 length; D3P2, digit 3 
phalanx 2 length; D4M, digit 4 metacarpal length; 
D5M, digit 5 metacarpal length; TIB, tibia length; 
PES, pes length; EAR, ear length. 
The skull, dentary and dental characters were 
measured to 0.01 an accuracy of mm, while the 
external characters were measured to 0.1 mm. 
Pelage descriptions follow the colour terminology 
of Smithe (1975). 
Adults were diagnosed as those specimens with 
the following sutures fused: basioccipital- 
basisphenoid, basisphenoid-presphenoid and 
palatine-maxillary. 
The effect of sex and island on all characters was 
examined by standard multiple regressions (where 
all effects were assessed simultaneously) for those 
islands for which we had specimens of both sex. 
Pantar Island was excluded because we had only 
female specimens from that island. 
Stepwise canonical variate (discriminant 
function) analyses (DFA) were run for skull, 
dentary, dental and external body characters using 
all characters measured, except for tibia length 
[because it had a significant interaction (P<0.01) 
between island and sex], for males and females 
combined. 
STATISTICS: RESULTS AND DISCUSSION 
Univariate statistics 
Mean, standard deviation, minimum and 
maximum values and sample size for each island 
are presented in Table 1 for all characters 
measured. 
Multiple regressions 
Multiple regressions were run for skull, dentary 
and external characters on four islands (Lombok, 
Flores, Lembata and Alor). The results of this 
63 
analysis are presented in Table 2. Because of the 
number of interactions being tested some of these 
tests may be significant by chance alone at 0.05 >P 
>0.01. Consequently, significance levels for this 
analysis were set at P<0.01. 
Sex 
No characters were significantly influenced by 
sex alone, but there was a significant (F, 17 = 6.046; 
P = 0.005) interaction between sex and island for 
tibia length. This interaction resulted from the tibia 
length for the male being much longer than the 
female (56.3 v. 50.4) on Flores Island, whereas on 
the other islands they were subequal in length. 
Island 
Some dental measurements only were 
significantly influenced by island alone. These 
were: C'M 2 length (F 317 = 10.79, Pc.001); C,M 3 
length (F 317 = 44.52, P<.001); P 3 breadth (F 317 = 8.94, 
P = 0.001); and M 1 breadth (F 317 = 12.54, P = 0.001). 
Multivariate analysis 
Because of the absence of any significant 
influence of sex, apart from the significant 
interaction between sex and island for tibia length, 
males and females were combined for all 
characters, except for tibia length, for the following 
DFA. Tibia length was excluded from this analysis. 
The DFA was first run for the remaining 27 
characters for all islands (Lombok, Flores, Lembata, 
Adonara, Pantar and Alor). This analysis extracted 
two significant functions which explained 92% of 
the variance (Figure 2). A total of 86.2% of 
individuals was correctly classified to their 
appropriate island. Three clusters were apparent 
from Figure 2. These were Lombok Island, Alor 
Island and a third group comprising Lembata, 
Flores and Pantar Islands. The DFA run on these 
three above groups, using all 27 characters, again 
extracted two significant functions that explained 
100% of the variance, with 96.5% of individuals 
correctly classified to their appropriate group. Only 
Table 1 (continued) 
c,m 3 
P’L 
P 3 B 
M'L 
M'B 
FA 
D2M 
D3M 
D3P1 
D3P2 
D4M 
D5M 
TIB 
PES 
EAR 
11 
11 
11 
11 
11 
11 
11 
11 
11 
11 
11 
11 
11 
11 
11 
22.69 
3.88 
2.97 
5.13 
2.80 
117.4 
57.8 
80.4 
54.8 
81.8 
80.2 
84.9 
52.5 
29.5 
26.8 
0.29 
0.15 
0.11 
0.28 
0.13 
2.6 
1.7 
2.8 
1.3 
1.7 
2.7 
2.6 
1.3 
0.8 
0.7 
22.19 
3.61 
2.77 
4.68 
2.67 
113.6 
55.3 
76.3 
52.4 
77.7 
75.2 
81.2 
50.1 
28.7 
25.6 
23.20 
4.08 
3.17 
5.51 
3.09 
121.1 
60.5 
85.9 
56.5 
84.4 
85.0 
90.3 
54.5 
31.0 
27.6 
18 
18 
18 
18 
18 
00 
18 
18 
18 
18 
18 
18 
18 
18 
18 
21.09 
3.81 
2.74 
4.82 
2.53 
114.6 
56.2 
78.9 
53.2 
81.9 
78.5 
82.9 
51.2 
28.9 
27.5 
0.54 
0.16 
0.11 
0.33 
0.10 
4.3 
2.2 
2.5 
2.3 
2.6 
2.9 
3.1 
2.3 
1.0 
1.0 
19.28 
3.53 
2.51 
4.20 
2.29 
107.4 
50.9 
73.2 
49.2 
76.3 
72.3 
75.6 
46.4 
26.5 
25.9 
21.87 
4.14 
2.92 
5.36 
2.64 
122.0 
60.1 
83.4 
56.1 
86.2 
83.9 
87.7 
56.3 
30.2 
29.5 
