2 
D.J. Kitchener, L.H. Schmitt, P. Strano, A. Wheeler, A. Suyanto 
Figure 1 Locality oi Rhinolophus simplex and R. borneensis specimens used in this study. •, Rhinolophus simplex simplex ; 
■, R. s. parvus ; ★, R. simplex keyensis; ▲, R. simplex subsp. nov.; and A , Rhinolophus borneensis importunus. 
Terminology used in the description of skull, 
dentary and dental (skull) characters and external 
characters follows Hill and Smith (1984). Pelage 
descriptions follow the colour terminology of 
Smithe (1975). 
Adults were diagnosed as those specimens with 
basioccipital and sphenoid bones completely fused 
and epiphyseal swellings absent from metacarpal 
joints. Additionally two adult age classes were 
established based on extent of wear on M 2 
hypocone as follows: young adult, no wear or little 
wear such that the worn area is still elevated above 
the unworn hypocone basin; for adults, worn 
surface area of hypocone below level of unworn 
hypocone basin. 
The effect of sex, adult age classes and taxon on 
skull, dental and external characters was 
investigated by stepwise multiple regressions on 
taxon, sex and age for five taxa. These were 
Rhinolophus megaphyllus (Queensland); R. borneensis 
importunus (Java)-, R. simplex simplex (Bali, Nusa 
Penida, Lombok, Sumbawa, Moyo, Sangeang, 
Rinca, Flores, Lembata, Alor and Sumba); R. 
simplex parvus (Timor) and R. simplex subsp. nov. 
(Savu, Roti and Semau). R. simplex keyensis was not 
included because tire sample size was so small. 
Further, for the three R. simplex subspecies 
considered, the effect of sex, age and island on 
skull dental and external measurements was 
examined using multiple regressions. Examination 
of the residuals from regression analyses gave no 
indication of heteroscedasticity. 
Canonical variate (discriminant) analysis (DFA) 
was computed on skull and external characters 
separately, with males and females combined, 
using the SF’SS PC* program. 
Cellogel electrophoresis of homogenised liver 
was used to investigate genetically determined 
protein variation using the techniques described in 
Richardson et al. (1986). This permitted the 
investigation of variation at 30 presumptive loci. 
Genetic variation was assessed on 101 specimens, 
including some that were juvenile and not included 
in the morphometric analyses. The proteins scored, 
with Enzyme Commission Numbers and Locus 
Symbols in parenthesis, were: aconitate hydratase 
(E.C.4.2.1.3; Acon-1 & Acon-2), adenosine 
deaminase (E.C.3.5.4.4; Ada), carbonate 
dehydratase (E.C.4.2.1.1; Co), diaphorase 
(E.C.1.8.1.4; Dia), enolase (E.C.4.2.1.11; Enol), 
fructose-1, 6-diphosphatase (E.C.3.1.3.11; Fdp), 
fumarate hydratase (E.C.4.2.1.2; Fum), glucose-6- 
phosphate dehydrogenase (E.C.1.1.1.49; G6pd), 
glyceraldehyde-3-phosphate dehydrogenase 
(E.C.1.2.1.12; Gapd), guanine deaminase 
(E.C.3.5.4.3; Gda ), aspartate aminotransferase 
(E.C.2.6.1.1; Got-l and Got-2), a glycerophosphate 
dehydrogenase (E.C.1.1.1.8; a Gpd), glucose- 
phosphate isomerase (E.C.5.3.1.9; Gp-1), isocitrate 
dehydrogenase (E.C.1.1.1.42; ldh-1 and ldh-2), 
lactate dehydrogenase (E.C.l.1.1.27; Ldh-1 and Ldh- 
2), malate dehydrogenase (E.C.l.1.1.37; Mdh-1 and 
Mdh-2), mannose-phosphate isomerase (E.C.5.3.1.8; 
Mpi), purine nucleoside phosphorylase (E.C.2.4.2.1; 
Np), peptidase (E.C.3.4.13.11; Pep-A; E.C.3.4.11.4; 
Pep-B; E.C.3.4.13.11; Pep-Cl and E.C.3.4.13.9 
Pep-D ), 6-phosphogluconate dehydrogenase 
(E.C.1.1.44; 6Pgd), phosphoglucomutase 
(E.C.5.4.2.2; Pgm), superoxide dismutase 
(E.C.1.15.1.1; Sod). 
Chi-square was used to test for significance of 
contingency tables. Tables were reduced when 
more than a quarter of the cells had expected 
values less than 2. When expected numbers were 
small after the tables were reduced to 2 x 2, exact 
probabilities were computed using twice the 
probability of the observed tail. Methods used to 
estimate heterozygosity within populations and 
genetic distances between populations were those 
of Nei (1978). These produce "unbiased" estimates. 
F-statistics were computed by the method of Weir 
and Cockerham (1984), which take into account 
