On some Bats of the Genus Rhinolophus, with Remarks on their Mutual Affinities, and Descriptions of Twenty-six new Forms.
Author
Andersen, Knud
text
Proceedings of the Zoological Society of London
1905
1905-12-31
2
75
145
journal article
http://doi.org/10.5281/zenodo.3757451
d81354de-a914-4582-98be-811adbff11df
3757451
General
Remarks
on
the
Rhinolophus lepidus
Group.
The ancestral species.—
The ancestors of the
simpleX
and
lepidus
groups were very closely related. The latter had a projectingconnecting process, a slightly smaller skull and teeth. But the general
shape of the skull,
the
dentition,
the nose-leaves, apart from the process and a very slight
difference
in the shape of the sella, the ears, the
wing-structure
, the length of the tail, and, we might even say, probably the size, were either identical or extremely similar in both of these extinct Bats.
The place of origin.
—There can scarcely be any doubt that the
lepidus
group originated much farther westwards than the
simpleX
group. If we regard Japan as a continental group of islands, and put aside Java, on account of its peculiar geological history, we still find, not only the most primitive, but in fact
all
the species of the
lepidus
section on the Continent. It is only the section which has spread over the adjacent larger islands, one of which (Sumatra) has comparatively
recently
been continental, while another (Java), probably in a more remote period, seems to have been connected with some part or other of Indo China; and only one form, still so closely related to the Java species as hardly to be specifically different, has found its way so far eastwards as Lombok. The hypothesis, therefore, cannot be called unfounded, that
of the two ancestral species, the ancient
1
simplex
” and the ancient 11
lepidus
,
” the former was Eastern
in range (Austro-Indo-Malayan),
the latter Western
(
Oriental
).
Differentiation*.—
From a systematic point of view I found it convenient to divide the
lepidus
section into three “types
”
; I think that, phylogenetically speaking, there are two only: the
lepidus
and the
minor
type. The former, as coming nearest to
simpleX
in the proportionate size of the skull and teeth, is, probably, the more primitive; it is now distributed over the Indian Peninsula
(
lepidus
),
the Himalayas
(
monticola
),
and
Malacca (
refulgens
).
The latter, the
minor
-type, has spread from the Himalayas (
minor
) eastwards through S. China to Japan
(
cornutus
);
it is represented on the now quite isolated Anambas Islands
(
“
minutus
”);
its
occurrence
in Java is not surprising, considering
*
Compare
the
diagram
on
n.
138.
the faunistic affinities of that island; and it has established itself on the western coast of the Indian Peninsula (
gracilis
).
I have but very little doubt that now, when attention has been called to the differences of all these forms of the minor-type, it will be found also in other parts of the Indian Peninsula.
If any inference can be drawn from fragments of a skull and the external characters, the subbadius-type would appear to be an offshoot of the minor-type: already in
minor
and
cornutus
the process is a little sharper-pointed than in
lepidus
; in
subbadius
and
monoceros
this tendency is carried much further.
The skull of the species of the
acuminatus
section (Java- Lombok, Sumatra-Engano) is of the
lepidus-ty
pe; the process too; the colour remarkably like that of
refulgens
.
This leads me to suppose that
acuminalus
and its allies
(
sumatranus
,
calypso
)
are scarcely more than giant representatives of the lepidus-type.
It is the
subbadius-
type which, from a zoogeographical point of view, is by far the most interesting: it has spread southwestwards over a vast part of the Ethiopian Region, and westwards over the Mediterranean countries:—
(1) The
empusa-type.—
Rh.
empusa
*
and
blasii
have progressed further on the way already indicated by
Rh. subbadius
.
They have the small skull and the small teeth characteristic of
minor-
subbadius
; in the shape of the skull there is no essential difference;
the dentition is identically the same
; the process is that of a
sub badius; the sella is deltoid,
that is: the tendency, in the
subbadius-
sella (as emphasised above), towards assuming a subacute summit has been further developed; and we still see
the constriction at the middle of the sella.
But
empusa
and
blasii
are (as always the Ethiopian and W. Palaearctic species) in several points more highly developed: III.2 is lengthened (about, or more than, 1 the length of III1.); also IV.2 is very much longer (not far from twice the length of
IV
1.).
Rh.
empusa
is, however, an inhabitant of Nyasa- land, far S. of the Equator,
Rh. blasii
of the Mediterranean Subregion; thus, the two extremely closely allied species are now separated by an enormous tract, where no relative appears to occur. As we now know that they are descendants of the Oriental subbadius-type, the explanation seems to be quite clear: one branch spread south west wards, into the Ethiopian Region, and developed into
Rh.
empusa
(slightly more primitive dentition; shorter ears, broader horse-shoe); another westwards into the Mediterranean countries,
Rh. blasii
.
There is an instructive fact connected with these two Bats: I believe them to be comparatively recent intruders into their areas;
Rh.
empusa
is known from one specimen only, from the very
East
of Tropical Africa;
Rh. blasii
is much more common in the
Eastern
Mediterranean tract, and still it does not seem to have reached Spain t.
* Andersen, Ann. &
Mag
. Nat. Hist. (7) xiv. (1904) p. 378 (there is a p. 380: the length of the mandible is 2
’
, not 13*1 mm.).
f
Not recorded in Cabrera Latorre’s “Quirópteros de Espana,
”
Mem. Hist. Nat. ii. (1904). I am also not satisfied that there is any reliable the African coast of the Mediterranean.
misprint
on
Soc. Espan.
record from
(2) The
landeri-euryale
type.
—
The Ethiopian
Rh. landeri
(Fernando Po, Gaboon),
h.
lobatus
(Lower Zambesi to Mombasa), and
Rh.
dobsoni
*
(Kordofan) have the small skull and the small teeth characteristic of
minor-subbadius
; the same shape of the skull;
the same dentition
(no vacillation in the position of p); the process is that of a
subbadius
.
In so far there is no difference at all between this section and the former (
empusa-blasii).
But in the shape of the sella and in a certain peculiarity in the wingstructure they have taken a course of their own:—We have seen, in the
simpleX
group, a progressive development from a sella constricted at the middle, through a parallel-margined stage, to a pandurate sella; we have seen in the
lepidus
group, too, the constricted sella
{minor)
modified into the parallel-margined (
gracilis
); the Ethiopian species
here under consideration
represent the
third and
final stage, the pandtvrate sella.
In addition to this: in all of them IV.1
is peculiarly shortened’,
less than (extremely rarely, as a slight individual atavism, equal to) half the length of IV2. As in
Rh.
empusa
and
blasii
,
III.2 is lengthened.
Rh. euryale
,
from the Mediterranean Subregion, is so extremely closely allied to the above-named Ethiopian species that it shares with them
all
essential characters (even
the highly peculiar shortening of
TV.
1),
with one exception:
it has retained the parallelmargined sella.
Summary.—
When discussing the affinities of the Ethiopian species of the
Rh. simpleX
group (above, pp. 117-20), I arrived at the conclusion that they are undoubtedly derived from Oriental types, and that, most probably, the ancestral species have spread directly from South Asia into the Ethiopian Region.
As will be observed from this, a study of the Ethiopian representatives of the
Rh. lepidus
group leads to quite the same result: they have their closest known allies in the Oriental Region, but they are, without exception, considerably more highly developed than any of their Oriental relatives. Bats of the subbadius- type have evidently spread from some part of South Asia south westwards into the Ethiopian Region
{
empusa
;
landeri
,
lobatus
,
dobsoni
),
and westwards over the Mediterranean countries (
blasii
;
euryale
).
Of all the species of the
Rh. lepidus
group only one has found its
way
to Lower Egypt,
Rh. euryale
.
It is a species exclusively Mediterranean in range, and unusually liable to differentiation into slightly differing local forms I.
Its presence in Lower Egypt is easily explained by invasion from the adjacent Asiatic coast of the Mediterranean, where it is very common (specimens from Lower Egypt are indistinguishable from the Palestine form,
Rh. e. judaicus)
J.
* Thomas, Ann. &
Mag
. Nat. Hist. (7) xiv. (1904) p. 156.
f Andersen andMatschie, “Ueber einige geographische Formender Untergattung
Euryalis”
(SB. Ges. naturi'. Fr. Berlin, 1904, pp. 71-83).
I Although it is beyond the strict limits of the present paper, I propose to insert a few words on the remaining Ethiopian species of the genus:—The
aethiops
section (
Rh. aethiops
, hildebrandti,
and
fumigatus)
are very closely related to the Himalayan
Rh. maorotis,
but much more highly developed in the dentition, the wing-
The probable affinities and phylogeny of the principal forms of the
Rh. lepidus
group are expressed in the subjoined diagram (Ethiopian types marked with an asterisk)