Small Pteropus from Timor 
149 
Table 1 Canonical Variate Function Coefficients from 
DFA between six taxon ( Pteropus lombocetisis - 
Timor; P. 1. lombocensis; P, 1. heudei; P. g. 
griseus; P. griseus pallidus and P. t. temmincki) 
and based on five cranial characters (see text). 
Standardised values followed by (in brackets) 
unstandardised values. 
Character 
Function 1 
Function 2 
P 3 breadth 
-0.6906 (-1.2756) 
0.5497 (1.0153) 
Rostrum length 
0.7794 (6.6908) 
0.3124 (2.6814) 
Greatest skull length 
0.7440 ( 0.6660) 
0.3350 (0.2998) 
C,M 3 length 
Mesopterygoid 
-0.5234 (-0.8375) 
0.1362 (0.2180) 
fossa breadth 
0.3390 ( 1.1297) 
0.3364 (1.1209) 
Constant 
-23.8467 
-50.7967 
Variance explained (%) 
60.7 
33.0 
unidentified MZB Timor group; P. 1. lombocensis; P. 
1. heudei; P. t. temmincki; P. griseus griseus; and P. g. 
pallidus) and five selected characters (rostrum 
length; P 3 breadth; greatest skull length; C,M 3 
length; and mesopterygoid fossa breadth). This 
DFA extracted four significant functions, which 
combined explained 99.9% of the variation. The 
separation between these six taxon/groups was 
clearly apparent from the plot of Function 1 and 2 
(Figure 2). The unidentified Timor group clearly 
clustered close to P. /. lombocensis and P. I. heudei 
and was a form of P. lombocensis. Function 1, which 
explained 60.7% of the variance, clearly separated 
the P. lombocensis from both P. temmincki and P. 
griseus. The characters loading most heavily (>0.5) 
were P 3 breadth, greatest skull length, rostrum 
length and CjM., length (Table 1). Function 2, which 
explained 33.0% of the variance, clearly separated 
P. temmincki and P. griseus. The character loading 
most heavily (>0.5) on Function 2 was rostrum 
length (Table 1). A total of 93.6% of individuals 
were classified to their correct group/taxon. Two 
of 20 P. I. heudei misclassified to the Timor form of 
P. lombocensis and two of the 10 P. g. griseus 
misclassified to P. g. pallidus. 
The Pteropus lombocensis Group 
The sample of one male and nine female P. 
lombocensis specimens from Timor preclude an 
analysis of sexual dimorphism in this sample. 
Flowever, males and females were combined in 
this sample following the absence of observed 
sexual dimorphism in P. I. lombocensis and P. /. 
heudei by Kitchener el al. (1995a). 
A DFA based on the three forms of P. lombocensis 
(the Timor form, P. I. lombocensis and P. /. heudei - 
including Komodo) and five selected skull and 
dentary measurements (C,M 3 length; M 1 length; 
mesopterygoid fossa breadth; braincase breadth; 
and rostrum length) extracted two very significant 
Functions. Function 1, which explained 84.9% of 
Function 2 
Figure 3 Plot of Functions 1 and 2 from DFA based on 
the three P. lombocensis subspecies (P. /. 
lombocensis, P. 1. heudei and P. /. sallottii) and 
five selected skull characters (see text). Island 
codes as for Figure 2. 
the variance, separated each of these forms (Figure 
3). The characters loading most heavily (>Q.5) on 
Function 1 were C t M 3 length and braincase breadth 
(Table 2). Function 2, which explained 15.1% of the 
variance, partially separated P. 1. heudei from the 
other two forms (Figure 3). The characters loading 
most heavily (>0.5) on Function 2 were 
mesopterygoid fossa breadth, M 1 length and C.M., 
length (Table 2). The DFA allocated all specimens 
to their correct group. 
Summary of the multivariate analysis 
The above analyses indicated that the MZB 
Timor specimens clustered closely in discriminant 
function space with Pteropus lombocensis but were 
Table 2 Canonical Variate Function Coefficients from 
DFA between three groups of P. lombocensis 
[(i) Timor; (ii) Lombok; and (iii) Komodo, 
Flores, Lembata, Pantar, Alor] and based on 
five cranial characters (see text). Standardised 
values followed by (in brackets) 
unstandardised values. 
Character 
Function 1 
Function 2 
C,M 3 length 
1.0847(3.1533) 
-0.6713 (-1.9513) 
Braincase breadth 
0.7276 (1.6337) 
0.2222 (0.4990) 
Mesopterygoid 
fossa breadth 
-0.0025 (0.0080) 
0.8745 (2.8395) 
M' length 
0.1323 ( 0.3851) 
0.8612 (2.5064) 
Rostrum length 
-0.1786 (-0.3192) 
0.4564 (0.8157) 
Constant 
-98.8921 
-11.6914 
Variationexplained(%) 84.9 
15.1 
