Myotis adversus comple: 
209 
11) - the one extracted had a greatest length of 
0.73. 
Pelage colour variable - dorsum ranges from 
Russet to Burnt Umber to Dark Grayish Brown. 
Ventral surface with basal hairs Cinnamon Brown 
or Dark Grayish Brown tipped with Pale Neutral 
Gray, or light Neutral Gray. Patagium and ears 
Russet or Fuscous. 
Distribution 
Queensland and Northern Territory, Australia. 
Etymology 
Named after Mr Greg Richards for his studies on 
the chiropteran fauna of Queensland. 
Myotis macropus (Gould, 1855) 
Vespertilio macropus Gould, 1855: un-numbered 
page of text. 
Syntypes 
Includes BMNH No. 53.10.22.32; skin 
("alcoholic"), skull separate. 
Type locality 
South Australia 
Specimens examined 
See later section. 
Diagnosis 
Myotis macropus differs from Myotis adversus and 
Myotis moluccarum as detailed in the earlier 
diagnoses of these species. 
Description 
The morphology of Myotis macropus is as 
described earlier for Myotis adversus, except for 
differences noted in the earlier diagnosis. Also the 
baculum of the Victorian specimen (C25641) is long 
(0.81), broad and a spatulate type 1 form (Figure 
11) rather than types 2 or 3 typical of M. adversus. 
Distribution 
South Australia and Victoria, Australia. 
DISCUSSION 
Myotis adversus (sensu lato) in the study region 
comprised three species and six subspecies. One 
specimen from New South Wales also appeared 
representative of Myotis adversus and is 
morphologically closest to M. a. tanimbarensis; it is 
not allocated to a subspecies. 
It appears, then, that all three species in this 
species complex occur in Australia - Myotis 
moluccarum (northern Australia), Myotis adversus 
(New South Wales) and Myotis macropus 
(southeastern Australia). 
Myotis adversus is widespread and appears to 
alter little in morphology from Java through Nusa 
Tenggara to Alor Island. This is indicated by the 
fact that the cluster of the large sample of 
specimens from Alor Island in discriminant 
function space incorporates all the other islands in 
Nusa Tenggara (Figure 3a,b). Wetar Island 
(Maluku Tengarra Administrative province), 
immediately to the east of Alor Island, is the first 
population of Myotis adversus in this island drain to 
noticeably diverge morphologically. This 
divergence was considerable, such that the Wetar 
individuals that misclassified to the Alor Group 
based on skull dentary and dental characters, 
comprised only two of 18 individuals, and based 
on external characters, only one of 16 individuals. 
The Yamdena population (Kepulauan Tanimbar) 
further to the east again, and part of the outer 
Banda Arc of islands, also diverged further 
morphologically from the Alor Group of 
populations and also was quite distinct from the 
Wetar population. No individuals in the Yamdena 
population were misclassified to these other 
populations in the DFA. It is probably that this 
morphological divergence of the Wetar and 
Yamdena populations reflects their relative 
geographic isolation from other islands to the west. 
Alor Island, and many of the islands further to the 
west in the Inner Banda Arc, were either joined or 
separated by small water gaps during the last 
glacial maxima about 18,000 yrs BP, when the sea 
level fell by about 120 m. Wetar Island, however, is 
separated by a wide stretch of deep water of ca. 
900 m depth (Indonesian Naval Hydrographic 
Survey maps, 1991) (Kitchener et. al 1990, Heaney 
1991). Yamdena Island, in the Outer Banda Arc, is 
the most geographically isolated M. adversus 
population (except for the New South Wales 
specimen, if it is indeed M. adversus); further it has 
never been connected by dry land to these other 
island populations. However, distance of present 
day water gaps between populations, or closeness 
of past connection during the Pleistocene, are not 
the sole explanation for the morphological 
divergence of the Wetar and Tanimbar 
populations. This is because the Timor and Savu 
populations, which are also isolated 
geographically, both by relative wide current water 
gaps and by deep seas, do not appear to have 
similarly diverged morphologically. Perhaps the 
slightly drier climate of the Wetar and the wetter 
climate of Yamdena Islands compared to the 
islands to the west (Oldeman, 1980) has resulted in 
differential selection pressures on the morphology 
of the population of both Wetar and Yamdena 
Islands. 
The distribution of Myotis adversus extends from 
Java along tire island chain of the Lesser Sundas as 
far as Tanimbar only. Myotis moluccarum occurs on 
