16 
D.J. Kitchener, L.H. Schmitt, P. Strano, A. Wheeler, A. Suyanto 
— R. borneensis 
— Alor 
r- Flores 
- Semau 
_L Roti 
p Lombok 
- Rinca 
- Sumba 
- Lembata 
r— Moyo 
• p Sumbawa 
- Sangeang 
- Timor 
— Sawu 
016 014 012 01 0 008 006 0 04 002 
GENETIC DISTANCE 
-c 
Bali 
Nusa Penida 
Figure 5 Dendrogram computed by UPGMA from the genetic distance (Nei standard distance, unbiased) between 
island populations of Rhinolophus simplex and R. borneensis from Java. 
The characters loading most heavily (>0.6) on 
function 1 and presumed important discriminants 
between R.s. simplex and both R. s. parvus and R.s. 
subsp. nov. are snout to vent length, SV and 
vertical sella height, VSH (Table 5b). Characters 
loading most heavily on function 2 (>0.6) and 
presumed important discriminants between R. s. 
parvus and R.s. subsp. nov. are snout to vent length 
and pes length, PES (Table 5b). 
GENETICS 
The gene frequencies are presented in Table 6. 
Fifteen of the 30 loci scored showed variation 
within islands. The genotype frequencies for all 
occurrences of polymorphism within an island did 
not differ significantly from the Hardy-Weinberg 
expectations. Mean heterozygosity levels are 
presented at the bottom of Table 6. They fall within 
the usual range observed for mammalian 
populations (Nevo, Beiles and Ben-Shlomo 1984). 
Much of the variation within Rhinolophus simplex 
was due to inter-island differences. F-statistics 
revealed four loci that had F ST greater than 0.1. 
These were Acon-2 (0.83), Idh-2 (0.63), Pep-D (0.16) 
and 6Pgd (0.22). The mean F^ over all loci was 0.42 
with bootstrapped 95% confidence limits of 0.09 
and 0.67. However, for most loci, there was little or 
no variability within or between islands and the 
unbiased Nei genetic distances between islands 
was low, being generally less than 0.04 (Table 7). 
These distances, together with those estimated 
from a sample of 18 individuals of R. borneensis 
from Java were subjected to cluster analysis and 
the resultant dendrogram is presented in Figure 5. 
This dendrogram reveals the integrity of R. simplex 
as a species distinct from R. borneensis. 
SYSTEMATICS 
Rhinolophus simplex simplex Andersen, 1905 
Rhinolophus simplex Andersen, K., 1905: 76, PL 3. 
Holotype 
British Museum No. 97.4.18.4, adult female, in 
alcohol, collected June 1896 by A. Everett. 
Type locality 
Lombok I., Nusa Tenggara, altitude 2500 ft ( ca. 
830 m). 
Diagnosis 
Rhinolophus s. simplex differs from both R. simplex 
parvus and R. simplex subsp. nov. in averaging 
larger in all skull, dental and dentary 
measurements, except for the posterior width of 
the sphenoid/pterygoid bridge; SW, and external 
measurements, except pes length, PES, and basal 
sella length, BSL, (Tables la, b). It differs almost 
absolutely from R.s subsp. nov. in tibia length and 
forearm length (see Table 1). IjM,, longer relative to 
outside cochleae width, nasal inflation breadth, 
C'M' length and outer M’M 3 width (Figures 6, 7a, 
b, c, respectively). It differs from R.s. parvus in 
having snout to vent length longer relative to pes 
