i

4

:i^

i

THE

RHOPALOCERA OF JAVA

DANAIDAE, SATYRIDAE, RAGADIDAE,

ELYMNIADAE

BY

M. C PIEPERS AND P. C T. SNELLEN

With the collaboration of H. Fruhstorfer

WITH 8 COLOURED PLATES

THE HAGUE

MARTINUS NIJHOFF

'913

THE

RHOPALOCERA OF JAVA

\J.3

THE

RHOPALOCERA OF JAVA

DAN AI DAE, SATYR I DAE, RAGADIDAE,

ELYMNIADAE

BY

M. C PIEPERS AND P. C. T. SNELLEN

With the collaboration of H. Fruhstorfer

Si

the HAGUE
MARTINUS NIJHOFF

•9'3

INTRODUCTION.

Whilst I was engaged in the preparation of this, the third volume of my
monograph of the Rhopalocera of Java, my collaborator P. C. T. Snellen
died. The systematic portion of this work, however, had already been completed
by him, so that its continuation need not, therefore, be interrupted on that
account. On the other hand I shall surely miss the aid of his extensive know-
ledge of the Malayan lepidoptera.

Before proceeding to a general discussion of the Rhopalocera, now to be

considered, in the manner followed in the two previous volumes I deem it

necessary to point out that for comparison I have been able to avail myselt

of the work of Prof. Dr. Seitz " Die Gross- Sckmetterlinge der Erde " in which

the Malayo-Australian Danaidae and Satyridae have already been dealt with by

my collaborator H. Fruhstorfer. For although this book, intended chiefly for

identification, differs totally in character from mine, this does not prevent it from

being very useful for my work. I do not, indeed, agree in every respect with its

contents, especially as regards the practice of basing a subspecies on every

slight difference in colour or form of wing in butterflies of the same species

and to apply to such forms distinct names. This method of procedure is at

present much in vogue in the domain of entomology and since I do not follow

this fashion kind critics have pretty clearly expressed the opinion that as I pay

so little attention to local forms I do not, evidently, keep in touch with the

prevalent attitude in this science. The matter, however, is this that this attitude I

conceive to be a fashion or an expression of exaggerated speculation rather than

a serious contribution to science. Where real local forms occur these do not

escape my attention, but a slight difference of colour, even when it occurs in

different localities, does not, in a scientific sense, merit the name of local form.

It is a case of an opinion based on f;Use premises which, in its practical

I

11

expression, has been subjected to much exaggeration and has, consequently,
degenerated into mere speculation. The attitude assumed in this connection is
that the form, and especially the colour pattern of the butterflies, originate,
chiefly if not entirely, through the influence of the environment and become
modified also in connection with it. The nature of this influence — cold or
heat, drought or moisture, light or food, or anything else — is in no way
thoroughly investigated but is left to the fruitful source of the imagination. It
is true much stress is laid, in this connection, on the results obtained from
experiments in which pupae of European Rhopalocera have been subjected
artificially to varying degrees of cold and heat, but conceived in a very super-
ficial manner and interpreted without knowledge of the phenomenon of colour
evolution, indeed without really grasping the evolutional nature of the morpho-
logical changes; these results, consequently, as I have already explained in
detail else-where, are altogether wrongly understood ; in reality the said erro-
neous opinion can find no support in them. Although all these factors may
now and again exercise some influence on the form and colour of butterflies,
as will be considered presently, by far the principal factor in this respect is
their hereditary morphological evolution and, therefore, as regards the colour,
the process of colour evolution. The numerous small differences in individuals
of the same species only represent so many different stages in this evolution
which, of course, in individuals living in isolated localities, may occur somewhat
more strongly developed but without this being the actual result of the difference
of locality. In an article by Krapotkin in the N'ineteenth Century for September
1 90 1 , entitled Recent Science, I find it stated that from a quantity of land snails
collected by Gulick on a mountainous island of the Sandwich Archipelago it
was shown that each valley possesses a peculiar form of these mollusca with
many varieties, and that according to the opinion of Hyait, who investigated
this collection, this proves conclusively that all these forms cannot possibly have
been caused by the climate since this is the same in all these valleys. To
attribute it to difference of locality is neither to be thought of. Between indi-
viduals of one species of one butterflies living together on the same island,
several instances of which 1 have already indicated amongst the Pieridae of
Java, such differences occur likewise; differences between the sexes are even
very common, known as sexual dimorphism although in reality having no
relation to difference in sex as such ; on the other hand the same stage of evolution
may be attained simultaneously by individuals living in different regions and
give rise to the same forms. The expression "local form", therefore, in itself
is not altogether correct and if it is desired to attach to this the scientific
meaning of forms caused principally through local influences it is not applicable

ni

to the differences in question. To designate these as subspecies for that reason
only cannot, therefore, be justified and the creation of all the special desig-
nations is, consequently, usually superfluous. In some cases where species
whose different forms must of necessity be distinguished, as for instance with
regard to the polymorphism in Pap. MEiViNON L., a trinomial system may certainly
be of use even for indicating forms which are not subspecies. But this is
simply a question of practical utility ; if interpreted generally, however, according
to the present fashion — when, for instance, seperate names are assigned to
the numerous forms of Zygaena and Parnassius — it is made to appear as
if a matter of scientific importance were involved whereas, in fact, only confusion
is occasioned. There are already sufficient names.

For this reason I do not desire to keep in touch with this so-called prevalent
attitude of entomology. Quite recently — in Entom. Mitteihingcn, I, 6 —
Prof. C. Emery, of Bologna, has protested against this pseudo-nomenclature
which simply occasions confusion.

Although in working out the Rhopalocera of Java in connection with the
systematic division followed by the late Mr. Snellen, the Danaidae, Satyridae,
Ragadidae, and Elymniadae have been regarded as so many separate families,
I would, however, in accordance with later views in this respect, include the
latter two in the Satyridae, in so far as it concerns my general discussion in
this introduction and, therefore, take into account only two groups, that of the
Danaidae and that of the Satyridae. Both these belong to those Rhopalocera
which BoiSDUVAL assigned to the group siispoisi and are, therefore, fairly distinct
from his group succindi ; this is especially manifest in the earlier stages, not
only with regard to the pupae, to which these designations have reference, but
also to the larvae. The larvae of the IMalayo-Australian Danaidae at least —
those of the African and American forms I am not sufficiently acquainted
with — are characterized by the possession of shorter or longer fleshy processes,
such as do not occur in the larvae of the Papilionidae and Pieridae or of the
Hesperidae. It is true the most ancient forms of larvae in the genus Papilio,
especially the Ornithopiera, do possess fleshy processes but of a totally different
character, being straight and rather stiff in appearance, and not in any way
capable of voluntary movement as, to a certain extent, is the case in the larvae
of Danaidae, evidently as a vestige of a former, much more mobile, condition.
In the larvae of Satyridae spiny processes occur at the head and tail, while in
those of Nymphalidae similar processes are found in various shapes on other parts,
of the body but in the larvae of the families of succindi no trace of them can
be observed. A close relationship between these two large groups, which,

IV

based simply on the venation of the wings, has been postulated some time or
other, does not, therefore, actually exist.

As regards the group of Danaidae, this is of purely tropical origin ; the
few species which occur outside the tropics have, indeed, both in the old and
the new world, spread to the north and the south but their home is, never-
theless, in the tropics ; the tendency of spreading outside their centre of origin
is, indeed, not at all rare in this group ; one of the greatest migrators amongst
the Rhopalocera — Danais, (or Anosia) Plexippus, L. — even belongs to it.
The tropical origin at times manifests itself very stronghly in such colonisers.
Daxais Chrysippus, L. has spread, not only over Africa, and a part of Asia
as well as over nearly the whole of the Malayo-Australian faunal region, but
has also established itself in the extreme south of Europe, in Greece ; but,
whereas, in the tropics it occurs annually in several generations, in Greece it
has only one and occurs there in the imago state only in August, i. e. the
warmest season of the summer. In the tropics the Danaidae are, however,
numerous in the Malayo-AustraUan as well as in the Ethiopian and the Neo-
tropical regions ; evidently, therefore, the Danaidae must have originated under
a tropical climate in the southern hemisphere and must have spread east and
west by means of former land connections ; an adaptation to life in the northern
and southern non-tropical districts, however, has occurred in only a few species.
A ver)' ancient separation has, presumably, led to the independent existence of
the group now found over the whole of South America where, during a very long
period, it has, evidently, developed independently to such an extent that at
present it is scarcely considered to belong to the same family but has been
raised to a separate group under the title of Neotropidae. Nevertheless, some
features of the most ancient Danaidae appear to have been better preserved in
this group than is the case with its allies in the old world, for in the Malay
Archipelago some small butterflies occur possessing these same features and
which for that reason have to be classed with the group Neotropidae although
their home is situate in the region of the Danaidae proper. This can scarcely
be explained otherwise than by regarding them as relics of the original form
which have experienced little or no change and therefore agree to that extent
with the group separated at a remote age and now restricted to South America,
in which likewise some of the original features have been retained. In America,
on the other hand, Danaidae are found in the genera Lycorea and Ituna,
which must have arrived at a more recent period when the peculiar type of
Danaidae had already become developed in the old world, by land connections then

in existence, probably from Africa, and even some Danaidae belonging entirely
to one of the Malayo-Australian forms. To the latter, for instance, belongs the
great traveller Danais (Anosia) Plexippus L.

In the course of ages land connections have presumably existed repeatedly
between South America and Africa and it is very remarkable, therefore, how
the presence in America of butterflies of the Danaid type belonging to different'
groups indicates different periods of migration which becomes explicable only
through the repeated existence of such land connections at different periods.
The oldest of these connections existed presumably at that period when the
continents of America, Africa and Indo-Australia, now terminating southwards
in apices, were connected with the antarctic continent and with each other
consequently, during a tropical climate, a fairly remote period, but during
which the original Danaidae may well have been in existence. The two later
connections may well be presumed to have existed between West Africa and South
America. That the Danaidae have spread from the old world to America, and
not vice versa, appears from the circumstance that not only the Neotropidae but
likewise the others in America have acquired mostly the peculiar elongated
American form of wings, so characteristic of the Heliconidae, but which the
Danaidae of the old world do not possess, a phenomenon occurring also in
butterflies of other orders, as for instance the Pieridae, Acraeidae, and Nym-
phalidae, and in such a manner that this process in these has not advanced
to the same extent and some Pieridae are very clearly in a period of transition
in this respect. This reveals, indeed, a process of change, gradually continuous
and, consequently, where this has already been completed, initiated long ago
which, therefore, must point Hkewise to the existence of an ancient land connection,
although of a more recent date than that from which the Neotropidae resulted.
But some species of Danaidae occurring also in America, as has already been
stated, have all retained the form of wing of the old world, as for instance
Dan.\is (Anosia) Plexippus L. a fact which appears to indicate that they
must have been subjected for a relatively short period to the influences, to be
discussed more fully presently, which cause the form of wing in America to
change. This can only be understood on the assumption of another later
migration and, therefore, of a still later land connection in more recent times.
For the existence of these land connections moreover, there is much probability
on geological grounds.

A great tendency, or rather, adaptability for migrating evidently exists in
this family. One species, Danais (Anosia) Plexippus L., already referred to,
which, like Vanessa Cardui L. in Europe, migrates in large swarms, so-called,
has, like the latter, spread over a large portion of the world. In a similar

VI

manner it is evidently carried off at times by air currents to which it can offer
no resistance and thus travels occasionally eastwards across the whole width of
the Atlantic Ocean to the western coasts of Europe and North Africa ; it has
thus even been observed once to arrive amongst the dunes of Holland across
England and the North Sea. That these are no intentional migrations may
be inferred from the fact that all these individuals perish and are unable to
propagate the species since the food required by the larva does not exist in
Europe. To the westward, however, across the Pacific Ocean, in regions
producing the requisite food, it has spread in this manner to Australia and as
far as China, while other species, such as Danais Chrysippus L., inhabit so
extended an area, from Australia to Africa, Asia, and even South Europe, that
in that case also it must be attributed to an abnormal distribution. Danais
LiMxiACE Cram, likewise occupies a very extensive area, from the east of the
Malayo-Australian Archipelago as far as Africa, and the latter continent cannot be
its home on account of its form which is entirely of the Malayo-Australian type; it
must, therefore, have migrated thither. Over palaearctic western Asia and outside
the tropical part of Australia some species of Danais have likewise spread.

A striking difference is also to be observed in this family with regard to
the manner in which the process of colour evolution operates, especially the
diffusion and recession of black, giving rise to definite groups each character-
ized by a special colour pattern. After the occurrence of the remotest severance,
which gave rise to the separate existence of the Neotropidae, the Danaidae of
the old world followed a course of development of their own in which, however,
the process of colour mutation has moved on various lines, evidently under
special influences entirely unknown to us, and has produced, in this manner,
several groups. A portion of these (Lycorea Doubl., Itura Doubl.), as has
been stated, arrived subsequently in South America, has adopted, the same as
has occurred in the case of the Neotropidae, both the special " South American "
form of wing and the manner in which the process of colour evolution operates
there. In the Danaidae of the old world this process has moved on no less
than three different lines. In the genus Amauris, found in Africa, a peculiar
distribution of black occurs side by side with a fading of the original colour
to a complete white, which must doubtless be ascribed to local influences since
the same phenomenon occurs in African Rhopeilocera of other families, giving
rise to the alleged mimicr)- of species of Amauris, by female forms by Papilio
Dardanus Brown. In the Indo-Australian region, presumably in the western
part, in tracts between Madagascar and India, now covered by the Indian
Ocean — in Madagascar and the groups of islands to the north some forms
of EuPLOEA are still to be found — two lines in the process of colour evolution

Vlt

have been produced, that which occurs in the genus Euploea and that which
is to be observed in the genera Hestia, Ideopsis, and Danais. In Euploea
the black has entirely fused with the original red in such a manner that the
peculiar more or less dark red-brown, characterising this genus, has been
produced ; in some species, however, the red in this colour fusion has remained
so predominant in some spots that much of the original pigment has distinctly
been retained. In some species white dots make their appearance on this red-
brown, in others they become more numerous and larger, increasing to large
spots or even horizontal or vertical streaks; some portions of the wings,
especially along the margins, even become completely white ; this occurs most
markedly in the individuals of species living in the K6 islands, whose members
in other regions have not yet reached such a stage in the process of loss of
pigment. Other Euploeas fade more gradually ; thus E. Browni Godm. Salv.,
occurring in the east of New Ireland [New Mecklenburg], has practically
become entirely white, while of the former brown colour only small patches remain
on the under surface. The production of these white dots is evidently due to
no other cause than that in the process of colour evolution certain of the scales,
originally filled with red pigment, have reached a stage where the pigment,
fading gradually at first, has totally disappeared and in this manner the scale,
filled with air, produces the intense white colour, peculiar to such scales; the
further this process continues and the more the number of scales filled with
air increases, the larger and more numerous of course do the white spots
become. Whereas in some butterflies, as the Pieridae, after the disappearance
of the pigment the wing scales drop, sometimes causing a glassy appearance
at those portions of the wings, this does not appear to be the case in other
butterflies, where the scales, now devoid of pigment, become filled with air,
producing the intense white colour much brighter than the pigmental white of
the Pieridae, which occurs in many butterflies and especially in Danaidae.

It is very important in this connection to observe how these white dots
increase to form larger spots and even white streaks, in which case, therefore,
such spots do not arise from white streaks, as has been assumed in a well
known theory to be a fixed rule; also that in the production of such streaks
and spots in the species of some definite district a certain plan cannot be
denied, which goes to prove that this is also governed by influences peculiar
to the districts in question.

In the genera Hestia, Ideopsis, and Danais, however, this so universal
fusion of the black with the original red pigment has not taken place. In
these black entirely replaces the original colour in some parts, especially in the
apex of the fore wings and is diffused only partially over the remainder of

vnr

the surface of the wings, in some cases, as in D. Chrysippus L., to a very
limited extent but much more in others, such as D. Genutia Cram., following
espially the veins but leaving between them larger or smaller patches of
the original colour in which the fading process in continued. In the latter
species and some others it has become a bright orange and in others again
it becomes lighter in shade, as, for instance, in the Javanese D. Melanippus
Cram., while in a form of this species occurring elsewhere, distinguished
by the name Hegesippus Cram., this orange has become entirely white in the
hind wings.

In a great number of species, of which Danais Juventa Cram, may be
taken as the best known example, the original colour has changed completely
into white which re-appcars everywhere in peculiar streaks and spots through
the black, mostly faded to brown ; reaching the same goal, in fact, by another
road, as effected in Euploeas by the increase of white. In some species
remnants of the original red or its fading into yellow have been retained by
different methods, such as D. Tytia Gray and D. Aspasia F., the latter also
in Ideopsis Chloris Feld. where the black has again been diffused in another
manner as is also the case in Hestia, unless the colour pattern of Hesxia
is to be regarded as consisting of the remnants of the much diffused black
reduced for the greater part by the fading process, which is quite po.ssible.
The unusual size of the butterflies of this genus, indeed, justifies the assumption
that a very ancient form has been preserved in them ; they stand in the same
relation to the other Danaidae as ORNiXHOKrERA to the other Papilios and as
regards the last named it appears from the history of the development of the
larvae, as will be indicated when discussing P. Memxon L., that they have
originated from more ancient forms agreeing with the present day Ornithoptera.
The exceptional size of the Saturnias also supports this view, the grounds
for which, however, cannot for the present be discussed. Not only in mammals,
birds, and reptiles giant forms have existed in former periods of which only a
few relics have been preserved ; in insects this was also the case and the large
forms of Lepidoptera, now extant and constantly decreasing in size, must
similarly be regarded as such relics. In D. Chrysippus L. a species in which
the diffusion of black has been kept within narrow limits, the colour in several
individuals living sometimes, but not invariably, in different districts, exhibits
fading which now and again has become partially white and has thus produced
the imaginary species Dorippus, Alcippus, and Alcippoides.

How is it to explained that the process of colour evolution operates in such
different ways ?

In the first place it should be observed that however much variety may

IX

obtain in this respect, the phenomena occuring in connection with it are evidently
governed by fixed rules.

Not only that in each of these groups the process of change is evidently
in general of the same nature but this may also be observed in detail. Thus,
for instance, all the different white streaks and lines in species such as
D. Agleoides Felder and D. Similis Clerck bear the same character and this
again recurs in those of the 9 of Euploea Midamus L., a species in which
the c? follows the direction of Euploeas while the 9 follows principally that of
the species of Danais just referred to. The slight differences occurring here
are evidently only such deviations from a similar line of evolution which, in
each distinct species, arise from their special morphological peculiarities which
have to be taken into account in the course of the evolutionary process. I
have indicated this clearly in my treatise " Ueber das Horn der Sphingiden-
Raupcn" , which appeared in 1897 in volume XL of the Tijdschrift voor Ento-
mologie, where I explained how the so called horn of the caterpillar of the
Sphingidae, originally a long moveable organ furnished with spines, has gradually
become atrophied ultimately to disappear entirely.

A process, however, which has continued from the original Sphingidae
through all the genera and species which have issued from them and doubdess,
therefore, during several thousants of years until the present day, and which,
as far as known, has hitherto only in one species caused this horn to dis-
appear while the ontogenesis of the existing kinds of caterpillars shows how
this process has ever continued slowly in them all and how, consequently, all
kinds of forms of this organ, in various stages of atrophy, have arisen, all
simply phenomena of the continuation of the same process, the morphological
operation whereof must take into account the peculiarity, occuring correlatively,
of each species. It is, therefore, neither accident nor chaos to which these
differences, arising in the course of the process of colour evolution in Danaidae,
must be attributed ; they are governed by rules although their operation at times
acquires so intricate a character that to unravel it, while not impossible, becomes
extremely difficult.

The same may be observed, for instance, in the development of the white
in the wings of several species of Euploea. One species, very common in
Java, E. Leucostictos Gm., occurs also in many other islands of the Malay
Archipelago where it has received various names as, for instance, E. Viola
Butl. in Celebes. Another species of Java, E. Schlegelii Vol!., is also found
in Celebes where the name E. Gloriosa Butl. has been applied to it. Now
both these forms of Celebes differ from their allies in Java — in the case of
Viola especially at regards the 9 — entirely in the same manner, i. e. in

2

X

both the white spots are spread out into peculiar streaks (see PI. XII, fig. id,
PI. XIII fig. 15^).

This peculiarity, however, is not confined to them since they agree in this
respect with several other species peculiar to that island so that when they
are compared with each other they all clearly exhibit a special Celebes facies
to an expert Lepidopterologist. Similarly when the Euploeas occurring in the
Philippines are compared with the same species from other islands of the
archipelago a special Philippine type can be recognised in them, in this case,
however, rather on account of the size and increased number of the white or
bluish-white spots especially in the apical portion of the fore-wings. The most
notable example of this may be observed in the K6 islands. Three species
of EuPLOEA occurring in Java under the names of E. Leucostictos Gm.,
E. Mazares Moore, and E. Ci imena Cram, are met with in those islands in
the forms called E. Assimilata Felder, E. Hopferi Felder, and E. Eurypon
Hew, all differing from the Javanese forms in the same manner in having a
very broad white margin round both wings, a feature which also occurs in
some other islands, such as Banda, but neither in Ambon, Buru, nor New Guinea.
As regards E. Climena Cram., entirely brown specimens marked with some
white dots (PI. XI fig. \d) and others with partly white hind- wings (PI. XI fig. \B)
occur intermixed in Java, the specimens from Bavvean agreeing with the former
as do those from Ambon, (PI. XI fig. \c) while those from Flores, on the other
hand, have the white on the hind-wings as a part of the Javanese; in this species,
therefore, an evolutional diffusion of white exists which occurs in varying degrees
but reaching its climax in the Ke and some other islands. In the K6 islands
this is, indeed, the case with all these three species and, in addition, with butterflies
of other genera. Hypolimnas Anomala Wall (Antiopa Cram.) is a Nymphalid
of wide distribution in the Malay Archipelago which has assumed the type of
colouring of Euploeas, differing in this respect from the other members of the
genus to which its belongs, and thus, following in the same direction in the
process of colour evolution, beginning to exhibit, in the same manner as
Euploeas, in different districts a larger or smaller number of white spots,
especially on the under side of the hind-wings. In the islands referred to this
species acquires, just like the said Euploea species occurring there, a broad
white margin round all four wings. Moreover in a species of Danais, D. Schenkii
Koch, from the same islands white makes its appearance in a striking manner ;
the large Saturnia occuring there is also much more whitish than its allies in
the other Malayan islands.

On taking these facts into consideration it cannot be doubted that they are
all of the same nature and must, therefore, be attributed to a common origin.

XI

It is difficult, therefore, to assume otherwise than the existence of local influences
causing these changes of colour. When, however, one is familiar with the
nature of the evolutionary changes which cause the modifications of form and
colour, to which Lepidoptera are subject, it will be perceived that in reality
the changes here referred to do not go beyond the acceleration, retardation,
or direction in a certain channel of an evolutionary process of change already
existing ; they evidently agree, therefore, with those which are expressed in so
different a manner in the process of extinction of the horn of the Sphingidae
larvae, already alluded to, and are most probably, therefore, caused in like
manner by the attendant special influences. These are in that case, however,
unmistakably of a local character. Other phenomena exist which can likewise
be only attributed to local influences. Butterflies of the same species in Celebes
and from the Moluccas are frequently considerably larger than those of Java ;
in Flores, on the other hand, they are frequently smaller; why?

We know that adverse surroundings are able to modify in a wrong direction
the hereditary course of development of a child. But although we clearly
discern the existence of such influences we can only demonstrate a portion of
them whereas others, whose presence is equally certain, are beyond our ken ;
these are usually classed together under the general term " antihygienic condi-
tions". It is also known that such influences may be due to local causes and
when their nature is known this may occasionally be demonstrated. To assume
that the hereditary course of development of the forms and colours of butter-
flies also may be guided in another direction by influences proceeding from
their environment, is consequently by no means startling. Only, in order to
grasp this, it should be understood that these changes of form and colour are
no accidental phenomena but the expression of hereditary courses of evolutionary
development. Of this, however, lepidopterologists have as a rule no conception
whatever.

When I protested very strongly just now against the present practice of
attributing every little difference, especially in distribution of colour but also
in form between butterflies of the same species occurring in different districts,
in general to so-called local influences, without being able to adduce any evidence
concerning their nature except fanciful statements about droughts or rainy seasons,
it should not be assumed that I wish to doubt the existence of local influences
as such. With so extremely superficial and unscientific an exaggeration I do
not wish to associate myself; it is, moreover, not the only one of its kind
which may be observed in the speculative treatment of the differences in colour
in Lepidoptera, already alluded to. Some insular melanism, assumed indeed
but certainly not existing, has had to do duty in this connection in turn with

xn

the greater dryness of Eastern Java as compared with the western portion of
that island, an undoubted fact in itself but of no such importance as has been
attributed to it in this manner. Such superficial treatment, however usual it
may be, has, in my opinion, as little in common with serious science as have
the many so-called historical novels of Alexandre Dumas with the actual history
of France, although many people, without scientific education and decidedly
without scientific critical faculty, consider it as such in good faith and judge
by that criterion. It should not, however, be inferred from this that I absolutely
deny that specific influences can bring about such changes. I certainly do not,
for instance, deny that the greater or lesser degree of salinity of sea-water
exercises influence on the morphology of fishes and other marine animals.
Providing these influences are, however, apprehended in a scientific manner
and are understood to be factors which, in the course of the natural evolutionary
processes of form or colour, to which such species are normally subject, act as
stimuli, causing some retardation or acceleration in them or conducting this
course in a specific direction, and providing it be understood that this action on
the individuals is mainly governed by their individual susceptibility which
may vary in a high degree. For susceptibility is a quality, whose nature is
indeed very little known to us, but whose existence cannot be denied. No
stimulus, of whatever nature, can exercise influence unless the susceptibility for
it is present ; where this is not present originally it appears, nevertheless, that
where the individual has been exposed to this stimulus for a period, this quality
may even be generated. The requisite period again varies considerably in
duration and may certainly in some cases be of very great length.

Many experiments have been conducted in which especially pupae of Lepi-
doptera have been exposed for periods of greater or lesser duration to varying
degrees of heat or cold ; the most extensive and important are those carried
out b)' Dr. E. Fischer {Allgemeine Zeitschriji fiir Enlomo/ogie, VI n°. 20) and
latterly by Dr. Harry Federley. It was observed that in consequence the
imagines developed from these pupae exhibited differences in colour and some-
times in form from imagines developed under normal conditions, and owing to
the existing ignorance, to be discussed further presently, with regard to the
nature of the evolutionary changes to which Lepidoptera are subject and
especially concerning the nature of colour evolution, these colour modifications
were assumed to be due to the direct chemico-physical action of the said heat
or cold. The usual superficial reasoning relying upon this would, therefore,
attribute the production of colours in Lepidoptera — and even in animals
generally — to such direct climatic influences and this erroneous conception
has still many adherents. The fashion, alluded to, of referring each deviation

XIII

in colour and form to such climatic influences and of basing thereupon
species and subspecies is principally founded on this conception. With a
better insight owing to the knowledge acquired concerning the phenomenon
of colour evolution I have already in my treatise " Die Farbencvohition {Phy-
loorenk del- Farbeii) bei den Pieriden"', published in 1898, pointed out that
there can be no ([uestion here of direct special chemicophysical action but
merely of a stimulus exercised by the abnormal state of the artificially
produced condition in the course of the evolutionary process of change. But
this made no impression on the prevalent ignorance and when subsequently
Dr. Fischer beean to see that this influence did not act direct but had a
retarding effect on the formation of the colours and consequently produced
phylogenetically more ancient forms, his want of knowledge regarding the phenome-
non of colour evolution confused him sothat he considered both his forms B
obtained by moderate heat or cold, and his forms D produced by frost and
great heat, as having been caused by retardation whereas a more intimate
knowledge of this phenomenon admits only the former as recessive old forms
produced by retardation while it indicates the forms D as prospective forms
stimulated into development by acceleration. The fact, brought to light by
subsequent investigation by Dr. E. Fischer, that the same colour forms may
be produced both by heat and cold, completely decided in this matter. For
when such abnormal action only produces a disturbance in the course of normal
existing processes of nature which, according to circumstances under which it
operates, causes retardation or acceleration, the fact that in specific circum-
stances it may sometimes produce the one or the other cannot cause surprise.
It cannot, however, be reconciled in any way with direct chemico-physical
action. Other disturbing influences have gradually been shown to be equally
capable of producing the same results. At the International Congress of Zoologists
held at Bern in 1904, Dr. M. Countess von Linden stated that by with-
holding oxygen during the pupa stage, and doubtless, therefore, by a morbid
disturbance, she produced the same changes in the colour of the imagines as
had been obtained by Dr. E. Fischer by frost and great heat. My own
observations, to be referred to later, concerning the differences in colour and form
which occur in the same species of butterflies and have been erroneously termed
seasonal varieties, make it extremely probable that the origin of these differences
must be sought in the more nourishing nature of the food which the cater-
pillars consume during the rainy season which, therefore, also acts as a stimulus
promoting the development of such individuals whose susceptibilety enables
them to respond to it. Finally, from the pedigree breeding, carried out at
Batavia by Mr. Jacobson in connection with an account of Papilio Memnon L.,

XIV

also to be discussed presentl}', but already published in the Tijdschyift voor
Enfomo/ogif, it appeared to me that the presumably somewhat abnormal condition
produced in this animal by this nursing, however carefully conducted, had
caused sometimes a noticeable retardation in its colour development and had
thus produced a form, no longer existing in Java but still living normally else-
where, being doubdess more ancient phylogenetically than its forms occurring
in Java at the present day. Cold and heat, withholding oxygen, specially
nourishing food, more or less scanty supplies, and, as we shall see also, abnor-
mally strong illumination or darkness, therefore, may all act in a similar manner.
The different evolutionary processes of change, including that of colour evolution
may in their course be influenced by such stimuli and acquire a slower or more
rapid development.

Nevertheless, as I have already stated, the prevalent ignorance retains this atti-
tude and one finds repeatedly, especially in Germany, the influence of cold and heat
— alleged to have been proved by these experiments with butterflies — adduced
on all occasions. I have even met with it in the domain of ethnology. Prof.
Dr. Standfuss regarded the change thus brought about in these butterflies as
newly acquired characters ; when he succeeded in inducing them to pair and
in raising in this manner a second generation, corresponding in colour to the
parents, he saw in this a proof of the heridity of the acquired characters!
These butterflies, however, had simply, as regards their colour evolution, reverted
to an older stage of development. Being at this stage they naturally produced —
in like manner as their ancestors had done while at that stage — a progeny
which corresponded to it. This contained indeed ample proof of the fact that
they had by no means acquired new characters by the experiment but had
simply reverted to an earlier stage of development.

The direction assumed by these processes must, therefore, also be governed
by such stimuli, even though their nature is as yet unknown to us. That
specific directions of this nature do exist, observation teaches us ; the fact also
emerges that where they occur they are not restricted to definite genera but
make their appearance in animals otherwise different but living in the same
district, sometimes, indeed, in whole genera, but also in a few species or even
in one or other of the sexes of these. Where whole genera have assumed
such a specific direction it sometimes appears, as is the case with the more
ancient American Danaidae, that these are the very ones, if local influences
are assumed to operate here, which have been exposed to them during the
greatest period ; where, however, only a few species or sexes have assumed a
specific direction, this points unmistakably to a difference in susceptibility which
has not yet been developed equally in all individuals. In those butterflies,

XV

where this occurs, as In the American Dismorphias and the Malayo-Australian
EuPLOEA MiDAMUS L. it is clearly a manifestation of transition from an older
condition in this new direction.

These are the local influences, whose action manifests itself in accordance
with the individual susceptibility, which must be kept in mind when there is
question of the above mentioned accelerated development which produces the
alleged seasonal varieties.

It is worthy ot being noted here how little variation occurs in migratory
butterflies distributed over a great part of the world, such as Anosia (Danais)
Plexippus L., Pyrameis Cahdui L., or Lycaena Boeticus L. ; would not this
point to a very slight susceptibility to foreign influences in these species, thereby
explaining how they can adapt themselves in this manner to different climates ?

With regard to the peculiar elongated form of wing of the South American
Rhopalocera, so characteristic of the family Heuconidae, which is confined to
that part of the world, I have already alluded to the fact that this form of
wing has been acquired not only by the Neotropidae, a form of Danaid
established there for a very long period, but likewise by the progenitors, which
must have arrived from Africa much later, of the present genera Lycorea and
Ituna, and by several of the later arrivals amongst the Acraeidae, Pieridae,
and Nymphaltdae. By these last, however, by no means so generally, sothat
some species of the Pierid genus Dismorphia are evidently still in a period
of transition which is shown by the fact that in only one of the sexes this
change of wing form has commenced. While the Danaidae which must certainly
have reached America last, to which Danais (Anosia) Plexippus L. belongs,
have retained their original form of wing entirely unchanged. W^here this
process of change of form occurs in this manner it can hardly be explained
otherwise than that it is caused by the difference in individual susceptibility;
that only during a long continued existence under specific uniform conditions
this increases to such an extent that the form of wing in all the species becomes
thus modified but with an existence of shorter duration does not thus increase
in all the species and this modification is, consequently, produced only in
certain species and sometimes even only in one of the sexes of such species,
while with a still shorter existence it may not occur at all.

The peculiar modification in the form of wing alluded to must, therefore, be
considered as the response to a stimulus, directing the evolution of the form
of wing in a specific course. A stimulus whose character we may be unable
to define as yet but which is evidently restricted to South America, although
the region where its influence is felt is of very great extent.

With regard to the evolution of pigmental colours the same phenomenon

Xvt

may be observed in the old world. As we have already noted, in the African
Danaid genus Amauris this follows the same course which characterizes the
African Papilio Dakdanus Brown. Still more striking is the arrangement of
the pigments producing the peculiar colour of Euploeas. But while this
character occurs in the whole of this specialized genus the same colouring
appears in a few species of various genera of Rhopalocera, living in the same
districts with them, in various degrees of resemblance, sometimes completely
as in Euploea's, at other times in a lesser degree, or even in one sex only,
as is the case with Dimorphias in South America, whereas in the others this
course of the evolution of colours has not, or at least not yet, been followed.
Hypoumnas Antilope Cram. (Anomala Wall.) in this respect has come to
resemble most a Euploea. The colour is completely that of a Euploea ;
in the (3 even a strong interference blue has been developed on the upper side
of the fore-wings, as in Euploea Midamus L., but of a lighter shade, sothat
the expert who, as happened to me, sees it on the wing in broad daylight,
although not in the glare of the sun, immediately distinguishes' both species in
spite of all ideas of mimicry. The interference colours are governed by the
form of the scales and the difference in this respect in two not allied
butterflies will presumably remain constant, however much the pigmental colours
may change. In addition white begins to make its appearance on the universal
brown in these Hypolimnas, as in Euploeas; at first in the form of white
dots and, in one island more and in another less, in white streaks along the
inner margin of the hind-wings, just, therefore, where in Euploea Climena
Cram. Sepulchralis Butl. the white is most developed, becoming diffused into a
broad border surrounding all the wings, just as in this and even other species
of Euploeas, in the K6 islands. In various species of Papilio, such as
P. Paradoxa Zinck, P. Canopus Westw., and others the colour evolution
has in general followed the same course as in Euploeas ; in some species,
such as P. Caunus Westw. and P. Agestor Gray, this has been carried so
far, as regards the white, that they acquire a strong so-called mimetic resem-
blance in colour to certain species of Euploea. In the Zygaenidae, a family
of Heterocera which appears to be specially susceptible to such influences, the
same Euploea colour occurs in a few species, most so in Isbaria Midamia
H-Sch. and Isbarta Militans Butl. Another species of Hypolimnas, H. Bolina
has, indeed, not shown itself as susceptible to this influence as H. Antiopa
Cram, but has not been able to escape it entirel)^ ; amongst the different forms
of the 9 of that species one occurs, Perimele Cram, in which the Euploea
colour appears likewise. In the c? of a species of Elymnias, E. Undularis
Drury the colour development of Euploea is also present, but in the 9 that

XVIf

followed by the genus Danais ; this therefore, points to a transition phenomenon
as regards colour evolution the same as occurs with respect to wing form in
the fore-mentioned Dismorphias in South America, in which the one sex is
already under the influence, directed in a definite course, but the susceptibility
to it has not yet developed in the other, which, consequently, retains the ancient
form of wing unchanged. This case is analogous to that of Euploea Midamus L.
whose d" is a typical Euploea whereas the 9 principally exhibits the colour
adjustment obtaining in the genus Danais and, therefore, resembles D. Agleoides
Felder more than a Ei-ploea, pointing also to a state of transition on which
hypothesis the Euploea course must have been developed from the one peculiar
to the genus Danais sothat the 9 of E. Midamus L. and Elymnias Undularis
Drury have retained the old form which, according to the ontogenesis of the
Euploea larvae, appears, in fact, to have been the case.

The same occurs with the other course of colour evolution which, besides
the one alluded to, has been followed by the Malayo-Australian Danaids and
is most observable in the genus Danais. In addition to the genera Hestia
and Ideopsis it occurs also in the different species of Papilio living in the same
districts, such as P. Macareus Godt., P. Idaeoides Hew. and others; like-
wise in Nymphalidae in species of the genus Hestina Westw. and in Satyridae
in species of the genus Zethera Felder and even in the family Zygaenidae
already alluded to, inter alia in Isbarta Aspasia Sn. and Cyclosia Papilionaris
Drury. Such cases of uniformity in the process of colour evolution or so-called
homochromatism are, moreover, known in different genera and families in
South America. Bates has referred to it and Blanford discussed it in A'ature
of June 2o'ii 1895 ^n<^ indicated several very important instances of it at a
meeting of the Entomological Society of Londoti on May s'*" 1897. Its cause,
however, was unknown to them.

When we observe how the appearance of white, which in Euploeas we
know to be an expression of the course of the process of colour evolution,
manifests itself in the island of Celebes in two species of Euploea occurring
there, but likewise in other Malayan islands, E. Viola Bud. and E. Schlegelii
Voll. in a different manner from what obtains in those other districts, yet in
both deviating therefrom in the same way and in complete agreement with what
occurs in this respect in several other species of Euploea indigenous in that
island, while at the same time other species of that genus occurring in Celebes
as well as elsewhere do not exhibit this special modification, but resemble their
congeners in other islands, this can hardly be explained otherwise than by
attributing it to some influence operating locally in Celebes, an influence which
not only governs the course of colour evolution in the indigenous species but

3

xvni

has operated in like manner on the two species arrived thither from other
parts, without influencing others in which this susceptibiHty has not yet been
developed.

On these grounds we must, therefore, admit the existence of such local
influences operating on the evolutionary processes to which the Rhopalocera
are subject although we may be unable as yet to indicate their nature. When,
however, empty phrases or the forementioned erroneous conception fail to satisfy
us, this conviction, as clearly follows from what has here been formulated, can
only be based on the phenomena revealed by a ^tudy of the evolution both
of the forms and of the colours of butterflies, a study which must, therefore,
reveal the existence of the phenomenon of colour evolution also. It is, more-
over, not the only fact in this connection which can only thereby be explained.
In my monograph of the Pieridae of Java, on page 50, I explained in this
way the presence of the red dots on the wings of the European GoNOFrERVX
Rhamni L, which are doubtless well known but whose cause nobody has
hitherto been able to indicate, and on pages 41 and 50 1. c. the equally
known and equally mysterious 8 shaped metal- coloured spots on the hind-wings
ot some species of Colias. On page jj I pointed out how clearly this pheno-
menon manifests itself in the various colour forms and the remarkable colour
relics, connected with them, of the genus Hebomoia, such as the streak on the
upper side of the fore-wings retained in the specimens of a species from the
Philippines, Flores, Sumba, and in a lesser degree in those from Java and
how completely it explains this but only in accordance with my conception of
the problem. In many other places besides I have clearly indicated this process.
What is the reason that this theory of mine of colour evolution, for which I
have striven so many years, not with hollow phrases indeed, but with the
adduction of large quantities of material in support thereof, has received so
litde credence? Can it be because, unlike so many present day biological
theories which receive much support, it is not based on superficial reasoning
and hollow propositions supposed to be confirmed subsequently by isolated
observations insufficiently controlled but strongly biassed by preconceived
notions, but is the result of the unprejudiced observation of a number of
mutually corroborative facts? Because its examination requires a method in
which those are not versed who have been brought up in the present day
experimental methods — /. e. of investigating what has already a priori been
assumed — but are not accustomed to explain logically by means of extended
comparison facts obtained by unprejudiced observation? Perhaps so, for no

XIX

attempt at thorough investigation of its value or at serious refutation of its
propositions has ever been made. In his " Experhnentek entomologische Studien
vom physikalisch-chcmischen Standpunkf aus " Prof. Bachmeijew has indeed given
an account of it but this has not led to any serious discussion. The Professor
even apologises, as it were, for giving an account of it and when he calls my
views " very original " this may possiby be understood to be a polite term
for "absurd". For thus he regards them in his narrow-mindedness and thus,
he supposes, correctly perhaps, that they will mostly appear to other biologists
of supposed scientific standing. For their point of view, like his, is the physico-
chemical one, in which they have been educated ; outside this only soul-saving
scientific dogma they deny all science, and my views based, not upon such
pedantry, but upon independent observation are wholly outside that dogma and
even reject it as such. Such an attitude is, to them, absurd and, therefore,
unworthy of investigation. I will not deny that a certain amount of change
in this attitude appears to be imminent for quite recently the following sentence
in an English periodical drew my attention : " It is perhaps true that physio-
logists are on the whole less satisfied now than formerly with the adequacy
of the physico-chemical explanation of vital activities".

My opinions encounter, moreover, much opposition through the inability, so
general amongst biologist, of grasping the principle of evolutionary change, for
the phenomenon of colour evolution is simply one of its manifestations. I have
already stated previously that since Daravin's time the word " evolution, is met
everywhere but in spite of this the subject itself is as yet very little understood.
And no wonder. When new opinions, no matter on what subject, whether
religion, politics, science, or what not, make their appearance in the human
mind they are the result of a higher mental development, acquired in this
connection by certain persons and which, mostly favoured by special circum-
stances, have found recognition. But it does not follow from this that such a
higher conception penetrates the mind of every one. This can only be effected
after a shorter or longer interval when the general level of mental development
has been raised so far that this higher conception can be indentified with it.
Not until then can the mental processes move and work on these lines; and
until then this higher conception, while openly acknowledged and taught,
remains in reality, so to speak, unassimilated and incapable of being elaborated
by such persons who constitute the large majority in every region of mental
activity. Thus it will be seen, when Indian nations are subjected to an alien
higher civilization, what is called the polish of European culture is produced
in them — a superficial layer of imported forms, ideas, and conceptions —
without governing actual thought and consequent action. Thus the mental

XX

attitude of the parvenu remains generally on the inferior level peculiar to his
birth or former environment ; hence the French proverb : " Froitez le Russe, votis
trouvercz le Tar tare". No subsequent teachings of morality or religion alter the
nature of him whose inherited inferior moral development unfits him for
its reception ; his natural egoism makes him the born criminal in the well-
conditioned state ; it depends only on the conditions of life in what manner this
will find expression. In the domai^i of science the same may be observed ;
the disciples generally adopt the dicta of the master but they elaborate these
from their own cultural level. The theory of evolution, similarly, is at the
present day universally accepted scientifically which implies that every species
must be considered as being, as a rule, in a condition of continuous evolutional
change, but in spite of this the old view of immutability of species frequently
overrides that of biologists. Darwin as wel as Lamarck, indeed, conceived
living nature as being in a condition of constant evolutional change, interrupted
now and again temporarily by some cessation in development, but the theory
of mutation again assumes an unchanging condition as the rule, only now and
again interrupted by so-called mutations, approaching thus in this respect the
old view, just alluded to ; but since the cause of their appearance remains
completely hidden this theory strongly reminds us, in fact, of distinct creative
acts in which the Creator alone is either disregarded or indicated by a note of
interrogation. For this reason I can only consider this theory as a retrogression
in the domain of the doctrine of evolution.

Now when I consider the detailed observations made by lepidopterologists
on the frequently very slight modifications in colour and markings in many
species and note how, purely on such differences, all kinds of species and
subspecies have been based, I fancy I can see much of the old conception of
the immutability of species in this method. For if they had grasped the bio-
logical principle of the evolutional mutability they would have understood that
these numerous differences, each of little significance in itself and running into
each other, can be nothing but so many expressions by wich the same evo-
lutionary processes manifest themselves in proportion to the individuality of each,
processes amongst which, in Lepidoptera, that of colour evolution especially
comes to the front.

While the mutability of a species is, indeed, acknowledged it is yet considered
to be a fixed entity ; but that each individual, belonging to any species, inde-
pendently plays its part in the universal evolution process is still very little
understood.

The processes in question nevertheless operate in this manner. The power
governing the processes oi change, to which the species is subject, affects every

XXI

individual, but in proportion to its conditions of life and to its susceptibility to
this power which constantly varies in individuals. Hence each individual
assumes an independent station in the course of these evolutionary processes
and amongst the individuals, therefore, all stages may be encountered formino-
transitions to one another. Now in the same manner as the cinematograph,
by the combination of a number of impressions of successive stages, reproduces
a whole occurence in its natural sequence, the individual colour patterns indi-
cate to the person, possessing sufficient lepidopterological knowledge to combine
them, how this process of colour change, of which each pattern represents a
specific stage, operates.

Colour evolution teaches us how to explain the origin of existing colour
variety, somewhat in the same manner as embryology and comparative anatomy,
from forms in various stages of development existing side by side; it teaches
this, however, by observing the living forms and its study is thus identified with
that of the nature of evolution in general in which so much remains to be
elucidated, and is, therefore, of far-reaching importance.

Recently I read the following sentence, written by a naturalist : " The whole
colour question of the plumage is one of evolution " . With the colours of
Lepidoptera it is the same. Nothing is further from the truth than the super-
stition so general amongst zoologists, against which I have already protested on
page vi of the Introduction to my Monograph of the Java Pieridae, that the
colour of animals is accidental, devoid of biological value and, therefore, not
susceptible to evolutionary change. But all this and consequently the pheno-
menon of colour evolution is, of course, scarcely comprehensible to those to
whom evolution is, as I stated, not much more than a term. Recourse is then
had to unknown influences which, even if they be termed meteorological, differ
litde in reality from the supernatural with which the uncultured satisfies his
lack of knowledge.

A great part, in this connection, is probably played by a certain pedantic
shallowness, unfortunately, far from uncommon, which, however, assumes a
decided scientific guise and as such may exercise much influence on the semi-
learned. This manifests itself in the, to my mind unwarranted, weight attributed
to the only objection, of any importance, raised against my theory of colour
evolution. According to this theory, as I have summarily explained on page ix
of the Introduction of my Monograph of the Java Pieridae, the original pigment
was red, which gradually faded to white until the pigment disappeared in the
scales, and during this process a black pigment, moreover, increased at first
and then decreased. It is objected to this that the ontogenetic investigations

XXII

conducted concerning the origin of the pigmental colours in the pupal stage,
have established a sequence which does not accord with the one assumed by
me and that in these red certainly does not occur as the original pigment. How
does the matter really stand in this respect? When the haemolymph of the
pupa, penetrated into the scales, has been transformed into a granular pigment
it is of a uniform dull yellow or light drab colour. This can, in a certam
sense, certainly be termed the oldest pigmental colour but only when this
pigment, as Mayer ^) expresses it, appears as "ornamentation", does it acquire,
through chemical changes in the haemolymph itself, what he terms the mature
colours which, therefore, arise from the action of chemical reagents ; experiments
have shown that in this manner, according to the reagents employed, red-orange
or yellow can be produced. By the term "mature colours", therefore, is
meant the colours which are visible in the imago after it has left the pupal
envelop and it is only with these the theory of colour evolution is concerned ;
when, in discussing these I used the term original pigment, I referred to the
pigment which, in the original forms of Rhopalocera, besides some black pigment,
constituted the universal mature colour and which in the present forms has since
been more or less modified by the process of colour evolution, but not what
has been termed original pigment in the ontogenetic investigations which as
such never appears as a mature colour and can only be classed with it after
undergoing chemical change. In consequence of what I have just stated con-
cerning these mature colours it is, therefore, (juite possible that in the primary
butterflies this colour should have been red to become gradually changed in
later times into orange, yellow, and white, especially since the pigments of all
these colours, in so far at least as this has been investigated, as in the Pieridae,
do not differ chemically. These changes must, however, have proceeded very
unequally in the different groups of scales which alone can explain the fact that
several of these pigmental colours appear side by side on the wings of the same
animal. This inequality of its expression in time and space is, moreover, as
has been correctly observed years ago by Weissmann, a fixed character in all
evolutionary processes and does not prevent these colour changes in any way,
from being governed by one and the same principle or, as it is usually termed,
by a fixed law. I always hesitate, however, to employ this so usual expression
" law " in this connection, since I am only too well aware that non-lawyers

') I follow here principally the two treatises by Alfred Goi.DnoROUGH Mayer, " The devel-
opment of the iving scales and their pigments in butterflies and moths" and" On the color and color-
patterns of Moths and Butterflies" {Bulletin of the Museum of Comparative Zoology at Harvard
College, xxix, n". 5, xxx, n". 4). Mayer's term " ornamentation " is good in itself, providing it
is not understood to imply that the process in question occurs for the sake of ornamentation.

xxnt

and consequently as a rule also biologists are inclined to understand by this
term something immutable, something which cannot be modified in its mani-
festations according to circumstances which is certainly not the case with these
so-called laws any more than with human laws. Only the leading principle
remains unchanged ; the method of its execution, however, is regulated according
to circumstances. That this process is not only possible but actually occurs in
this manner is admitted by the theory of colour evolution on the grounds of
the observations on which it is based. I will return to the subject presently.

The ontogenetic investigations concerning the origin of the pigmental colours
in the wings during the pupal stage do not show this sequence. It is true
that one colour appears earlier than another in that stage but that the evolutional
sequence of the colours, as this occurs phylogenetically, should be reflected in
it I do not consider at alle essential.

The method of tracing phylogenetic changes by means of ontogenetic investig-
ations rests entirely on the so-called fundamental law of Haeckel. The cor-
rectness of this law I certainly admit and it receives complete confirmation
from my study on the horn of the larvae of Sphingidae. But this does not
alter the fact that inemploying it in a scientific sense judgment should be
exercised since a slavish pedantic conception of it can only lead to erroneous
results which are of no value as scientific arguments. Vestiges or relics of all
the processes occuring during the phylogenetic development of an animal species
are not by any means traceable in the ontogenetic investigation of an existing
form. That Lepidoptera like all other animals have developed from older forms
cannot be doubted ; this is, indeed, demonstrated by a study of the wing-
venation and of the other evolutionary changes mentioned in the introduction
to my monograph of the Java Pieridae. That the same is true also as regards their
colours is proved irrefutably by the study of colour evolution and is evidenced,
moreover, by the considerable difference in colour frequently exhibited by indi-
viduals of a species occurring in different, and often in the same, districts at
different times or even contemporaneously, although they all have doubtless
originated from the same primordial form. Prof. Weissmann admitted this
years ago. In caterpillars, whose various stages of development marked off by
their moults greatly facilitate the ontological investigation of the later periods
of their development, it may actually be observed, as for instance in the larvae
of the Sphingidae examined by me in his connection, that the colour gradually
changes. The assertion, however, that in observing the first appearance of the
colours in the wings during the pupal stage, in which they are developed, the
phylogenetic sequence of the origin and changes of these colours may be traced
does not by any means appear to me to rest on such a firm basis. As we

XXIV

have seen this sequence must have depended on the presence of chemical
reagents varying in the course of ages through vital requirements unknown to
us, whose action, moreover, each time affected, in an extremely unequal manner,
only certain groups of scales.

Is there sufficient ground, therefore, for assuming that when an animal, in
the course of attaining maturity, reaches the actual state of development acquired
by the species all these chemical reagents and the vital influences which excited
them will be repeated in the same order in its ontogenesis? It is true in the
case of the colour of the Sphingidae larvae, just alluded to, the old colour
occurs ontogenetically but apart from the fact that there is only question of a single
colour change here, there is no comparison with the great difference in colour
occuring in the pigments of the Rhopalocera ; in the development of the im-
agines, moreover, no well-marked stages, as are caused in the larvae by the
moults, are to be observed but the whole formation of the colour pattern is
completed in a very short time, according Dr. van Bemmelen within 36 hours
before the emerging of the imago. In the course of the various observations
made in this manner with reference to the origin of the colours in the wings
I could find, indeed, nothing which points to a phylogenetic continuation of
colour development.

As a matter of fact I can discover in the result of the investigations onl}-
indications concerning the manner in which the colours of the wings in certain
butterflies in their present stage of development have arisen and nothing which
points ontogenetically to an orderly change from the original to the existing
colour, in the manner this must have occured phylogenetically. To the alleged
negative result of these investigations in respect of the theory in question I can,
therefore, attach little value. The physiological fact that the pigmental colour
in the larvae of Lepidoptera can change gradually as an unmistakable mani-
festation of an evolutionary process has been definitely established as the result
of my studies in connection with colour changes in the Sphingidae larvae. Of
an adaptation, pure and simple, of the colour to the environment, as occurs
sometimes in certain larvae and pupae and which, in the usual superficial
manner, has in this case also been given as the explanation, there is certainly
no question. Now when an extensive series of interdependent and therefore
mutually corroborating observations, such as I have published, points to the same
conclusion in the imagines of Lepidoptera in general, the admission of this fact
is not to be made dependent on the (}uestion whether this can be established
ontogenetically by experiment. Insistance on this condition simply reveals a
pedantic narrowness of conception.

In reality these investigations have in this respect little value in themselves.

\xv

Not being acquainted, or only slightly so, with my observations concerning the
colours in Rhopalocera and the manner in which they change in individuals,
investigators have worked in this respect not only without reference to these
observations but even with a total disregard of them ; a different line of research,
as well as a careful selection of material specially adapted for investigation
would have been a great desideratum in this connection, but haphazard
work has mostly been undertaken in this respect which, moreover, is strongly
biassed for the greater part by false theories, such as those of mimicry and
natural selection, or those of the German Professor Dr. Eimer, and endeavours
above all to seek confirmation of the said theories. Where, for instance, the most
accurate of these investigators, Alfred Goldsborough Mayer, actually pays
much attention to the colour pattern of the mature imagines, he simply follows
Baieson and his wholly unexplained fantastic principle of variability, but is
unable to conceive these colour changes as being the expressions of evolutionary
processes in a specific direction. I am unable, therefore, to attach much value
m this respect to his investigations but I consider it of interest to draw attention
to one or two points in them which prove clearly that his very accurate obser-
vations, although not, in my opinion, correctly interpreted, agree with my
own as regards the process of colour evolution. In his chapter " General
summary of results believed to be new to science " he says under (8) {a) '• when
in process of disappearance, bands of colour usually shrink away at one end".
This agrees entirely with what I have noted when discussing the first commence-
ment of the formation of the spurious eye spot in Cyllo Leda L. And again
under (12) "A record of the characteristic markings upon the wings of the
Danaid and Acraeoid Heliconidae shows that, physiologically speaking, the
colours red, rufous, yellow, and white are closely related and that black is
quite distinct from these, being the least variable colour of all". This, surely,
is just what, according to my observation, constitutes such a prominent fact in
the theory of colour evolution.

I believe that with a view to the experimental ontogenetic investigations
referred to, attention may be drawn to a phenomenon which in my opinion is
too much neglected. It is true I do not attribute very great value to it but it
will not be amiss to consider it. I refer to what occurs in the appearance of
the mature colours. It is known especially in Coleoptera that after reaching
the mature stage their colour still requires some time before being completely
developed. Something similar I observed, as stated on page 62 of my Mono-
graph of the Java Pieridae, in the larva of Terias Hecabe L. Until the last
moult the larvae of this butterfly and those of T. Sari Horsf. have the heads
of the same colour as the body generally, /. e. green. But whereas in T. Sari

4

XXVI

Horsf. the heads remain the same after this moult in T. Hecabe L. they
become black. Not suddenly, however, for I observed that after this moult
had taken place the head of this larva was still green ; at first it became
gradually darker and not until after about an hour did it become black which
thenceforth remained constant; probably, however, it is such a dark green that
it appears black to the eye. In the larva of Papilio Agamemnon L. also I
remember making a similar observation. The pupa of the Java Acraea
Terpsichore Cram, has the ground colour of a beautiful milky white much
marbled with black and ochre-yellow. A couple of days before the imago
emerges this milky white changes into an intense rose and after the butterfly
makes its appearance no trace of either milky white or rose is visible on its
wings or on the pupa case. How is this to be explained ? I have been unable
to investigate it but it appears to me that changes occur here which have no
connection with the phylogenetic development of colour. It leads incontestably
to the assumption that a certain amount of influence of air or light is required
for the maturing of the colour as it occurs in the perfect insect; it may,
therefore, be doubted if such reaction really occurs in wings exposed artificially
during the experiment and whether the mature colours may be traced back
to the earlier stages rather than that they should originate in the present stage
of development of the chrysalis. For this reason I consider this phenomenon
deserving attention.

On the other hand there is a great deal which, in my opinion, confirms
the propositions of the theory in question. The most important is undoubtedly
the fact that it is becoming more and more manifest that the numerous colour
phenomena observed in Lepidoptera can only be explained by means of it.
This is especially striking with respect to the remarkable cases of colour dimorph-
ism or polymorphism which cannot be reasonably explained otherwise although
there is no lack of phraseology in this connection. Where such cases occur
regularly side by side as in Pap. Memnon L. they follow the rules of colour
evolution ; likewise where they make their appearance contemporaneously but
in different districts, as in Pap. Dardanus Brown ; and again where they are
restricted to specific periods by climatic influences, showing clearly which is
the more and which the less developed form, the same occurs. Finally where
by the action of retarding influences, of whatever nature, only a lower, and
therefore older, stage of development than the present normal one is reached,
this older stage agrees also with the doctrine of this theory. This alone enables
us correctly to understand the different results obtained in experiments with
heat and cold conducted in connection with the development of butterflies,
which are for the most part wrongly interpreted. A comparative study as to

XXVII

what must be considered relics amongst the colour spots in Rhopalocera proves
the existence of the evolutionary change of these in the clearest manner. Such
a study, however, only becomes possible of course when one has learned te
interpret these changes, not as the result of accidental climatic conditions or
similar airy assumptions — the ignorant would call it the action of spirits —
but as that of regular manifestations of natural evolution. In which connection
one may with advantage ponder over the words of Prof. Dr. Klaaisch when
proceeding to the ontoological comparison of the Neanderthal Gorilla type
with the Aurignac Orang type, {Die Umsc/iau, Sep/, j, igio), he states that
any one who wishes to form an independent judgment on this subject cannot
dispense with a thorough study of the skeleton of man and of anthropoid apes
and that sceptical phraseology will not do service instead. This, in judging
the colours in butterflies as in other matters, is in the majority of cases simply
overlooked for the sake of convenience.

The study of colour evolution invariably indicates that red must have been
the original pigmental colour, in the sense of mature colour as stated. But
this is not all. Mayer in treating of the " Quantitative Determination of Pig-
mental colors " states " to have analysed the colors of many butterflies by means
of the spectroscope and also by Maxwell's discs ", and that in this manner
he has " determined that the vast majority of the colors found in Lepidoptera
are impure; that is to say, they contain a large percentage of black and that
the so-called " blacks " found in butterflies, are rarely jet-black but almost
always only deep shades of brown " . Thus, for instance, he found in the
glaucous green of Colaenis Dido 29% black and in addition 24 "/o vermilion,
in the sepia-brown ground colour of Cercyonis Alope 71 "/o black and 21.5 "/o
vermilion ; in the tawny rufous colour of wings of Mechanitis Polymnia
4670 black, 40 "/o vermilion, 1470 lemon-yellow; and in the rufous red patch
on the upper surface of the fore- wings of Heliconius Melpomene 2 7 ^o black,
66.5 "/o vermilion, 6.5^0 lemon-yellow. In all these cases, therefore, the colours
are mixtures of red and black of which red constitutes a very considerable
portion and sometimes also yellow, all of which, therefore, completely accords
with the theory of colour evolution of an original red colour with the gradual
increase of black, and sometimes fading te yellow. Whence otherwise comes
all this red and why is it thus mixed with black? This is supplemented by
what Mayer further states about his Spectrum ana/ysis of colors oj Lepidoptera,
which leads him to the conclusion that " in general the colors of the wing are
not simple but compound ; that is to say, they are made up of a mixture of
several different colors" and in these red and yellow frequently occurred; it is
worthy of note in this connection that he found that " the spectrum of the rufous

XXVIII

ground color of the upper surface of the wings of Danais Plexippus consists
of all the red and yellow of the spectrum and about 75^/0 of the green".
Here we have to do, therefore, with a butterfly of the same genus Danais,
with which we were concerned just now and which is also closely allied to the
genus EuPLOEA.

On the one side we have, therefore, a number of very careful observations
which, on being correlated, completely elucidate and confirm each other, by
means of which a number of biological phenomena are explained, hitherto not
to be accounted for in any other way, and for whose interpretation all sorts
of unproved and even incomprehensible influences have been invoked. On the
other side we find some incomplete anatomical investigations, some technical
dexterity in which certainly cannot be denied, but whose relevance in this
connection is very doubtful and which were, moreover, accompanied by so litde
knowledge of what had already been proved by investigation and comparison in
this field, that this must of necessity have exercised great influence on the
interpretation of the results of these investigations. I believe the choice between
these two cannot admit of much doubt.

I think the foregoing observations are sufficient to convince the unprejudiced
that the theory of colour evolution rests on a firm basis and can in no way
be considered as refuted. I will not further dilate on this; those desiring further
information on this subject and as to what is to be observed in this connection
in other animals may find additional particulars in the chapter " Farbeuevohttioti "
in the second section of my work " AMi einmal Mi7nicyy, Sclcktion, Danviiiistnus "
published in 1907. I would only here quote the following from the treatise
published in 1 904 by Dr. Arnold Jacobi " Dt'e Bedeutung der Farben im Thier-
reich ". " The first pigments " this savant states "to arise, in the gradual develop-
ment of animals, appear to be those corresponding to the left side of the
rainbow and admitting, therefore, only rays of very great wave-length and very
little refrangibility and as a consequence red is the most common ground colour
to which yellow is gradually and green sometimes also added. That red is
the starting point of all colour development may be inferred from the fact that
the actual perception of light in our eye is connected with a finely divided
red substance, Rhodopsin, in the retina. Thus the primitive eye-spots in various
Infusoria are red and in Turbelaria and Chaetopoda, living in great depths
of water where light scarcely penetrates, as in the subterranean Aphides, the
eyes are also red, although the pigment in the eyes of their allies exposed to
light is dark.

Moreover, many animals whose stage of development has retained an ancient
simple character are red, such as, for instance, some lobsters; even our common

XXIX

marine and fluviatile lobsters and crabs have a red body colour although this
is hidden by a brown colour until the latter has been driven off chemically by
means of boiling. In relation with the change in the pigment of the eye many
aquatic animals which, when they occur at slight depth are colourless and
translucent, become red when they reach greater depths and darkness. In a
like manner many animals living in holes or inside plants exhibit this other-
wise so striking colour, such as the orange-red larvae of Cecidomya, those of
Trypanus Cossus L. and of many microptera as well as certain Indian snakes
(Uropeltidae).

Concerning the caterpillars of Danaidae little is to be noted. Those of the
genus Eui'LOEA appear to indicate that this genus has become specialised from
the genus Danais. It has, indeed, been already noted that the colour develop-
ment in EuPLOEA MiDAMUs L. 9 greatly resembles the one which obtains in
the genus Danais and that this makes it probable that this species remains
in the condition — as may be met with in others — in which one sex has
retained the older form, while the other has assumed the later one. The
Danais form in this instance, therefore, must be the older one from which
the Euploea form has been developed. Amongst the little known Euploea
larvae, which, like those of the genus Hestia differ more especially from Danais
larvae in the long fleshy dorsal processes, occurs one, /. e. Euploea Mazares
Moore, in which these processes are short and straight, corresponding, there-
fore, with those in the Danais larvae and having presumably remained in an
older stage which is that of the genus Danais. In very young specimens
of Euploea larvae, furnished with the long processes, they are equally
short and straight as in the Danais larvae. But considering that, in view
of the small size of these larvae, in any case the processes are of necessity
also very short it would appear somewhat risky to draw any conclusions from
this fact.

As regards the pupae of Danaidae much remains which requires elucidation.
Those of EuPLOEAs are conspicuous by their strong metallic lustre. Some,
such as those of E. Midamus L , completely resemble little clumps of polished
gold or silver, in others this lustre is interrupted by patches or streaks of a
dark or light brown or it is even more of a copper colour. At the fourth
International Congress of Zoologists, held at Cambridge, Mr. Edmond Bordage
stated that in the Mauritius he had reared in the dark larvae of E. Goudoti Bsd.,

XXX

whose pupae normally resemble polished gold or silver, and obtained pupae
having dark brown streaks and spots on this metallic lustre. These, therefore,
agreed with the normal pupae of E. LEUCOsricxos Gm. from Java. How is
this to be accounted for? It can hardly be attributed to colour. evolution since
this metallic lustre is undoubtedly of a structural nature whereas colour evolution
in only known in pigmental colours.

The pupae of some species of Danais also exhibit special phenomena. I
have frequently bred those of Danais Chrysippus L. and of D. Genutia Cram.
Of the former I obtained many pupae the majority of which were of milky
white, some retaining this colour while others subsequently became rose or
green. On the other hand I observed that of these pupae, reared in a clear
glass cylinder, which were of a beautiful green a portion subsequently turned
to a milky white while others under the same conditions remained green. The
bred pupae of D. Genutia Cram, were all green at first; some remained of that
colour but others subsequently became pale rose. None of these pupae contained
parasites. Whether they were reared in dark boxes or in clear glass cylinders
made no difference ; one beautiful green chrysalis of Danais Juventa Cram.,
formed in a very dark box, agreed completely in colour with one found by me
in a natural state. In all these pupae golden spots occur which appear as white
immediately after pupation and in which the metallic lustre only becomes
apparent later. How can these colour differences be explained? I believe in
order to arrive at this attention must be paid in the first place to what is
known concerning the colour phenomena occurring in the pupae of other
butterflies and that, therefore, I must take into account all investigations on the
subject, although these mostly have reference not to Danaidae, but to other Lepi-
doptera — Papilionidae, Pieridae, and even Heterocera. When some time back
I discussed the rearing by Mr. Edw. Jacobson of imagines of Papilio Memnon L.
I had also to concern myself with this question. My views in this connection have
since been published in the Tijdschrift voor Entomologie liii, igio. I will
briefly repeat them here. According to Mr. Jacobson the pupae — about 400 in
number reared by him at Batavia on its food plant had, almost without exception,
assumed the colour of their immediate environment, those attached to the
variegated greyish-brown branches of the citrus tree, on which the larva had
been reared, were of the colour of bark with green spots while those fastened
to leaves or green twigs were green. A few only had pupated against the
white gauze surrounding the larvae on the citrus branches and these were of
a dirty white without green spots. Mr. Jacobson expresses the opinion that
these results in general agree wMth those obtained by Prof. Dr. J. Vosseler
in breeding Papilio Demoleus L. in East Africa but do not correspond with

xxxt

my earlier observations in Java. During my stay in Java I did not indeed,
conduct actual investigations in this field, the importance whereof had not
attracted my attention at that time, but nevertheless I bred many larvae of
Papilio Memnon L. and P. Polixes L. for other purposes and my notes made
at the time are of value also in this connection. From these it appears that
in the pupae reared under varying conditions of light and darkness no influence
of this could be perceived, some being green others brown with green spots.
Of four pupae of P. Polites L. two, which had developed between dark green
citrus leaves in cylinders of clear glass, were of a striking dark green but two,
which had pupated in similar cylinders without leaves and whose environment
was greyish, were bark-coloured. Both Jacobson and Vosseler obtained similar
results with pupae reared completely in the dark and the latter expresses
astonishment that in that case such pupae assume the colour of their immediate
surrounding. Jacobson suspects that my negative results are due to the fact
that my experiments, unlike his, had not been conducted in full daylight. My
experiments, however, were certainly not conducted, like his, in the natural
state even if under white gauze, although by no means in the dark but in
cylinders of clear glass or in boxes covered with white gauze and in rooms
of houses where, indeed, subdued daylight only is admitted in order to avert
the great heat, but which are, nevertheless, as well lighted as an ordinary
mid-european dwelling. Moreover, in complete darkness similar results are
obtained as I have just mentioned. There is a great deal, therefore, still requiring
elucidation. Before proceeding I would, however, draw attention to one or two
points. So far as I can call to mind, and the illustrations in my possession
of Memnon pupae confirm this, the green chrysalides of this species are very
light, mostly yellowish green and by no means of the dark green of citrus
leaves; I also have a recollection that the pupae designated with the rather
indefinite term bark-coloured do not exactly conform to the colour of the
ligneous parts of the citrus tree. We must not lose sight ot this ; in such
matters we must beware of unconscious auto-suggestion.

A considerable number of observations and discussions on this subject have
seen the light. Many of the more important of these were available to me;
of a great many others, where this was not the case, I have been able to
consult the ample and apposite quotations published by Prof. P. Bachmetjew
in his " Expcrimentclle entotnologischc Studien von physikalisch-chemischen Stand-
punkt aus". In making use of these great circumspection must be exercised.
Many of the experiments have been conducted in so superficial a manner or
have reference only to certain animals or conditions that in this respect, where
the difference in susceptibility — not only between species but even between

XXXII

Individuals — leads to widely divergent, or even directly opposite results, litde
value can be attached to them. Not a few, even of the more important ones,
are governed by the suggestion of mimicry which, like every suggestion, not
unfrequently induces one to observe what is looked for sothat one cannot be
too careful in this respect. Others again, some of them of great importance,
are completely dominated by the preconceived notion that the colour mani-
festations must be explained by chromo-photographic action. The correctness
of this idea, however, is also ver)' doubtful and this should constantly be borne in
mind. Without losing sight of this we will proceed to examine these observa-
tions.

The fact that where a caterpillar at a specific period of its existence, when
it ceases feeding and prepares to pupate, is exposed to radiation from any
definitely coloured environment — especially during the twenty hours preceding
the last twelve hours before pupating — the colour of the resulting pupa
frequently adapts itself to its surroundings, may at the outset be accepted as
having been proved beyond dispute, especially by Pox:lton's investigations. It
further appears, however, that much difference in this respects exists amongst
the various kinds of caterpillars.

Of the older observations on this point, those made in 1876 by Mrs. Barber
are well known especially through Darwin having drawn attention to them.
According to this observer pupae of Papii.to Nireus L. reared against a twig
of the citrus tree, on which the larva lives, were light green ; between
dark green leaves dark green ; against dead twigs covered with pale yellowish-
green leaves they resembled the latter; while they were yellowisch against the
wooden frame of the case of the same colour. Red surroundings appeared to
exercise no influence but on the other hand a pupa attached to the junction of
purplish brown brick and wood composing the case was stated to have its
back the colour ot the bricks and the under surface that of the wood. The
latter observation, however, was doubted by Poulton since it did not agree
with his own experience; he justly refers to the fact that it is a common
occurrence that the colours of pupae differ greatly in the dorsal and ventral
regions and when it is borne in mind — what has, moreover, been proved
in other instances — that Mrs. Barber was a fanatical supporter of the
then novel theor}' of mimicry, I consider it advisable not to attach too
much weight to her observations. Mr. Roland Trimen subsequently con-
ducted experiments, on the pupa of Papilio Demoleus L. He found
these pupae yellow, green, or reddish-brown, according to whether they
wefe attached to bands of these colours fixed by him to the sides of the cage.
A black surrounding caused them to become dark but light red and blue

XXXIII

had no effect. Of more significance are the observations by Pgui.ton who
certainly is a strong and by no means unbiassed supporter of the said theory
but whose experiments have been conducted on so large a scale and with so
much accuracy that, even when we are unable otherwise to accept his con-
clusions, the facts observed by him merit every confidence. Six pupae of
Vanessa Io L., which occur in two colours in nature — dark green and more
rarely yellowisch-green — reared by him in a surrounding of yellowisch-green
tissue paper were all of that colour. In the pupae of Argynnis Paphia L.
and Vanessa Polychloros L., according to Wood and others the environment
caused them to become darker or lighter but otherwise the colour was not
affected. In Pieris Brassicae L. and P. Rapae L. the pupae become a
beautiful smaragd-green from a yellow or orange surrounding but a black or
white environment induced only darker or lighter shades ; the former also resulted
from every other surrounding except yellow or orange; blue has no influence
on Pieris pupae nor has orange on those of Vanessa. The most important
results were obtained by Poulton in experiments on the pupae of Vanessa
Urticae L., which he was able to carry out on a large scale. The larvae
which were the subject of these experiments were placed in green surroundings
in the hope of producing green pupae which in this species do not occur in
nature; but in vain, the pupae were only "somewhat darker than usual and this
suggested a trial of black surroundings, from which the strongest effects were at
once witnessed ; the pupae were as a rule extremely dark with only the smallest
trace, and often no trace at all, of the golden spots which are so conspicuous
in the lighter forms. A not inconsiderable number, however, retained the
normal light colour with golden dots. These results suggested the use of
white surroundings. The colours of nearly 1 50 chrysalides obtained under such
conditions, were very surprising. Not only was the black colouring matter as
a rule absent, so that the pupae were light-coloured, but there was often an
immense development of the golden spots, so that in many cases the whole
surface of the pupae glittered with an apparent metallic lustre as if they were
covered with gold-leaf". Sometimes the pupae were light pink in colour.
Finally a golden surrounding was tried and the same effect secured as with
white but with a great increase in the number of golden pupae. Poulton,
moreover, observed that whereas an orange surrounding induces a dark smaragd-
green in pupae of Pieris, that colour does not influence Vanessa pupae and
also that blue has no effect on Pieris pupae.

Concerning the pupae of Vanessa Io L., Dr. Kathariner {Bio/ogisckes
Centralblatt i8gg) has also published some observations in which, however, he
considers the rare green form a variety and the lighter or darker brownish-grey

5

XXXIV

form with a few metallic dots the normal one, while in my opinion the green
is the original form, now rarely met with, and the other the one which has
advanced further in the normal darkening process. According to his obser-
vations on a great number of pupae, those reared in broad daylight with light-
coloured surroundings were of the green form and those in darker surroundings
brownish-grey. In the green ones the usual metallic dots were present ; he
once found such a chrysalis in the natural state wich was entirely of a golden
lustre and on this occasion quotes the opinion of C. Ed. Venus that this
should be the result of an intense solar irradiation. The results mentioned,
however, only concern the large majority of the pupae, on which he experi-
mented ; contrary results occurred also, although in a much smaller number
and even when reared in complete darkness the green form occurred occasionally.

From larvae of Vanessa Cardui L. and V. Urticae L., which pupated in
a temperature of 37^ C. against white linen in full daylight, Standfuss obtained
pupae of a more or less white general shade but others reared similarly in a
temperature of 8^ — 23'' C. assumed the usual brownish- grey hue.

Prof. Dr. J. VossELER discusses in Zeitschrift fiir Wissenschafilkhe Insekten-
biologie. Band Hi, {igo/), p. 204, the pupae of Papilio Demoleus L., reared
by him in quantity in East Africa. Although a whole-hearted supporter of the
theory of mimicry and speculating on the subject to his heart's content, and
like Kathariner and others retaining the likewise antiquated tenet — attributing
the origin of the colours in pupae in relation to their environment to chromo-
photographic action — his observations on the matter are not devoid of im-
portance. He states that the pupae of this species vary from very bright yellow,
yellow, green, and bluish-green to bright and dark brown with green dots and
invariably marked with the same dark band ; the ventral side is brightest in
colour and thus most strongly imitates the colour of the environment; the
colours of bark and wood especially are imitated best, yet this imitation is
never absolute but is invariably confined within certain limits.

BoRDAGE also reared some pupae of Papilio Demoleus L. in the Mauritius
which, according to him, are grey or yellowish-green, the dorsal side some-
times more reddish-brown, while the ventral side is yellowish-green. The colour
of the environment where these pupae are formed, such as bark or leaves,
appeared to have no influence on the colour of the pupae, neither did light
or darkness, sothat under these conditions pupae of both colours occurred.

With reference to Acraea pupae from South Africa, Mr. Mansel Weale
informs us that in his experience a light or dark environment would cause these
to be lighter or darker in colour but that otherwise their colour did not change.

Finally some investigations have been instituted concerning the colour of

XXXV

the pupae of the South American Papilio Polydamas L. of which species it
has been stated by Fritz Muller that they are dimorphic as regards colour
and that both green and brown pupae occur. Also concerning the pupa of
the European Papilio Machaon L., which is likewise dimorphic, produces
green and brown pupae and similarly has not shown any symptoms of this
susceptibility. Various investigators have experimented with it in this connection.
I find it noted by a certain German observer that green as well as brownish-
grey or plain grey pupae occur in this species, some of the first named having
dark markings — all irrespective of the surrounding amid which pupation took
place, with this qualification, that those reared in a dark case, although showing
both the colours named, were in general somewhat darker. Cecil Floersheim,
in an essay entitled " Some 7iotes on the earlier stages of Papilio Macliaon, parti-
cularly with regard to the colour-dimorphism oj its pupa " occurring in The Ento-
mologist's Record and Journal of Variation, vol. xvii, {igo^), came to a different
conclusion. According to him the green pupae were formed almost exclusively
between green leaves but a few (2 to 11 5) against dark-brown twigs near the
soil, while on the other hand the brown ones occured mostly against these
dark-brown twigs and a few against green leaves but not against the light green
ones where the majority of the green pupae were found. Some fifteen, having
pupated against wood painted white and being exposed to the full glare of the
sun, were all brown and he, therefore, while admitting the influence of the green
leaves, does not consider that bright light promotes this adaptation.

Many similar observations are mentioned in the work by Wilhelm Petersen
" Zur Frage der chromophotographie bei Schmetterlingspuppen {18 go)" which,
however, are on the same lines as the others already referred to.

On quietly passing all these observations in review, ignoring for prudence
sake the first mentioned in which a strong desire is manifested of seeing what
the theory of mimicry suggests finding and provisionally also passing by the
quasi-alchemic ones of Poulton — to which I shall refer later — it appears
to me that they may all be summed up as establishing some amount of darkening
as the result of a dark environment or at least an environment making that
impression on the larva. This result being invariably in relation to the colour
which obtains normally in the species, as well as to the susceptibility to the
accompanying stimulus which always differs in each individual and to which is
due the fact, so astonishing to most observers, that experiments conducted
exactly in the same manner on various individuals not only differ somewhat but
show directly opposite results. In this manner the considerable difference of
temperature in the experiments related by Standfuss may possibly have occa-
sioned difference in susceptibility.

XXXVI

In this connection there is, therefore, question only of a greater or lesser
increase of black and this in relation to the susceptibility thereto, but this is
simply identical with the chief element of colour change, which I term colour
evolution, and which plays so important a role in imagines, which I first
observed in the larvae of Sphingidae and subsequently also in other larvae, and
to which, on page XXI of my introduction to the treatise of the Java Pieridae,
I attributed the increase of black in Pieris pupae. When we reflect that the
pupae which have been dealt with actually occur in two colour forms: green,
from yellowish to various darker shades, and grey, through various shades to
almost black, and that these two forms occur side by side in different indi-
viduals of the same species, such as : Papilio Machaon L., P. Polydamas L.,
P. Demoleus L., and P. Memnon L., and thus produce a pupal dimorphism,
the matter will admit of no doubt to any one acquainted with the phenomenon
of colour evolution. Wherever such a dimorphism occurs it points to the
existence of an evolutionary process of change in which a certain number of the
members of such a species have advanced further than the others which possessed
little susceptibility in this respect sothat two distinct forms, each exhibiting a
different stage in this process, have been produced. Exactly the same process
obtains in the case of the green Sphingidae larvae turning brown, which is
dealt with in my article " Uebcr die Farbe iind den Polyinorpliismus der Sp/iiugiden
Raupen ", and in which the course of the process can be distinctly observed
owing to the separation of the various stages of development by the moults.

In this case there is not only a very great difference — doubtless representing
many centuries of development — between the various species as regards the
stage of this evolution attained, but even between the individuals of the same
species there is a considerable disparity in this respect. The different shades
in which colours occur in pupae of the same species also point in that direction,
namely to a general darkening tendency which in different individuals, however,
is more or less developed. But to those well acquainted with the phenomena
of colour evolution this is by no means all. I distinctly remember, and Jacobson
also observes, that the dark brown-greyish pupae of Pap. Memnon L. exhibit
green dots ; Vosseler notes the same concerning the dark pupae of Pap.
Demoleus L. Whence these dots? They are relics or vestiges of the ancient
green colour of the pupae which in certain spots have resisted the evolutionary
darkening process; these are the same persistent spots to which I have already
referred when discussing the Pieridae, as regards the process of colour evolution.

In so far as the colour of pupae is still subject to this process and suscepti-
bility is present, its course may be accelerated by the influence of favourable
conditions of life. Now it appears that at a certain period shortly before

XXXVII

pupating a definite amount of illumination, in this case a darkened environment,
may bring about this effect, but that, on the other hand, an unusually strong
illumination or one caused by specific light rays corresponding to certain colours
of the environment, may retard or interrupt this process.

How and why? To a large extent the answer is still "we know not".
PouLTON, in order to prove that the influence of colour, alluded to, on the
pupa could not be the result of some response through the sense of sight on
the part ot the larva, covered the eyes of the latter with a certain varnish
during the said period before pupation so as to blind the larva. It may be
presumed that this was, indeed, superfluous; at that period when the skin —
and with it that of the head — is nearly severed from the body and has
probably but little connection with it, as is evidenced by the change of colour,
it may be assumed that the optical function no longer operates by means of
the eyes and that the pupal stage in this respect has already been assumed.
Nevertheless, even if we do not know in what manner, it may be taken as
very probable that the animal remains susceptible to influences of light, for we
are here concerned with a class of animals endowed with extreme sensitiveness
in this respect. Most of the Heterocera are unable to endure daylight and
hide away from it, some even passing the day in an extremely somnolent
state; on the other hand many swiftly fly towards any light in the darkness
such as the strongly lighted white tropical dwellings. Some Rhopalocera also
seek semi-dark spots in the dense forest while others again continually prefer
the full glare of the sun, many species being evidently unable to dispense with
it, sothat they cease to fly directly the sun disappears, even if only for a few
moments, behind the clouds, although the temperature may remain unchanged.
It cannot, therefore, appear strange that light, or the absence of it, should
exercise a strong influence on such animals and especially at a particularly
critical period of their life which would bring a particularly great susceptibility
in its train. In many animals devoid of eyes this is, indeed, also the case.

Now if an environment conducing to a decrease of light promotes the
natural development of the colour process in pupae, it stands to reason that
the reverse condition should likewise exercise certain effects. Hereto, more-
over, certain phenomena sometimes seem to appear which accord to such an
extent with other facts, whose occurrence has been observed through the action
of other abnormal influences on the pupae in the resuhing imagines, that they
also are by no means surprising.

Blue surroundings had no effect whatever on pupae of Pieris Brassicae L.,
and P. Rapae L., so that no susceptibility for this colour was present, black
or white surroundings, or even those of any other colour except blue and the

XXX vin

two colours to be referred to hereafter, resulted in the darkening of the pupae, /. e.
an increase in black ; yellow or orange surroundings, therefore radiating a very
bright light, caused the pupa to become dark green however. In the latter
some darkening, therefore, occurred also, but whence this green colour?

According to W. Petersen the colour of the pupa is due to the pigment
in the cuticula and the hypodermis. Now the pigment in the latter is green
in the larva and sometimes remains so in the pupa. If, however, an accu-
mulation of pigment occurs in the cuticula this green colour becomes darker.
Yellow or orange light, however, is said to prevent the formation of dark
pigment in the cuticula, causing it to become translucent when the pupa would
appear green owing to the pigment of that colour in the hypodermis. This may
well be the case but those who do not judge from a chemico-physical point of view
will be satisfied neither with this nor, therefore, with the chromo-photographic
explanation. It will be asked what, then, is the cause of the appearance of
dark pigment in the cuticula, as it appears to occur normally and is prevented
abnormally by such a particular light? What natural process is disturbed in
its normal course by this illumination? Petersen does not explain this; let
us see, therefore, if we may not be able to trace it.

The pupa of P. Brassicae L. is described by the Dutch Entomologist
Snellen as yellowish-green with black dots, that of P. Rapae L., as yellowish-
grey or brown with three yellow stripes, and that of P. Napi L., closely allied
to the latter, as dull green, finally that of P. Daplidice L, as green. Green,
therefore, appears to be the original colour of the Pieris pupae, partly turning
to yellow and in which, moreover, black commences to appear. The occurrence
of green in the forementioned experiments, therefore, can be nothing but the
reappearance of the colour peculiar to the pupae of P. Brassicae L. and
P. Rapae L. at an earlier stage of colour development, and is doubtless the
return to an earlier colour form similar to that occasioned in the colour develop-
ment in the imagines by the experiments with heat and cold on the pupae,
referred to on page XII supra, the B. forms of Dr. E. Fischer ; the result of
disturbance by abnormal influences in the natural process of colour develop-
ment, in the one case caused by heat or cold, in the other by too bright a light.

We must here draw attention to a series of other experiments.

In order to find out whether the colour of the environment also influences
the colour of the cocoons of Heterocera, Poulton made several trials. Thus
he found that larvae of Saturnia Carpini H. (Pavonia L.) formed very dark
cocoons in dark surroudings but white ones in bright light; that Eriogaster
Lanestris L. produced creamy white cocoons on white paper but dark-brown
ones between leaves ; that the cocoon of Halias prasinana L. is likewise brown

XXXIX

normally but white in white surroundings, and that the same occurs in the
case of the cocoons of Liparis auriflua W. V. and Rumia Crataegata L.
Bateson, however, in 1892 published in the Transactions of the Entom. Society
of Lotidon the results of some experiments conducted by him in connection with
the cocoon of Eriogastkr lanestris L. He obtained, indeed, the same results
but he expressed the opinion that the cocoon was normally dark coloured and
that the dark pigment is destroyed if during the formation ot the cocoon the
larva is placed in an abnormal condition through any disturbing factor, such
as being removed from the food plant when nearly full-grown or by the presence
of parasites.

Dr. Harry Federley, in an article in the Mcddc/anden af Societas pro Fauna
el Flora Fennica, igog — igio, expressed yet another opinion; he also obtained
similar results in his experiments but he thinks these are to be attributed,
chiefly at least, not to the influence of the light but to that of the greater or
lesser humidity ; when the white cocoons, obtained in this manner, were sprinkled
with water they acquired the brown colour one or two days after. It is
difficult, however, in my opinion, to imagine that in the many experiments
which have produced the said results the lack of humidity could invariably
have played such a role without it being noted and I believe, therefore, that
this observation must be considered as an independent phenomenon. Bateson's
interpretation could not, of course be reconciled with the opinion of Poulton,
so strongly suggestive of the theory of mimicry, as regards the cause of this
phenomenon. The latter, therefore, at once proceeded to contest it and those
Transactions for the same year contain an account of new experiments on the
same larva carried out by him, in which it was subjected during the formation
of its cocoon to various disturbances without inducing it to become white. He
thus considers to have sufficiently refuted the interpretation by Bateson.

It would appear to me, however, that Poulton may have been mistaken
and that Bateson's interpretation is really correct intuitively, if I may so call
it, /. c. without quite understanding its real bearing, while Poulton has over-
looked the principal disturbing factor in the case. Would not the presence of
bright, or at least white, light occasion an important disturbance in such a
heterocerous larva which in that conditions has continuously lived in daylight
but whose organisation of vision now becomes so changed that in the imago
stage it is only adapted for darkness and which, consequently, for its pupation
intentionally seeks dark places or protects itself against daylight by means of a
dark cocoon? I think this may be considered far from improbable and that
in this manner all these cases of white cocoons obtained on white paper or

XL

in white boxes can easily be accounted for. That bodily functions — and this
applies especially to secretions — are not exercised, or only partly so, under
abnormal conditions is indeed no uncommon phenomenon ; thus the secretion
of dark pigments might well be impeded by the disturbance here referred to,
either absolutely or to such an extent that the cocoon threads, originally white,
might become coloured in which latter case insufficient moisture of this secretion
might be the determining factor and Federley's experiment would, therefore,
be to the point here. This would again be a case of disturbance in the
natural course of development caused by white light, which would very well
accord with the forementioned interpretation of the action of yellow and orange
light on PiERis pupae. This appears to me to receive equal confirmation from
Poulton's experiments with the pupa of Vanessa Urticae L. As we have seen
by exposing these pupae to bright white light he obtained a few pink ones but
by far the greater number were very light in colour with an extraordinary
development of the golden spots, even causing a complete gilt appearance of
some of the pupae ; illumination with gilt surroundings, thus irradiating a bright
yellow light, further increased the number of strongly gilt pupae. In this
butterfly a very intense white or yellow light, however, appears to be required
for this ; simple white surroundings, such as white paper which influences the
cocoons of Heterocera, appears to be insufficient ; pupae of Pieris species and
of Papilio Machaon L. placed against a white wall became darker, not light ;
pupae of Pap. Memnon L. attached to white gauze assumed a dirty white.
Similar surroundings, however, may, as we have seen, produce different effects
in different species.

PouLTON was particularly pleased with his quasi-alchemic discovery of the
almost completely gilded pupae and, in the same manner in which phenomena
of mimicry are so hotly chased, the appearance in nature of golden spots, such
as occur in the pupa of Vanessa Urticae L., for instance, is attributed with
great ingenuity to a similar illumination or even where it may be observed on
a much larger scale as in South America — or for instance in the Malayan
EuPLOEAS — it is interpreted as a vestige of earlier conditions of similar illu-
mination ! But upon serious reflection there is no ground whatever for attributing
these strong and wide-spread metallic lustres, such as occur for instance in the
pupae of EuPLOEA and Atella, to such influence ; moreover the same occurs
in other life-stages of Lepidoptera. In the case of larvae, indeed, few instances
are known, although I recollect referring on page 56 of the Monograph of the
Java Pieridae to the steel-blue lustre of some cateq)illars of Callidryas Pomona F. ;
in imagines, however, many examples are met with ; the metallic spots on the
lower surface in some species of Argvnnis and those on the upper surface of

XLI

Plusias being the best known ; reference may also be made to the strono-
metallic gloss in species of Morpho and in several Lycaenidae. In the Vanessa
pupae also it is doubtless simply a question of prismatic colour phenomena of
whose nature we are still ignorant and which have no relation to the direct
influence of light but which may be excited abnormally by abnormal disturbing
influence of light in so far as the susceptibility for it exists, as in the pupa of
Vanessa Urticae L. Disturbance of the natural course of development, there-
fore, is the cause in this case. This reacts on the colour of the pupae in
the presence of abnormal influence of light, in the same manner as may be
witnessed with respect to the colour and form in imagines resulting from
various influences, as indicated on page XIV stipra. The occasional appearance
of a pink colour under the same influences is doubtless to be interpreted in a
similar manner.

The experiments in connection with influences of light have, therefore,
produced much confusion in the science of lepidopterology ; I hope to be able to
shed some light on the subject. It should be observed that a knowledge of the
phenomenon of colour evolution is closely associated with it.

Fortwith to explain all the phenomena to be observed in this connection
would, indeed, be too great a task and more than could reasonably be expected.
The explanation of some, however, appears not impossible.

The observations by Bordage indicate that in the normal condition Euploea
pupae are of a generally dark brown colour on which a strong metallic lustre
has a tendency of spreading more and more. Of the different species, some
are in a more advanced stage than others as regards this process. Now in
the case of pupae normally produced in daylight, when this process takes place
in the dark a disturbance is caused resulting in a decreased distribution of the
metallic lustre.

The Memnon pupae, like those of P. Machaon L., P. Philodamas L.,
and P. Demoleus L., are dimorphic, i. c. light green and dark greyish or
brownish, which is termed bark-coloured. A dark environment may cause the
former to turn to dark green, as has been observed in the case of the two
pupae of P. Polites L. ; moreover, with ordinary treatment or even when reared
in dark boxes, green as well as greyish or brownish coloured pupae may be
produced; very susceptible individuals only in that case acquire a dark colour.
A particularly great susceptibility to this evolutionary change seems to occur
in P. Memnon L. the same as in P. Machaon L., hence the slightest darkening
of its environment, when pupation takes place against a branch or bark, is
sufficient to cause them to become brown -grey, except as to the persistent green

6

XI.II

spot, alluded to iis a vestige of the old colour. The others, not subjected to
such surroundings, remain light green. A pronounced white environment, too,
appears to have a disturbing effect ; hence a few pupae formed against the
white gauze turned a dirty white, /. e. somewhat darkened white without green
spots. The phenomena noted by Kathariner in the likewise dimorphic pupa
of Vanessa Io L. are thus also explained.

With regard to Danais pupae I am still groping in the dark. The fact
mentioned by Poulton that white light induced the production of a few pink
coloured pupae of Vanessa Ukticae L. is not without interest in this connection,
since there appears to be but little difference between this and the more rosy
colour of a portion of these dimorphic Danais pupae. Bordage obtained in
this species faint rose, light green, more rarely yellowish-white and sometimes
even red pupae; the latter were, probably, infested by parasites and invariably
produced imperfectly developed imagines. He thinks that larvae which have
pupated against brightly coloured or metallic-lustrous surfaces mostly produce
green pupae and against dark coloured surfaces mostly white or rose-coloured
ones. With weak illumination against black surfaces and still more in total
darkness the rose-coloured and white pupae greatly increased in number. Sub-
sequently he found that of a number of pupae placed in golden-lustrous sur-
rounding during a day and night ^js had become green and Ys rose and white.
This does not, however, shed much light on the subject and certainly does not
account for the changes observed by me. My observations in this respect,
however, were accidental and were not the result of experiments conducted
intentionally with that object in view. In order to be able to form a sound
judgment such experiments still require to be carried out. Many other investig-
ations also are necessary in this field. We are only at the commencement of
the enquiry. In spite of the many experiments conducted in this respect con-
cerning the facts, or of their extent and accuracy, incontestably deserving every
praise, such as those by Poulton for instance, I believe that, as regards the
interpretation of these facts, in general not much of consequence has been
contributed. Poulton and others, completel)^ obsessed by the theory of mimicry,
endeavour to explain the observed facts simply by mimicry fictions ; even the
chromo-photographic interpretation still counts too many adherents. For this
reason I can attach but little value to the important study in connection with
this subject contained in the work, already referred to, by Wilhelm Petersen.
This savant completely adopts this attitude, the phenomenon of colour evolution,
and with it every attempt to interpret the colour phenomena on evolutional
lines, being wholly foreign to him. The chromo-photographic theory proceeds
from the physico-chemical point of view, which is simply based on a vague

XLIII

hypothesis by which the bearing of the phenomena of life can only be partly
explained. While this theory may, in itself, not be so impossible, as Meldola
and PouLTON appear to think, it can only be applied in connection with the
evolutionary changes to which the colour as much as the whole form of each
animal is subject and which may differ considerably not only in the greater
divisions but even in individuals. Where it does not take this into account I
believe consequently that it cannot lead to the desired goal and that all investig-
ations and observations conducted from this point of view lead a priori on
the wrong track. Even resemblance of the colour of pupae to their environ-
ment, the conception of so-called bark-colour, must be studied with greater
attention, since a great deal of superficial observation has been brought to bear
on the subject; moreover, the same remarkable agreement observed in many
larvae has to be studied more accurately and may prove ot great service for
comparison in this connection. Nearly alle observations noted in this respect
have taken place far too much under the suggestion of mimicry to allow of
an unbiassed judgment being formed. It should, therefore, be borne in mind
that this resemblance to the environment occurs, not only as regards colour,
but equally as regards shape, as for instance in the case of the larvae of
geometrae resembling twigs, which can hardly be attributed to the influence of
light rays. Moreover, chemical and microscopic investigation is indispensable
here. We know that the colours of imagines, except such white as is produced
by the filling with air of the scales, are of a pigmental or structural nature.
We know something about these pigments but our knowledge is still very
incomplete, and as regards the occurrence of structural colours too we are as
yet very ignorant. Without a doubt pigmental colours occur in caterpillars
but others exist also even on a large scale, but what do we know of these?
And as regards the pupal colours, observation leads us to suspect, indeed,
something similar but concerning their structural colours too we know practically
nothing. Our knowledge generally with regard to these colours in still extremely
insignificant. I have already referred to this at the conclusion of the chapter on
the influence of light in my work '' Noch eiiima/ Mimicry, Selektion, Danvinis)iiits" ,
published in 1907. A considerable amount of specific and accurate investigation
is still needed in this respect. Only then shall we make further progress with
the interpretation of the colour phenomena in the pupae of lepidoptera.

Even the urgent recommendation of further study in this field on these lines
has its value; with this I propose, therefore, to terminate my reflections.

XLIV

Now as regards the Satyridae, on proceeding to a general consideration of
this family, two of the subjects, already referred to when treating of the Danaidae,
come to the front, in respect of which clearer ideas are required than usually
obtain in order to understand the phenomena which present themselves so
prominently in the butterflies of this family. First, that of colour evolution
which in this family plays so important a role and frequently produces colour
phenomena which, without a knowledge of this process, are inexplicable. Secondly,
the mistaken notion, already referred to, concerning the alleged seasonal varieties
which, in judging the differences existing between butterflies of the same species
in this family, has attained a truly fearful extent.

We will first consider colour evolution. There are surely few things which
to every lepidopterologist, who is concerned with the study of the Indo-
Australian fauna, appear equally striking as the great variation in colour and
marking of the wings of Cyllo Leda L., especially on their under side. This
is not, indeed, confined to this species, since in Kallima paralecta Horsf,
for instance, the colour of the underside of the wings also offers much variety,
but in the former it is much more pronounced and on account of the common
occurrence of this butterfly this rivets the attention more. No other phenom-
enon which presents itself in connection with the subject, I venture to say,
hcis been less capable of interpretation. Of course there has been no lack of
phrases concerning the influence of the environment etc. in this connection but,
although it is not impossible that this may exercise some influence, it is by
no means certain that this is so ; in any case this assumption does not really
help us to an understanding and the theory of mimicry enables us least of all
to explain the origin of so many differences. The fact that such resemblance
in colour with its environment may sometimes afford a butterfly some protection
against its enemies, in so far as these pursue them by sight only, may be
taken for granted but it does not explain in what manner this arises. With
the aid of the theory of colour evolution, however, this truly very intricate
phenomenon may be elucidated, and, at least at present, exclusively by means
of it, which consequently constitutes a very powerful argument for the correctness
of this theor}'. The value of a theory can, indeed, be gauged only by what
it enables us to elucidate.

When treating of this species in its proper place I will fully explain my
meaning in this connection, but since this has a general bearing on the value
of the theory in question I have thought it useful to refer to it here.

An explanation of many other phenomena, which are particularly conspic-
uous in the Satyridae, must also be looked for in the theory alluded to.
Midway between the dark-coloured species of a genus so typical of this family.

XLV

like Mycaiesis, occur some whose wings show a fair amount of yellow and
even orange; in Java, indeed, this is the case only in a couple of species but
in others, not occurring in that island, it is found in a much greater degree.
Whence this difference? Since we have already traced the course of the
process of colour evolution in many other butterflies it is easily assumed that
the original Satyridae were also red and that subsequently the red of these
butterflies turned to orange and yellow in the usual course; that, moreover, at the
same time a great increase of the black pigment occurred causing the colour
of the wings of many species to become darker in varying degrees of intensity
while in others, which in consequence of the inequality, invariably characterizing
the appearance of evolutionary processes, not having advanced to the same
extent, much yellow or orange has been retained, so that, therefore, these colours,
where they are present, must be regarded as relics of an older condition and
the butterflies in which they occur exhibit a less advanced stage in the evo-
lutional extension of black. This is, indeed, in complete agreement with the
theory of colour evolution and with what may be observed in this respect also
in other butterflies. But here we find an instance where from a comparison
with some closely allied butterflies the course of this process may be observed
in a particularly striking manner. The orange spot near the apex on the upper
side of the fore-wings in Cyllo Leda L. (PI. XV fig. 30^), wich in some
specimens, however, is larger and clearer than shown in the figure, is nothing
but a vestige of a much greater amount of orange such as still occurs in many
races or species of Cyllo (PL XV fig. 307, ;-). In some this appears in the
form of a fairly broad streak running transversely across the wing surface and
it may be observed here how first in some individuals this streak is invaded
and covered at its lower end by the black pigment, exactly in the manner as
we have observed that it was correctly noted by Alfred Goldborough Mayer ;
it then (PI. XV fig. 3or) encroaches on the remainder which becomes covered
bit by bit until but little is left, as is frequently the case in Cyllo Leda L.
(PI. XV fig. 30c), which, moreover, also gradually becomes covered completely
by the black pigment in the individuals of this species further advanced in the
evolution (PI. XV fig. ^oe). If, in fact, the various stages, as they occur in
these butterflies, were to be reproduced cinematographically in succession, a
correct idea could be formed of the course taken by the process of colour
evolution here being enacted. Of the various stages in this process instances
have been retained in the various species, races, or individuals.

We have here a specially apposite illustration of the course adopted by
the process of colour evolution which, like that of the "streak on the edge
between the red which has remained there and the white which has become

XLVI

the general colour" in individuals of Iphias (Hebomoia) GLAUcrppE Bsd., occur-
ring in some regions, as referred to on page 35 of my treatise o^i the Java
Pieridae, is particularly adapted to afford an insight into this process of change
and the origin of the colour markings on the wings of Rhopalocera.

I have referred above to the broad transverse band which in many Rhopa-
locera devides the upper side of the fore-wings into two parts. Such bands
occur, as I have already noted when treating of the Hesperidae, in many Rhopa-
locera, among Satyridae in Neorina and especially in various species of Debis,
sometimes in both sexes, at other times only in the 9. this band having some-
times resolved itself into dots or stripes, the one and the other clearly indic-
ating various stages in a process of development whose nature, however, is
not clear. In the instance referred to of the butterflies of Cyllo it is, never-
theless, clear that we are concerned with a phenomenon of colour evolution
and it is, therefore, probable that these bands in other Rhopalocera must be
interpreted in the same way. How does this process operate, however, in this
instance? Why is it that with the spreading of the black pigment in Cyllo
butterflies just this band has remained intact so long? One would be inclined
to assume that this spreading took place from both extremities towards the
middle; such a spreading of the black pigment from the apex towards the
base occurs indeed in the 9 of Thyca Belisama Cram., but in Hebomoias and
other Pieridae the original colour, on the other hand, is clearly driven towards
the apex. As a rule the band in question is white or at least of a very light
colour and therefore indicates the occupation of a region where the original
colour has remainnd exempt from the invasion of black and has gradually
faded. But all this does not make it clear to me why this has occured in
these instances.

For an explanation of the origin of the so-called eye-spots, a phenomenon
also of frequent occurrence in Satyridae, a knowledge of colour evolution is
likewise requisite ; this phenomenon, and therefore everything related to it, may
better be discussed in connection with the erroneous conceptions concerning the
so-called seasonal varieties.

To this I will, therefore, now proceed. On pages XIV to XIX of the intro-
duction to my treatise of the Pieridae of Java I have already declared myself
to be opposed to the prevalent misconception concerning the dry and wet-seasonal
forms; this is more especially conspicuous with reference to the Satyridae.
Following in the footsteps of de Niceville, Moore in his Lepidoptcra indica has
adopted a number of well separated forms of the dry and the wet season and
among these the Satyridae play an important role ; others again have followed

XLVII

Moore, Bingham being one of these, while Fruhstorfer goes still much further
in this direction and thus bases on supposed seasonal varieties of Satyridae
various subspecies. But I am convinced this rests entirely on a misconception
even when there may be some substratum of truth. In the same introduction
I have already referred to the fact that many observers have noted the so-called
seasonal forms side by side at the same time in various districts ; even de Nice-
viLLE, according to Moore, admitted that the dry and wet-season forms prevail
during their respective seasons, but are by no means strictly confined to them,
although at the same time he endeavoured to account for this by the circum-
stance that even during the dry season showers of rain occur occasionally and
that the wet season is sometimes interrupted by periods of drought, a matter,
however, not of so much significance that the fact in question, which, more-
over, does not occur now and again only but happens constantly, is to be
attributed to this. Nor does this in any way explain how both so-called forms
are met with at the same time — by no means of rare occurrence — and is,
evidently, based on the erroneous assumption that temporary meteorological
influences are to be regarded as the determining cause. According to Fruh-
storfer, Dr. Martin noted with reference to the Sumatra form of Ypthima
Baldus (Morus Fruhst.) that no dry-season form exists of this but that, never-
theless, the ocelli on the under side of the hind-wings are sometimes strikingly
smaller than usual. With respect to the Pieridae I have published in the work
referred to, some important observations in connection with this subject together
with my ideas concerning the origin of the differences here alluded to. My
observations on the Java Satyridae fully confirm these ideas.

As will be further noted when treating the species separately, these alleged
seasonal forms are by no means distinctly separated but run into each other
through transitions and are, moreover, found frequently side by side. It cannot,
indeed, be denied that as regards colour some greater development may be
observed during the wet season in many individuals of the same species than
is the case during the dry season. But seeing that this is by no means the
case in all individuals, nor confined to the different seasons, it is impossible to
assume the existence of such well marked dry and wet-season forms, as is done by
the above-named entomologists. This difference is only manifested in the manner
indicated by me on page XVIII of the introduction to my monograph of the Java
Pieridae as regards Calledryas Pomona F. ; in the Satyridae it may especially
be traced by a study of the ocelli which are very characteristic in several genera.
For this purpose, however, the origin of these ocelli must first be well understood
and this is only made possible by the knowledge of the process of colour evolution.

The so-called eye-spots or ocelli occur in many butterflies. Before proceeding
to discuss these I would caution my readers against the existence of what I
will call pseudo-ocelli which being confused with the true eye-spots, causes some
perplexity in explaining the origin of the latter since this does not accord with
that of the former. Those designated as pseudo-ocelli by me are simply spots
produced on the spreading of the black at the expense of the original colour,
already partly faded, by the latter crowding in a more or less rounded form
in which the imagination endeavours to trace a resemblance to an eye. To
this may be referred the so-called eye, so conspicuous on account of a certain
development of interference colours in that region, on the upper side of the
fore-wings of the European Peacock butterfly (Vanessa Io L.) which owes
its popular name to that resemblance ; the genesis of this spot is demonstrated
by the less developed forms, produced by breeding, of this butterfly, illustrated
by the famous entomologist Prof. Dr. M. Standfuss in his Handbucli der
paldarktiscJien Gross-Sclnndterlinge. The same applies to the spot near the apex on
the upper side of the fore-wings in Cyllo Leda L , referred to above. For although
these spots also owe their origin to the process of colour evolution their round
shape is to a certain extent accidental and is by no means sharply defined;
in this respect they differ at once from the sharply marked ocelli, generally
round but sometimes oblong, composed of concentric colour rings, and which
have originated in a different way although by the same process; moreover,
they must not be confounded with what I call for that reason pseudo ones.
The true ocelli occur in may Lepidoptera ; some genera, such as Parnassius,
Tenaris, Morpho, and Brassolis, as well as, among Heterocera, Saturnia,
are specially characterized by them ; in a smaller form, but otherwise of an
identical nature, they are frequently met with, inter alia, in some genera of
Satyridae, which leads me to discuss them here. Darwin paid attention to
the ocelli and initiated the confusion between the true and spurious ones.
Discussing the ocellar markings on the feathers of certain birds in his work
The Descent of Man in connection with his ideas concerning sexual selection,
he considers them in relation to those of certain butterflies and especially to
the spurious ones of Cyllo Leda L. Whether these spots and the ocellar
markings in birds are, however, as has been assumed, indeed identical in origin
may, in my opinion, be considered very doubtful; the true ocelli in Lepidoptera
have evidently nothing in common with these ocellar markings of the feathers
in birds. W. Bateson, in his "Materials for the Study of variation" , devoting
a special and otherwise very important study to this subject, and being unac-
quainted with the process of colour evolution, entirely adopts the attitude of
Darwin in the matter.

XLIX

The eye-spots, which I term true ocelli, are characterized, as has been
stated, by various concentric colour rings and, as Bateson correctly observes,
strongly suggesting the notion " that the whole series of rings may have been
formed by some one central disturbance, somewhat as a series of concentric
waves may be found by the splash of a stone thrown into a pool ". Now
the production of colour rings round an excisting centre is not entirely unknown
in Lepidoptera. In my paper Over dc o?ihvikke/i>ig van eenige Javaansche Papi-
lionidenrupscn published in 1888 in volume XXXI of the Tijdsehrilt voor Ento-
mologie I showed, for instance, how in the caterpillars of Papilio Agamemnon L.
and P. EuRYPYLUs L. (Jason Esp.) a ring of coloured pigment is formed
round the base of the spines on the third thoracical segment ; a similar ring
spot is also found in the larva of Ptehogon Oenotherae Esp. round the horny
last vestige of the horn peculiar to the caterpillars of Sphingidae. It appears,
therefore, that where on the body surface a certain induration occurs inter-
rupting or disturbing the normal distribution of the pigment a ring-shaped
accumulation of the latter round the obstruction results. This, moreover,
apparently occurs not only in caterpillars ; the origin of the 8 shaped figure
on the under side of the hind-wings in Callidryas Scylla L. and C. Pomona F.
I considered, when treating of the }ava Pieridae, could only be accounted for
in this way, /. c. by assuming that in certain places of the wings small indu-
rations — probably traceable anatomically — are formed, around which the
pigment present begins to accumulate in ringshaped fashion. The same or
something similar, it may be assumed, causes the production of the ocelli in
the wings of butterflies and a knowledge of the process of colour evolution
strongly supports this assumption. It is, indeed, by no means impossible that
in some genera or species certain anatomical impediments against the normal
distribution of the pigment, as this occurs under the stress of the process of
colour evolution, operate in specific places on the surface of the wings,
corresponding to the above mentioned indurations to that extent. The fact
noted by Bateson that in Satyridae the ocelli occur invariably on one of the
creases or fold marks of the wings between two nerures, /. e. just, therefore,
where such a disturbance would take place, deserves consideration in this
connection, as well as the observation by the same author that such ocelli on
the upper and under sides of the wings, indeed, sometimes occur which completely
coincide but that this is by no means always the case and consequently the
assumed disturbance may possibly occur in one of the two sides of the wings only
which, indeed, agrees entirely with the fact that the process of colour evolution
pursues its course independently with regard to either ot these surfaces. Some
other explanation in this connection may, however, present itself, /. c. that to

7

which I referred when dealing with the Java Pieridae where I applied the term
" persistent " to these spots which I looked upon as being portions of the wing
surface where the original colour had resisted the evolutional invasion of black
much longer than in other parts, the cause whereof must probably be sought
in some special condition of the tissues. However this may be, it is certainly
reasonable to assume that when in the stress of the process of colour evolution
the black pigment endeavours to spread over the original colour and encounters
such a disturbance, induration, or persistent spot, this pigment will be distributed
around it and an island will be formed of the original colour surrounded by
black. We find in fact that the colour of this enclosed centre accords with
this proposition as a rule, sometimes, however, circumstances may arise when
this is not so clearly perceptible. In the ocelli of Parnassii the centre
frequently exhibits for the greater part the original red colour, only just in part
faded to white. In the various genera, however, in accordance with the same
absence of uniformity, invariably to be witnessed in the course of an evolutionary
process, the process in question has doubtless frequently taken a special course.
Thus, for instance, it is remarkable how these ocelli on the hind-wings of
Parnassii correspond in form and manner of disposition to the black spots on
their fore-wings ; the former are evidently just such spots as the latter in which,
however, the black pigment has not been distributed to the same extent and
where, consequently, in the formation of the ocelli the black pigment has not
exercised as much stress as has been the case in other genera. In some
Satyridae the course of this process can be clearly discerned. The accom-
panying illustration (PI. XV fig. 506) gives an enlargement of the ocellus, indicated
on PI. XV fig. 30rt by number 5, on the under surface of the hind-wings of
Cyllo Leda L. a clear white centre will there be observed round which a
fairly wide ring of intense black has gathered while outside this again a narrower
yellow ring has been formed. Now this centre is doubtless the vestige of the
induration or persistent spot where the original colour was not — or at least not
completely — covered by the black pigment, but where this has gradually faded
to white in conformity with the rule of the process of colour evolution or has
already been destroyed and replaced by scales simply filled with air which, in
view of the intensity of this white, I do not consider improbable.

For this reason it not unfrequently happens, as Bateson observes, that the
centres of ocelli are deficient in or destitute of scales. Around this centre the
black pigment, which was checked in its even distribution, has become accumu-
lated, thereby causing the production of a fairly broad ring of black much more
intense than the black pigment occurring elsewhere in the colour of the wings
in this species. This intensity clearly indicates that an accumulation of this

11

pigment has taken place here and this accounts for the narrow yellow ring again
surrounding this black. For the black pigment, which there as elsewhere in
the wings, had first covered the original colour, has, under the said stress,
thence pressed towards the centre, where consequently, it has increased most
and has thus produced the intense black colour but at the same time it has
retired from the place where it occurred first and in this manner has caused
the original colour — now faded to yellow — which formerly it covered, to
reappear as a narrow ring.

Conceived in this fashion the colour arrangement of this ocellus is easily
understood without having recourse to chemical action to which Bateson draws
attention. Providing, of course, one is acquainted with the phenomenon of
colour evolution and has been taught to interpret the distribution of the black
pigment over the original, or subsequently faded, pigmental colours in a manner,
which in fact reminds us of the flowing forward of a liquid, as the expression
of the stress of this evolution.

Since this is the case with this ocellus it seems reasonable, as regards other
true ocelli, to explain the cause of their origin in a similar manner even though
the course of the process of colour evolution in every species, or at least in
every genus, may not be so clearly traceable and adventitions circumstances,
such as the appearance of interference colours, may impede its investigation.

Other ocelli exist which must also be classed with the true ones but which,
in part at least, have presumably originated in a different way. I refer to
those occurring in many caterpillars of the Sphingidae, and which have played
an important role in the fantastic theory of mimicry. As far as I have been
able to find out they sometimes start from a colour spot on each segment,
the remnants of the longitudinal lateral stripes occurring in earlier stages of
development and which subsequently either disappear entirely or at least strongly
diminish ; these remnants appear to assume a round shape during the growth
of the caterpillar and in this manner form large round ocelli on such articu-
lations which increase inordinately in size. The course of this development,
however, requires further and special investigation.

The ocelli, first referred to, are of great importance for a correct judgment
on the question of the alleged seasonal forms. For it is asserted that these
markings only occur in the wet season while in the dry-season forms of the
same species they are absent, and that consequently the forms of the dry and
the wet season can be distinguished by the absence or presence of the ocelli.
It is true this assertion is based upon a certain amount of correct observation ;
but this has become the subject of much wild speculation so that in the end
much misconception has arisen. Much stress has frequently been laid upon

LII

the excessive variability of the ocelli but in the interpretation of this phenomenon
little progress has been made, because, as 1 have already pointed out, so little
attention has been paid to the fact that every animal organism is subjected to
continuous changes of form, appearing in a greater or lesser degree in various
individuals, and that changes of colour come under this category, from which it
follows that where in consequence of colour evolution special forms have been produ-
ced, as is the case in butterflies with ocelli, these changes are particalarly striking.
Individual differences exist invariably ; no two butterflies, any more than two men,
being completely alike ; some difference in colour or marking is, as a rule, to be noted
also between two butterflies of one species. With regard to the forms created by the
evolutional stress of a special process of change of form such individual differences
must consequently occur. Its greater or lesser progress is always governed by
the peculiar constitution of each individual whose origin is never identical with
that of the other individuals and also by the very different individual suscep-
tibility for this special stress of development. These must, therefore, of neccessity,
reflect the fact whether the process of the evolution occurring here has more
or less advanced in some individual and, therefore, the degree of its develop-
ment. In one the ocelli will consequently be larger or more numerous than
in another; in some they will exhibit a particular distribution while in others
again they will be scarcely apparent or altogether absent. This is, in fact,
what a study of the ocelli brings to light. In size they differ considerably
in individuals of the same species. Bateson, on page 295 of his important
work already quoted, figures four specimens of Satyrus Hyperanthus L. all
differing in the number and size of the ocelli ; three of these he notes as having
been taken on the same day in the same district in England, " so that here
no question of seasonal or local difference is necessarily involved". Indeed,
in view of the present tendency of basing so-called sub-species and even species
purely on such differences, this observation may usefully be reiterated. Besides,
how insignificant actually are these differences. The specimen marked n*^. Ill is
the only one of any importance because practically no ocelli are to be seen
in it. This form occurs very little and is evidently one in which the colour
development has been much retarded ; it clearly corresponds in the present
species, producing but a single generation in a year, to the form Levana in
the Vanessa species yielding two generations each year. In Cyllo Leda L.
such specimens occur not infrequently and here it may be observed how they
arise through retarded colour development and grade into one another with
more advanced forms. As regards the other three, which are in a further
advanced stage of development, there is only a slight difference in respect to
the size of the ocelli; in the one which is most developed, marked IV, by the

LHI

side of two fairly large occelli in the centre of each of the hind-wings a third
appears, being confluent with one of the former. The latter phenomenon is
one of evolutional change of form not of infrequent occurrence where the ocelli
are strongly developed and often to be observed on the lower surface of
Lycaenidae, which has been noted as such by Prof Dr. Couvoisier of Basle
in his paper " lleber Zeic/nmiigs-Abcrra/ionrn bci Lycaeniden" {Zcihchrift fi'ir
Wissenschaftliche Insektenbiologie, igof). This confluence of ocelli may plainly
be observed in many Ypthima species ; in some specimens they may thus be
seen as two ocelli joined side by side, being already united where they are in
contact ; in more advanced stages they are amalgamated into one ocellus which
is still elongate in shape, however, while next this becomes more and more
rounded in outline until finally a single large ocellus results. This does not,
however, contain a single, but two nuclear points, the nuclei of the two original
ocelli; which, therefore, demonstrate the fact of this confluence. Not always,
however, is the presence of two white dots in such an ocellus due to this
origin. In Neorina Chrisna Westw. a large ocellus may be observed on
the apical portion of the fore-wings on the upper surface. This ocellus is of
an intense black with yellow border and sometimes with a white nuclear dot in
the centre of the black. Now two larger white dots invariably occur, not in
the centre, but in the upper and lower part of the yellow border. Consequently
they are certainly not nuclei. Moreover, they are not placed within the ocellus
proper, but why such white scales should appear just in these places remains
unsolved. The difference in the occurrence of these ocelli in butterflies of the
genus MoRPHO, quoted by Bateson, is also similar in character. The species
of this genus mentioned by him are closely allied, and they are, therefore,
subjected to the same evolutionary processes ; in which connection sometimes
a few more ocelli occur in one individual than in another.

These individual differences in size or number of ocelli occur also to a
considerable extent in the Java Satyridae, such as Mycalesis Perseus F. ; this
is most conspicuous in Cyllo Leda L., so common in that island, and a careful
study of this butterfly clearly demonstrates that in this case they are purely
phases of colour development, and that of some unlimited or irregular so-called
variability there can be no question. In Cyllo Leda L. the lower surface
exhibits on the fore-wings 4 and on the hind-wings 6 ocelli which I have indi-
cated with numerals in the accompanying illustration. Of those on the fore-
wings n^. 3 is the largest and the others are mostly very small. On the
hind-wings n''. 5 is the largest, n*'. 7 and n". g being next in size and the
remaining three considerably smaller; n**. 10 is sometimes duplicated and n". 8
mostly placed further towards the outer margin than the others. Individual

LIV

differences in this respect occur in Java only exceptionally ; in races or closely
allied species in the east of the Archipelago, in which the ocelli are invariably
larger, greater differences in this respect are met with, although the character
of these spots remains the same. The stage of development ot these ocelli, however,
is not identical in the several individuals, yet they are invariably in the same
mutual relation, n*^. 5 on the hind-wings being always the largest, n^. 9 coming
next, etc. They thus occur in the same relation in various stages of develop-
ment ; at the most advanced stage such an ocellus has the appearance in
miniature of the enlarged illustration given on PI. XV fig. ^06, but in less advanced
condition these colour rings are only partly distinguishable or not at all ; at a
still earlier stage they appear as yellowish dots or specks, and in proportion
to the lower stage of development they decrease in size ; the larger being reduced
to specks while the smaller disappear entirely but all in the same mutual ratio ;
if only one yellow speck remains visible it is found at the spot where in case
of complete development the largest of the ocelli — n^. 5 on the hind wings —
occurs. The action of a regular process of colour mutation is clearly mani-
fested here but there is no question of irregular changing caused by so-called
variability.

In a species like Cvllo Leda L., where the degree of its colour develop-
ment as well as that of its ocelli can thus so easily be observed and which
at the same time occurs so generally in various districts and at various times, the
question whether or not the existence of ocelli characterizes the wet-season
variety, ought to be capable of being demonstrated.

Now from the examination of a great number of specimens, available for
comparison, it is perfectly clear to me that such is not the case. It is true
the best developed ocelli occur in the wet season while ocelli met with in the
dry season are always less developed and mostly reduced to specks, wholly or
partly, or they may also be wanting in part or even entirely. But transitions
occur and, moreover, the rainy season in Java gradually passes into the dry
season while, finally, butterflies with very slightly developed ocelli are by no
means unfrequently met with in the middle of the wet season. It is, therefore,
clear that the process of colour evolution, in the way this, too, decidedly mani-
fests itself in the ocelli, operates in Cyllo Leda L. most strongly during the
wet season, but presumably because, above all, the food is more nutritious at
that time — since in the wet season the whole vegetation is much more robust
and the lar\'ae then met with have a much stronger appearance — and not
by reason of any direct specific meteorological influence. In the course of
this evolutionary process the individual susceptibility too undoubtedly plays an

LV

important part and its progress in relation thereto in certain individuals may
at the height of the wet season be almost or entirely suspended. Notwith-
standing the fact that, therefore, the wet season undoubtedly exercises some
influence, if only indirectly, it is by no means clearly reflected in the appearance
of the ocelli and the assumption of the existence of sharply defined seasonal
varieties as the results is unwarranted since these do not, in Java at least, actually
exist; which, moreover, completely accords with what I have ascertained in this
respect in the case of the Pieridae. The study of the ocelli in other Satyridae,
such as Mycalesis Perseus L. and species of Ypthima, leads to the same result.

The larvae of the Satyridae give occasion to some observations. In the
genus Ypthima the same colour dimorphism of green and brown larvae appears
to exist as was the subject of my careful investigations in the Sphingidae
larvae in Tij'dsclirift van Entoniologic, XL, and subsequently observed by DiErzE
{Deutsche Entom. Zeiiims;, Iris, XIV), in the larva of Tei'Hrolystis Druentiata.
As has been demonstrated by the study of the Sphingidae larvae, it can only
be a question of a regular, and therefore not an accidental, but evolutional
colour change, in short, of colour evolution. A striking fact, moreover, in
connection with these larvae — at least in so far as they are known, since
our knowledge in this respect is still very incomplete — is that whereas the
heads of the European Satyrus larvae, according to the illustrations given,
appear to be round, those in the genus Ypthima, while being round likewise,
are provided with two small spiny processes, while in the genus Mycalesis the
heads become bifurcate, and in the other Satyrid genera Cyllo, Debis, Orsitriaena,
as well as in the genus Elymnias, probably belonging to that family also, the
branches of these processes develop into true horns to which the processes
on the posterior extremity correspond in formation, terminating as they do in a
more or less developed bifurcation. The same is to be observed in a more
advanced stage in the Nymphalidae amongst which in the genus Neptis the
heads of the larvae are still more decidedly bilobed but in the other genera are
invariably provided with projections or so-called horns of various shapes, while
presumably in correlation with these various projections or spines are also
produced on the back and sides. It appears to me that this affords strong
confirmation of the suggestion that the family Nymphalidae has developed from
that of Satyridae which deserves the more consideration from the fact that some
systematists desire to place the genus Amathusia, considered to pertain to
Nymphalidae, for other reasons among Satyridae and in this respect, therefore,
a difference of opinion exists which is invariably the case where transition
forms occur which clearly demonstrate that in fact no definite limits exist.

LVI

Concerning so-called mimicry phenomena, where these are supposed to occur
in Danaidae or Satyridae, I will briefly refer to them without, however, deeply
going into the subject, because, as I have already stated in the introduction
to my monograph of the Java Pieridae, I do not consider it a subject for
scientific discussion but merely a matter of belief against which reasoning is of
no avail and which w'ith its supporters will probably die a natural death. In
1 910 an attempt was made again to bring the subject on the carpet and this
was undoubtedly one of the principal reasons which led to an international
entomological congress being held, all the principal supporters of this theory
having turned up in a solid body. I was kindly invited to take part in the
preliminary deliberations concerning this congress and I was also offered the
chairmanship of the section in which the subject of mimicry would definitely
come up for discussion, but while fully appreciating this courtesy I have thought
it expedient not to accept the invitation. In the first place I have long since
arrived at the conclusion that for the discussion of scientific subject congresses
are altogether unsuitable. Differences of opinion on really scientific subjects
can only be discussed with advantage by those who are acquainted with the
pros and contras and who can adduce their arguments against a theory they
feel bound to contest, in a detailed and thorough manner; but with verbal
discussion this is impossible since the time for carefully weighing the adversary's
contention is lacking, and the proper place for one-sided propositions undoubtedly
is in scientific periodicals where they can be considered quietly and at leisure.
Congresses are the product of our democratic age, a kind of popular assemblies
where superficiality sits enthroned, but such discussion cannot be reconciled
with the conception of serious knowledge. Thus the question of mimicry can
only be solved by serious scientific investigation. In the second place I consider
such investigation no longer required since this has already taken place. The
propositions in connection with the subject have been refuted by numerous
writers after an examination, as careful as it was comprehensive ; against the
facts on which the latter have based their opinion the supporters of the theory
of mimicry have never adduced any serious arguments; the opponents of this
doctrine are therefore entitled to consider them incapable of discussing this
theory in a scientific manner and to look upon them as supporters, not of a
scientifically formed doctrine, but of a creed, and against a creed no argument
avails. Everything they continually bring forward is nothing but the repeated
marshalling of new facts which under the dominance of their creed they represent
as mimicry but without ever adducing anything of the nature of proof to
support their explanation or to refute the contending interpretation of others.
It could not, therefore, be reasonably expected that a discussion of the subject

LVir

at that congress should lead to any other result and it is on that account
that I did not desire to take part in it.

Although following the systematic arrangement of the late Snellen I have
found it necessary somewhat to modify it in connection with certain points which
I would further have discussed with him if he had lived. After consulting
Mr. Fruhstorfer I have had to include the species Ypthima J areas de Nic,
whose occurrence in Java was unknown to Snellen ; moreover, after careful
study of much more extensive material than was available to him I have
excluded from the present work the two species Ypthima Nigricans Sn. and
Cyllo Zithenius Herbst. The advice of Mr. Fruhstorfer and the inspection
of his extensive collection have been of great service to me.

POSTSCRIPT.

The issue of the present volume has been considerably delayed. Although
otherwise completed at the end of 191 1, the printing of the plates has encountered
many difificulties and consequently publication could only be effected in
1913. As a result, the report of the proceedings of the International Ento-
mological Congres held in 1 9 1 1 having reached me in the course of
191 2, I was able to acquaint myself with its conclusions on the subject of
mimicry, discussed by me at the end of the Introduction vide page LVI.
This circumstance induces me here to add a few observations on the subject.

I there stated that for two reasons especially no useful purpose could be
served by such discussions. First, because, as I have already stated repeatedly,
the subject in question appears to me already to have been shelved in a scientific
sense and will, therefore, probably die a natural death ; so that the fact that a
few adherents continue to support it needs no further consideration. Secondly,
because experience has shown that on their part no scientific reasoning, no
adduction of proof or fundamental refutation, is to be expected but only records
of fresh observations which, having been explained from their one-sided point
of view, are regarded as confirming their theory which, however, with a
different interpretation is by no means the case, while they fail to refute the
latter. The report of the Congress, in my opinion, completely confirms my
attitude in this connection.

8

Lvm

I stated at the time that the inception of the Congress had created in my
mind a strong impression that the idea of energetically taking up the defence
of the theory of mimicry had played an important role in it. And what
actually took place? Messrs. Dixey, Poulton, Trimen, Jordan. Rothschild,
and Marshall, being some of the oldest and keenest defenders of the theory
of mimicry, attended the congress in Brussels ; of these three spoke on the subject.
To me this seemed to be a cruisade conceived with deliberation, a serious
attempt to stifle once and for all the biological heresy which during the last
few years had become more and more pronounced. Now what was the effect
produced? What was it that the whole remainder of the entomological world,
appearing at this congress in large numbers and acquainted with the fact that
the theory of mimicry would intentionally be brought on the tapis, considered
it worth while to adduce in support or against it? A single speaker, besides
the three forementioned, addressed the Congress on the subject; one other
speaker made a few unimportant observations in opposition to it. With all
this ostentation this was all the result produced and I consider, therefore, I
was justified in asserting that the theory of mimicry is moribund and that in
the domain of science it has ceased to be of any account.

From the beginning the supporters of the theory of mimicry have continually
adduced facts, sometimes isolated, sometimes in whole complexes, observed by
the same person, which would, according to their interpretation, establish the
existence of mimicry. Later and more accurate observation, however, has
demonstrated the invalidity of this reasoning. Sometimes it was proved that
the alleged observation had been of an extreme superficial nature or had been
carried out under suggestion of the preconceived notion of mimicry. In other
instances, whereas the observation itself was indeed accurate, it was found that
its interpretation as mimicry originated from the same idea, and that this could
quite well be explained in a different manner, but could not in itself demonstrate
the existence of mimicry and was only then to be attributed to it if the postulates
of this biological theory had, at the outset, themselves been proved, which,
however, has never yet been done. In this manner the so-called facts of
mimicry in South America of Bates, Fritz Muller, and others, those in
Africa — especially concerning Papilio Dardanus Brown — and the numerous
Indo-Australian instances, chiefly contributed by Wallace, have been brought
forward. Further investigation, in which, chiefly as regards the two latter, I
haven taken an active share, has refuted all this to such an extent that the
value of the theory in question has greatly depreciated with the majority of
serious biologists, if indeed it has not been completely demolished. In spite
of this the supporters referred to, again present the selfsame doctrine, not

LIX

indeed accompanied by arguments absolutely essential to it but simply, as on
former occasions, by an appeal to alleged facts.

Some of these are new and so far in themselves irrefutable. For this purpose
their observations would first have to be controlled. Even though the bona fides
of Dr. Wiggins need not be questioned he is clearly what Mr. Jordan terms E.
Haase, an ardent believer in mimicry, and that little reliance is to be placed upon
observations made under this obsession is known in numerous instances. The
accuracy of these observations requires to be controlled by other independent
observers who are sceptical on the subject ; not until then shall we be in a
position to estimate their value. It may, however, even now be assumed that
these facts, even though they be correctly observed, can assuredly be explained
without having recourse to mimicry. In reality they exhibit no character other
than that pertaining to many other facts observed elsewhere, which clearly
proves that a similar mutual resemblance in identical circumstances may indeed
occur through a similar line of development being influenced in the same
manner, without there being any question, however, of mimicry. Resemblance
does not constitute mimicry but resemblance considered under the suggestion
of mimicry creates alleged instances thereof

While once more discussing the subject of mimicry I propose here cursorily
to subject the proceedings in this connection at the said Congress to a critical
examination ; more especially to demonstrate that on this occasion the requisite
arguments were totally lacking.

On August y^ 19 lo Dr. Dixey gave a lecture on "Mimicry" . He does
not, however, commence, as a basis of his observations, by pointing out the
scientific necessity of acknowledging the existence of this alleged biological
phenomenon, in other words by indicating the scientifically sound, or at least
most credible, grounds on which this necessity is based.

To this he makes no allusion whatever, he simply assumes its existence a priori
and attempts to extend the knowledge of this phenomenon by connecting with
it various simple resemblances observed. He asserts, in fact, that the said
resemblance are so numerous and exhibit such peculiarities that it will not do
to attribute them merely to accident which, although nothing but a vague
expression, certainly does not sound incredible but which can in no case imply
that it can only be accounted for by the existence of mimicry; a great number
of similar facts have indeed long since been explained in other ways. He next
gives a detailed description of such cases of resemblance which, by the way,
contain inaccuracies or assertions which have long since been refuted to which,
however, no further reference can here be made. One is struck, however, in

LK

this connection with his want of knowledge as to what in this domain has been
investigated, discussed, and refuted by many entomologists, other than English,
which, naturally, entails a very limited judgment on the subject. He thus
arrives at the following ten propositions, which I will quote here.

1. The cases of resemblance between distinct kinds of insects are very
numerous — too numerous to be accidental.

2. These resemblances are to a very great extent independent of affinity.
Some of the most striking are those between insects of different orders.

3. They are peculiarly liable to occur in insects of the female sex.

4. They are, speaking generally, found only between the inhabitants of
the same region.

5. They may affect one phase of a seasonally dimorphic insect differently
from the others.

6. No structure or detail of organisation is involved in these resemblances
exept in so far as a modification therein may assist in producing a superficial
likeness in aspect or behaviour.

7. In the production of these resemblances the same effect is often brought
about by different means.

8. Every transition exists between a likeness which is so remote as to be
fairly disputable, and a resemblance which may even deceive a skilled observer.

9. In some cases there may be great disparity in point of numbers
between the forms linked together by community of aspect. In other cases
the numbers may be nearly equal.

10. The combinations of two or more forms resembling- one another are
in many cases not isolated, but are often connected with other combinations
by a more or less complete series of gradations.

Dr. DixEY calls these facts, but it is clear that they are not facts in a
scientific sense ; they are purely propositions recapitulating what according to
his notion appears to result from^ certain observations; propositions, whose
accuracy is not only considerably to be discounted, but which, moreover, are
of far too uncertain and subjective a character to allow scientific — /. e. logical —
conclusions to be deduced from them. But this course is not adopted by
Dr. DixEY. He simply proceeds to discuss the theory of geogrophical influ-
ences, the artist theory of Mr. Abbott H. Thayer, and the opinion that these
resemblances are to be attributed to internal causes, winding up with the decla-
ration that he associates himself with the opinion of Bates and Fritz Muller.
But while his arguments against the artist theory contain a great deal of truth,
this much cannot be said, as regards the remainder. His reasoning practically
consist of a priori adducing the well known Darwinian theories of utility and

LXI

natural selection, neither of which, however, is scientifically established ; the old
assertion that the fact of some butterflies being distasteful to birds constitutes
an argument in favour of the theory of mimicry, he also considers as practi-
cally proved. Yet this has in reality been long since sufficiently refuted. In
conclusion he declares the observations thus adduced to aeree well with his
" facts " quoted supra and he professes to demonstrate that they receive strong
support from these " facts " . We may, however, quite well leave out of account
what Dr. Dixey adduces in this connection since neither his " facts " nor the
observations in question have any scientific value. This applies equally to the
whole of his lecture, which is characterized as much by a lack of precise
logical arrangement as by a complete absence of the requisite critical faculty.
The theory of mimicry — like any speculative doctrine whatever — does not
lend itself to demonstration by means of a mass of ill-digested observations. Now
what cannot be proved adequately may at the utmost be acknowledged by
science as hypothesis but even this must rest on a sound basis. When any
phenomenon cannot be explained in any other manner a conjecture so soundly
based and, therefore, very admissible may in this respect pass muster as
hypothesis. As soon, however, as any other equally admissible solution presents
itself such hypothesis falls to the ground.

On the same day Prof. E. B. Poulton gave an account of Dr. C. A.
Wiggin's Researches on Mimicry in the Forest Butterflies of Uganda (1909).
These observations have reference to the joint occurrence ot some species or
forms of the Acraeine Genus Planema with butterflies belonging to other
genera, which at the same places strongly resemble one another, from which
fact various considerations concerning the existence of mimicry are deduced.
But in all these cases it is assumed a priori that the butterflies in question
represent models and mimics respectively ; this should, however, be proved first
for until this has been done these observations are without foundation. No
attempt, however, is made to establish this proof. Only from a confused
mental process the idea can arise that these resemblances, in themselves, however
remarkable they may be, have of necessity been produced by mimetic action.
From these alone this conclusion can never be drawn. Prof. Poultox, there-
fore, has likewise failed to adduce any proof of the theory of mimicry on
this occasion.

Mr. J. K. Jordan, also on the same day, discussed " The systematics of
certain Lepidoptera ivhich resemble each other, and their bearing on general
questions of evolution " .

Lxir

This discourse was decidedly on a higher level than the preceding, and
contained a great deal worthy of attention. As regards the theory of mimicry,
however, it was greatly lacking in force as well as in logical argument. Instead
of the latter he employed propositions devoid of scientific foundation — nothing
but hollow phrases — therefore a priori reasoning, while he entirely omitted
to make the requisite examination of the ways in which the opponents of the
theory in question explain the phenomena which are indicated as mimicry and
their refutation consequently was not attempted. Whether it be owing to
inability or to prejudice, when no cognizance is taken thereof the treatment of
the subject becomes of necessity so infomplete that no value can be attached
to it. Mr. Jordan professes to find a proof of the existence of mimicry in
the circumstance that in several cases where the models belong to different
species their mimics, on the other hand, are simply forms of one single
species, these forms not possessing the characters of incipient species and
their origin, therefore, being unexplainable otherwise than by natural selection
on account of their mimetic protection. This reasoning, therefore, likewise
proceeds a priori from the proposition that these butterflies are each others,
models or mimics respectively and that their resemblance cannot be attributed
to other causes. The correctness of this proposition should, however, be first
established scientifically and its proof, therefore, produced. But this again is
not attempted by Mr. Jordan and his observations are, therefore, valueless.
Mr. Jordan looks for the force of his reasoning, to a great extent, in the
Darwinian theory that every new variation occurs because of the utility which
accompanies it and in that of natural selection. Conceived in so general and
far-reaching a sense, however, these theories are not tenable.

Whosoever observes the phenomena of nature from an independent stand-
point and without being influenced by prevalent theories is bound to arrive at
the conclusion that a certain number of variations are useful to the creatures
in which they occur, while others must be considered detrimental or at least
in no wise to satisfy a specific want. It is by no means to be assumed, there-
fore, that new forms of necessity are referable to their utility. Resemblances
may, consequently, very well occur without being of any advantage. The
doctrine of natural selection does not possess the value either which the sup-
porters of the theory of mimicry attach to it. This doctrine rests entirely on
that of the struggle for existence, it is, in fact, a natural corollary of it but
only in the original Darwinian sense, according to which the whole of animate
nature was regarded as waging a fierce mutual contest which was supposed to
lead to the sole survival of the fittest. Soon, however, the error of this con-
ception concerning this so universal, exclusive, and fierce character of nature

LXIII

became manifest. Darwin himself in his later years toned down his views in
this respect and others have continued this process so that at present the expression
" struggle for existence " is not meant to imply much more than a general
term for the obstacles of all kinds with which every being has to contend in
the course of its life, without invariably being of so fierce and aggressive a
character. The doctrine of natural selection is not, however, based on the
words " struggle for existence " but on their meaning and when, therefore, this
no longer corresponds to the old drastic definition, this fierce, agressive
character implied by it, can no longer be assigned to natural selection, nor
the great influence originally attributed to it and assumed to play such an
important role in the theory of mimicry. This assumption nevertheless retains
currency in this theory although being devoid of scientific basis.

On the other hand Mr. Jordan takes no account of the views I have
published several years ago concerning colour evolution ; besides the theory of
mimicry he discusses, it is true, some different views as to the alleged mimetic
phenomena but is silent on this subject, notwithstanding the fact that it is just
this theory which affords quite a simple explanation of all the cases of resem-
blance which he considers so remarkable, although it must be admitted that
his argument would certainly thereby be considerably weakened. In mimetic
phenomena colours play the chief role, but as long as one only possesses
confused ideas concerning their existence — as long as one does not grasp,
for instance, that pigmental colours form a physiological unit governed by
definite rules of development — one will be unable to form a clear judgment
on these phenomena. A study of colour evolution alone enables us thoroughly
to understand the nature and progress of evolutionary phenomena and will
induce any one who reflects on the subject to come forth from among the great
host of naturalists, whom I had in mind when I repeatedly stated that since
Darwin's time one meets continually and everywhere the term "evolution",
but that this is by no means the case with persons who possess some clear
notion as to its nature. It is only through this study, moreover, that one
becomes acquainted with its exceedingly uneven progress and that one thus
acquires a much truer insight as to the nature of variation, which plays so
important a role in present day biological problems. Thus the nature of
dimorphism and polymorphism will be understood in quite a different sense than
is mostly the case at present. A new factor has arisen in the scientific study
of biological phenomena and amongst these that of mimicry; this factor has
to be refuted or admitted but where it is simply ignored no claim to a scien-
tific handling of the subject can be made.

While, therefore, fully admitting the justice of several observations and

LXIV

views of Mr. Jordan and being, moreover, entirely on his side on some points,
such as the rejection of the modern mutation theory, I am bound to declare
that I am unable to attach any scientific value to his opinions on the subject
of mimicr}' and that his lecture has by no means established the justness of
this theory.

On August 4''' Mr. W. Schaus addressed the Congress, the title of his
discourse being " A qiioi sert le Mimetisvie ?" The speaker mentioned a great
number of lepidopterological observations made, in his capacity of field natu-
ralist, during a long stay in South and Central America, in connection with
the various assertions which have been made on the subject of mimicry;
observations with reference to the alleged facts — to which great weight is
usually attached in this respect — so much at variance that their connection
has completely dissipated his former belief in the value of so-called protective
colours. His communication on the subject was indeed of great importance.
Primarily because we do not meet here with such loose, isolated — and mostly,
therefore, as inaccurate as incomplete — observations which are so frequently
made by mere amateurs or zoological globe trotters possessed of but very
little sound lepidopterological knowledge, but with a deliberate expos6, the
result of observation during many years with a thorough knowledge of the
butterflies to which it has reference, which on that account deserves much
consideration, but also because these observations tally in so remarkable a manner,
even to minute details, with those carried out by myself and others in like
manner with reference to the Indo-Australian fauna, which have for the most
part already been published ; observations which from a believer in the theory
of mimicry have turned me into an antagonist. Such agreement of opinion on
this subject, evolved quite independently, having reference to a different tropical
fauna, and based on similar knowledge and experience, is surely of no little value.

The statement, inter alia, by Mr. Schaus, that on several occasions he had
witnessed butterflies with so-called protective colours being attacked nothwith-
standing by enemies. Prof Dr. Seitz met by the observation that in order to
contest the theory of mimicry it is not sufficient to prove that a protective
colouring does not protect against all enemies but that it must be shown that
this would not avail against a single enemy and he quotes some instances
where, according to him, this is supposed to be the case. Prof Ssrrz thus
completely departs from the canons of proper scientic discussion. The burden
of proof would lie upon any one asserting that a specific colour affords pro-
tection, even if only against certain enemies, and it is only when this has been
demonstrated that the antagonist of this theory should show the fallacy of the

LXV

said proof. But this has hitherto never been attempted, nor does Prof. Seitz
attempts this as regards the cases cited by him ; because resemblance exists
protection, according to him, must be assumed. The facts observed by
Mr. ScHAUs, however, prove the invalidity of this assumption, and the statement
by Prof. Seitz in opposition to them is therefore, of no value.

Mr. Roland Trimen expressed astonishment that it is possible to deny the
existence of mimicry in view of such instances as are provided by Papilio
Dardanus Brown. This naturalist appears to be unaware that these supposed
instances have been refuted by me long since. With regard to the statement
by Mr. Schaus, that only in exceptional cases has he observed birds attacking
butterflies, he does not consider it improbable that birds prey on butterflies
mostly in the early morning before the latter are on the wing and that for that
reason this is not frequently observed. He further makes a couple of obser-
vations in which simply on account of resemblance the existence of mimicry is
again a priori assumed.

The Hon. W. RorHSCHiLo mentioned a couple of cases of resemblance
which in his opinion could only be explained by mimicry.

Prof PouLTON was unable to agree with either M. Abbott H. Thayer or
Mr. Schaus in their interpretation of the subject of mimicry since a number
of authenticated observations had convinced him of the contrary.

Dr. DixEv stated his opinion that observations such as those of Mr. Schaus
are not irreconcilable with the theory of Bates and Fruz Muller.

Mr. Marshall finally quoted several instances in which birds had been
observed to prey on butterflies. He appears to forget, however, that if this
indeed possessed the significance attributed to it on behalf of the theory of
mimicry, it would not happen in a few instances only but would be of universal
occurrence and this, in view of all the statements that have been advanced against
this assertion, is evidently not the case.

The latest on this subject which has come to my knowledge is a commu-
nication by A. M. Banta in an article entitled : " T/ic distaste fulness of Angsia
Plexippus" and published in N'atiire of May g'*^ 191 2. It is to the following
effect : " For a number of years trained experts of the United States Depart-
ment of Agriculture have been engaged in determining the food of native birds
by examination of the contents of thousands of bird's stomachs collected in all
sections of the country and in all seasons of the year. The bearing of these
findings upon the question of butterflies as food for birds has recently been
summed up by one of these experts as follows : " Four records of birds eating
butterflies are all that are afforded by the records of the examination of more
than 40000 stomachs in the Biological Survey".

9

LXVI

Further discussion did not take place because the time had arrived for the
members of the Congress to pay a visit to the Brussels Museum of Natural
History, which was in fact far more useful, for the discussion called forth by
the discourse of Mr. Schaus was really of little account.

To sum up, only an unfavourable judgment can be pronounced in general
with regard to the views brought forward at the Congress in defence of the
biological superstition which is called mimicry. This cruisade has evidently
miscarried completely. Doubtless its ardent believers will still uphold this theory.
At the second International Entomological Congress held at Oxford they have
returned to the charge in the same manner. It will doubtless long continue
to supply the need for zoological romance in popular periodicals but by the
serious, scientific entomologist it can no longer be acknowledged.

M. C. PlEPERS.

D A N A I D A E.

Genus EUPLOEA F.

EuPLOEAs are slowly hovering, not high-flying, butterflies settling on flowers
and seeking mostly shady places, some, such as E. Leucostictos Gmel., frequently
entering dwellings. They are met with, moreover, both in the plains and in
the mountains, being found even at an altitude of 2000 M. ; some species
occur everywhere while others are restricted to definite areas, which may probably
depend on the presence of the food plant. Among Euploeas also some speci-
mens are considerably smaller than others. According to Moore they belong
in Ceylon to the migratory butterflies. An observation made in December 1885
by Mr. T. Ottolander — already communicated by me some time ago —
is to the effect that in the province of Besouki, Eastern Java, he witnessed the
migrating of Euploea Leucostictos Gmel. on several days between 8 and 12
o'clock in the forenoon when the sun shone but no high wind prevailed.
Mr. G. A. ScHOUTEN likewise noticed the migrating of two species of Euploea at
the same period near the lake Ranau, in the province of Palembang, in Sumatra.
I have never myself observed the migrating of Euploeas in Java, although, at
the places in that island where I resided, Euploea leucosticos Gmel. was very
common ; nor have I received any confirmation of the fact from any other
observers and I do not deny that to my mind this renders the accuracy of these
observations somewhat doubtful. May it not have been that these Euploeas
simply happened ,to be particularly numerous at these spots without, however,
all flying in the same direction, a process characteristic of the phenomenon
usually indicated by the term " migration " }

The (5, on being taken in the hand, protrudes its genitalia in the form of
long yellow processes. I have never noticed them, however, to emit by this
means the strong scent which Dr. Fritz Muller states to have observed in

Brazilian Danaidae. According to Fruhstorfer all Euploeas emit a strong

smell, sometimes resembling the scent of flowers but frequently very repulsive,

the latter being the case especially in E. Midamus L. Longstaff, however,

took two c? and four 9 of this species at Hongkong, of which only one 9 had

such a powerful acetylene-like odour as to be clearly perceptible when in the

net. Except in a couple of instances where evidently these butterflies had

recently been feeding on excrements and clearly carried the smell with them,

I have never been able to notice that they emitted any particular smell, not

even the species just referred to, although I have certainly handled hundreds

of it ; I can scarcely believe, therefore, that they should be protected against

the attacks of birds by an unpleasant smell or taste. De Niceville, it is true,

makes this assertion, but this entomologist always labours strongly under the

suggestion of mimicry and observations in relation to their obsessions by such

persons are not much to be trusted. When treating of the Hesperidae, we have

already seen an instance of this in the case of the larva of Hidari Trava

Moore. But such insects, having only a very attenuated body together with

large inconvenient wings, can hardly be a favourite food with insectivorous birds.

Moreover, on two occasions I have seen Euploea Rafflesi Moore being caught

and eaten by a bird (Edolius). The tenacity of life in these butterflies is very

striking, since it is next to impossible to kill them, like other Rhopalocera,

by compressing the thorax; they certainly remain motionless a short time but

soon recover completely. On the other hand a drop of chloroform applied

to the side of the thorax kills thems at once. On my arrival at Batavia on

the morning of January iS''' 1863, after a sea voyage of three months, I went

to the Hotel des Tndes where a room, communicating with the garden, was

placed at my disposal. Here I observed my first Malayan butterflj' flying

round a shrub and, naturally, I could not resist the temptation of catching it

with the aid of my hat; it was a Euploea Midamus L. 9. I squeezed the

thorax of the insect until it was motionless and, as I thought, dead and fixed

it with an ordinary pin, for want of better, to a piece of board which I found

on the spot, provisionally placing the board on the top of a wardrobe. This

happened about 1 1 o'clock in the morning, shortly after which I left my room

to take luncheon and upon returning afterwards I noticed the board still in the

same place but both butterfly and pin had disappeared. I concluded that a

native servant had thrown away the insect. The next morning, however, about

10 o'clock I observed a similar butcerfly flying round the same shrub; I caught

it and it proved to be the identical one, the pin still through its bod}-. With

pin and all the creature had managed to detach itself from the board, had

passed the night in this state and then flew about as if nothing had happened !

When twelve years later, during a temporary return to Europe, I gave a lecture
at a meeting of the Nederl. Entom. Vereeniging on some of my experiences
in connection with Malayan insects, a translation of which appeared in the
Entomologist for November 1877, I was afraid to communicate this fact. It
appeared to me so incredible that I feared I should not believed and that. In
consequence, the reliability of my other communications might be doubted. It
was not until later, when I came across similar instances of the tenacity of life
in EuPLOEAs in Moore's Lepidopicra of Ceylon, communicated by Dr. Thwaites,
that I ventured to publish my own experiences in this connection. It has also
happened to me subsequently that I received from the mountains a tin of
butterflies in papers which had been lying between large quantities of naphtaline
powder, certainly not less than five days ; while examining them a Euploea
suddenly, on the paper being opened, flew out and escaped at once through
the window. This insect certainly had also had its thorax squeezed several
days previously and yet it well managed to survive its imprisonment in the
paper in the naphtaline-saturated atmosphere.

On page 5 of the Trans. Entom. Soc. London igo^ Mr. John C. W. Kershaw
states that while in China he observed a specimen of Epeh<a maculata which
had captured a Euploea Amymone Gdt. and had attackned the under side of
the abdomen and a part of the thorax, but upon releasing the butterfly from
its toils and placing it upon his hand it suddenly flew off and settled on a
high tree.

Trimen observed analogous cases with reference to specimens of Danais
and Acraea which upon being released from the pin, on which his native hunters
had stuck them, flew away as if nothing unusual had occurred.

In the case of two species I have repeatedly observed a yellowish-white &g^
deposited on the under surface of a leaf. I noticed the 9 placing herself near
the margin on the upper side of the leaf and thereupon curve the abdomen
round the edge and press it against the underside.

The larvae are provided with several pairs of appendages, whose number,
dimensions, and mode of attachment, vary in the different species. In the majority
of species, whose larvae are known, these processes in the full-grown larva are
long and four in number but in that of E. Mazares Moore they are short,
straight, and only three in number, thus agreeing with the processes in the
genus Danais. This rather points to the larvae of Euploea originating from
those of the genus Danais, especially since the young lar\-ae of the former
genus all have these processes short and straight and those of E. Mazares
Moore have, therefore, not kept pace in this evolutional change in that respect,
although the imago has assumed the new form, a fact which, moreover, occurs

in other cases. In the genus PAPiLro many instances of this will be observed,
giving rise to transition forms which distinctly indicate the course followed in
this development. According to Wood Mason the first pair of these processes
is movable ; I have observed this in only one species although I have reared
large numbers of these caterpillars, sothat this faculty appears to be in a state
of retrogression, being now found only in a few species or even in certain ,
individuals only. Moore in his Lepidopiera Indka states that the larvae
occur on Asclepiadaceae but I have found them on various plants.

The pupae are simply attached by their caudal extremity ; they are of stout
build, without projections, being rounded in outline, frequently with a bright
metallic lustre. The observation by Moore, concerning the pupae of Nym-
PHALiDAE, that the narrowness of the ventral side of the abdomen clearly indicates
that they are descended from a butterfly whose pupae were attached by means
of a girdle thread, applies equally to these pupae. The decrease in the faculty
of spinning, as this manifests itself in the pupae of the section Su.pa/si of
BoisDUVAL in comparison with those of his section siiccindi, is indeed, as I
have already stated, evidence of more advanced evolution in this respect, indi-
cating, therefore, that those forms in which it occurs must be of later development
than the others.

I. Climena Cram. (PI. XI, fig. \a, \h, ic, id).

Cramer, IF, p. 207, //. 389, £, F, (1782). . .
Butler, Proc. Zool. Soc. London, 1866,/. 2^2, fig. 2(3
Felder, Novara Lep., p. 335, (1864 — 1867). . .
Snellen, Tijdschr. v. Ent., 34, /. 231 (1891) .
Fruhstorfer (Seitz, Grosssc/im, d. Erde), p. 226, //.
86c (1910)

Papilio Climena.
Euploea Sepulchralis.

„ Zinckenii.

„ Sepulchralis.

„ Climena.

The Javanese specimens belong to the form Sepulchralis Butl. (Zinckenii
Felder).

Fruhstorfer distinguishes the specimens from E. Java as a separate form,
under the name of Ferissa, from those of W. Java, which he calls Sepul-
chralis. Without justification, however, for, as is shown by the accompanying
illustration (//. XI, Jig. la, \b), the increase in white occurs likewise in some
specimens from C. Java; it is, in fact, nothing but a manifestation, asserting
itself earlier or later in the various individuals, of the fading process of colour-
evolution. How uneven the course of this process is may be observed from
the fact that whereas all specimens from Bawean known to mc do not exhibit

this increase of white, my specimes from Flores, on the other hand, resemble
those of Java which do show it.

W. J. Batavia (3 — 14); Buitenzorg (365); Depok (95).

C. J. Touban (2) on the north coast; Magelang (500); prov. Madioun ;
prov. Tegal (Lucassen).

E. J. Kediri (64); Banyoupoutih (45) in the prov. Probolingo ; Semarou
mountains (800); Ardjouno mountains [Hekmcijo).

Snellen has noted with respect to this species :

" The Javanese specimens differ from the type from the Mokiccas only in
being smaller (Expanse of wings 55 — 62 mm. against 75 — 78 mm.). In some
specimens white occurs in addition on the hind wings ; Macleari Butler from
Christmas Island and Eurypon Hew. from the K6y Islands are probably also
forms of Cllmena Cram " .

I am unable to confirm Fruiistorfer's statement that this butterfl}' is
common at Batavia ; I never found it very common there, on the contrary, it
appeared to me much more common at Touban and at Kediri than in W. Java.

2. Malayica Butl. (PI. XI, fig. 2a, ib).

Moore, Cat. Lcp. E. J. C, I, /. 132 (1857) . . Euploea Ochsenheimeri.
Butler, y'otirii. Linn Soc. London, 14, /. 297 (1878). „ Malayica.

Snellen, Midden Sumatra, Lep., /. 1 2, pi. 2, Jig.

12 (?, (1882) „ Ochsenheimeri.

Distant, Rhop. Mai., p. 22, tab. 2, fig. 7, o"

(1882— 1886) „ Malayica.

Fruhstorfer (Seitz, Grossschm. d. Erde), p. 231,

//. 8o<^, (19 10)

Snellen notes the following:

" It is not the species to which Lucas applies the name Ochsenheimeri" .

For this reason Moore's name, Ochsenheimeri, although being the oldest
and his description agreeing well with the Javanese specimens, cannot be
accepted. Fruhstorfer, assigns to the latter the local name Hypanis,

W. J. Gedeh mountains (1000); Tjampea (160).
C. J.?
E. J. ?

I received the larva from Tjampea. Its back is velvety-black, except at the
first thoracic segment which, like the sides andundersurtace is of a dark orange

colour. On the black of each of the bod)- rings occurs a wide milky-white
transverse band attenuated at either end; this band on the second thoracic
segment is narrower. The head, the thoracic legs, and lower part of the pro-
legs are black. A pair of long black appendages project on the back from the
second and third thoracic, and from the second and penultimate abdominal
segments, the first pair being the longest. I noticed the caterpillar moved the
two foremost of the processes independently, but not to such an extent as
to bend them.

It feeds voraciously on the leaves of a species of Apocynaceae, a wild climbing
plant, evidently related to the food plant of E. Rafflesi Moore. It pupated on
the i^' September, the colour of the chrysaUs being a mixture of brown and black
with a golden and silver lustre, while the imago emerged on the lo* September.

3. Deheeri Doherty.

DoHERTY, Journ. Asiat. Soc. Bengal, 60, /. 163 (1891) . Euploea Deheeri.

Pagenstecher, ^d'/^r(5. y\^a^.f. F<';-. AW?^/-/&. 51,/. 188 (1898).

Fruhstorfer, (Seiiz, Grossschn. d. Erde), /. 231 (19 10). „ „

Fruhstorfek applies the local name Lamos to the Javanese specimens.
The species, discovered originally in Sumbawa, is recorded by Fi?.UHsroRFER
from E. Java, the specimens captured by him there, and which are in his collection,
being the only ones known from that island. They differ but slightly from the
specimens taken in Lombok. He has found it impossible to have an illustration
made of it.

Snellen notes here :

"This species, with an expanse of wings of 62 — 66 mm., has about the
form of wing of E. Malayica Bud. but is much smaller, the median area of
the wing, especially on the under side, is distinctly darker while the light dots,
a transverse row of which in front of the posterior margin of the hind-wings is
twice suddenly interrupted, arepurplish-white in colour. In the larger Malayica
the same transverse row of dots is curved round above, then straight, while the
colour is rather somewhat yellowish-white. The fore-wings, moreover, are
sometimes a little darker towards the external margin, but not in the median
area. In the 9 the inner margin of the fore-wings is quite straight but it does not
differ from the c? either in colour or marking. On the under side of the hind-
wings may be observed behind the dark median area an ill-defined, curved,
lighter transverse band. According to Dr. Pagenstecher's notes the species
appears to vary somewhat. It belongs to the section Adigama Moore, together

with Malayica Butl. It is doubtful whether the 9 described by de NiciSville
(Journ. Asiat. Soc. Bengal, 66, p. 674, 1897) belongs to the present species ".

4. Crameri Luc. (PI. XI, fig. 3a, ^6).

Lucas, Revue et Mag. Zoo!., 1853, /. 318

Felder, Wicu. Entom. Monatschr., IV, /. 398 (i860) .
Distant, R/io/^. Mai., /. 23, tab. 2,X^r. 4, c? (1882— 1886)
Moore, Proc. Zoo/. Soc. London, 1883,/. 267,//. 2<^, Jig. 5

„ Lcp. Ltd., I, /. 76, //. \c), fig. I, \a — d (1890)
Fruhstorfer, Berl. Ent. Zeitschr., 43, /. 187 (1898) .

„ , (Seitz, Grossschm. d. Erde), p. 229, //.

%2b, 86d (1910)

Euploea Crameri.
Bremeri.

Tronga

Crameri.

Euploea

Snellen has noted the following in connection with the present species :

" The typical form of E. Crameri Luc. does not occur in the Philippine
Islands but in Borneo, from which island Moore described it. It differs from
the Javanese form in being larger (80 — 82 mm. against 70 — 77 mm.) and in
being of a somewhat darker colour. The white spots on the fore-wings, more-
over, are almost limited to the apex, being also larger ".

These notes do not agree with the description by Fruhstorfer quoted snpra,
who appears to know Javanese specimens only from the Tengger mountains in
East Java which he designates by the name Tenggerensis.

W. J. Soukapoura, in the prov. Prayangan ; vicinity of Tjiletou or Sand
Bay on the south coast.

C. J. Babakan, prov. Banjoemas, on the sea coast, {^acodson).

E. J. Tengger mountains {Fruhstorfet).

5. MiDAMUs L. (PI. XI, fig. \a, \b, i,c, \d, 4c).

Linnaeus, Mus. LMd. Ulr., /. 251 (1764) . . .
SuLZER, Abgek. Gcscli. d. Lis., />. 144, //. 16, fig

4, 5, c? 9 (1776)

Cramer, II, /. 45, //. 127, C. D. (c?) (i779) • •
III,/. 132, //. 266, C. {d) (1782). . .
Fabricius, Ent. Syst. Ill, \, p. 40 9 (1793)- • •
HoRSFiELD, Cat. I^p. E. I. C.,pl. I, fig 10, ioa(i828)
Moore, Id. I, p. 133, //. ^, fig. 10, \oa (1857). .
AuRiviLi.ius, Rec. Cn't., /. 61 (1882)

Papilio Midamus.

„ Mulciber.
„ Basilissa.
„ Claudia.
Euploea Midamus.

10

8

Distant, Rhop. Mai., /. 24, tab. 2, /ig, 8, 9 6 9

(1882 — 1886) Euploea Midamus.

Staudinger, ^.vtfA &-^;//<'/'/.,/. 51,//. 25,c3'9(i884 — 1888) „ „

Moore, Le/^. Ind., I, p. 100,//. 1%, fig. \, \a, 1^(1890). Trepsichrois Midamus.
Fruhstorfer, (Seitz, Grossschm. d. Erde), /. 249,

//. 85a (1910) Euploea Mulciber.

Fruhstorker separates the specimens from Java under the name of Basilissa.

Snellen has noted regarding this species :

" Although Linnaeus, in some of his later works, confused the present
species with another Euploea (Super ba Herbst), it appears to be beyond
doubt, {vide Aurivillius, Rec. Crit. 1. c), that in the older work concerning
the Mus. Ludw. Ulr. a single species was described for which the Swedish
Professor cites Mulciber Cram, as the typical figure. The Javanese specimens
clearly differ, at least as regards the c?, from those of the South Asiatic Continent,
Malacca, and Sumatra (Mulciber Distant). They are somewhat larger, the
blue of the fore-wings is more purplish, their light spots are smaller, and the
median area contains less white. Tho white marking of the 9, moreover, is
somewhat finer. Midamus is a widely distributed species of which the following
are probably local forms : Diocletia Hbn. from the Philippines, Verhuelli
Moore from Nias, Mindanaensis Semper from Mindanao, and Gelderi Sn.
(DoNGO Doherty), from Flores.

W. J. Batavia (3 — 14); Buitenzorg (275); vicinity of the Tjiletou or Sand
Bay and the Pelabouan Ratou or Wynkoops Bay on the south coast (150);
Depok (95); Patjet (11 14); Sindanglaya (1074); Salak mountains (780); prov.
Prayangan (1500).

C. J. Semarang and Touban on the north coast; Jokjokarta (113); prov.
Tegal; Magelang (500); Bodjonegoro (258); Nousa Kambangan (Jacobson) ;
prov. Madioun.

E. J. Sourabaya on the east coast, Banyoupoutih (45) in the prov. Pro-
bolingo ; prov. Madoura ; Banyouwangy.

One of the commonest butterflies of Java, occurring everj^where, in the
lowest-lying districts as well as in the higher mountains and likewise met with
in many other islands of the Malay Archipelago with but slight variation.

A striking difference in colour exists in the two sexes, viz., that whereas
the c? entirely exhibits the Euploea type, in the 9 the process of colour evolu-
evolution, especially as regards the hind-wings, has evidently largely followed
the direction as it occurs in man)' species of the genus Danais, such as

D. Agleoides Felder for instance, so that the 9 much more resembles the
species of Danais than the c?. I have already discussed these different directions
of the process in the Introduction. Supporters of the theory of mimicry, who
maintain that the d of this species emits such a repulsive odour, being conse-
quently unpalatable to birds and that on account of the protection resulting
therefrom other butterflies have mimicked them, ought to explain why the 9 of
the same species has not availed itself of this protection but has on the contrary
mimicked other butterflies, even though generally these also are stated to be
unpalatable. Moreover, the beautiful dark blue structural colour, occurring on
the upper side of the fore-wings in the d — and the shade of which in the
specimens from Java differs from that in the Bornean ones — recurs, although
in a much less pronounced fashion, in the 9 and rather points to a similarity
of the scales in both sexes, since it is not improbable that these structural colours
depend on the external character of the scales. These 9 vary, i". in respect
of the darkness of the ground colour; 2*^. as regards the greater or lesser
intensity of the blue sheen just referred to, which may even be entirely wanting ;
3°. in the greater or lesser distinctness of the white stripes of the hind-wings,
which frequently are interrupted here and there, sometimes narrower and at
other times again wider. All these differences, however, may occur indiffer-
ently in specimens of East or West Javanese origin, and by no means constitute
distinct races or forms, as Fruhstorfer assumes, being simply individual
variations governed by their special development in the process of colour
evolution in each case. The unpleasant smell attributed to this species has
also been already dealt with in the Introduction.

The oviposition in this species has been observed by me and I succeeded
in rearing the caterpillars from the egg ; they eat voraciously and their develop-
ment is very rapid. During the rainy season at least, when their food may
be presumed to be the most nourishing, the time between leaving the egg and
the appearance of the imago would probably, in free nature, not exeed a
month. The larvae are found especially on the Waringin (Urostigma Benja-
MiNEA Miq.) and other species of Urostigma but also on other plants. As
has been recorded from Ceylon by Moore in the case of the indigenous E. Asela,
the present species is also frequently found in Java on the Nerium Oleander L,
an ornamental plant much grown in that island. The coloration' of the young
larvae does not differ, as such, considerably from the full-grown ones but while
being practically of the same type in all a fairly large amount of individual variation
exists, especially as regards the markings which at one time may be white and
at others again yellow or orange, while the black pigment also appears to be
much more prominent in some specimens than in others; alle these differences

lO

evidently indicating the various stages in the process of colour evolution.
The colour pattern is, consequently, difficult to describe and it will be more
satisfactory, therefore, to refer to the illustrations. The head is invariably
black or reddish brown, peculiarly marked with white or yellow lines. From
each of the second and third thoracic and the second and penultimate
abdominal segments a pair of fairly long processes extend from the back, those
of the second abdominal segment being more curved forward than the others.
In not quite full-grown individuals these processes are entirely black but in the
adults they are more or less dark red with black extremities. The handsome
pupae have the appearance of burnished gold or silver with some brown streaks.
When the larva has just recently moulted the pupa is flesh-coloured on which,
however, the metallic lustre soon appears.

The figures of the larva and pupa given by Horsfield are very inaccu-
rate and those by Moore, being copied from Horsfield, are no better;
my own illustrations are much better, although, having been executed in Java
b)- persons without any knowledge of the anatomical structure of caterpillars,
little attention has been paid to the division of the segments and, consequently,
the attachment of the third pair of processes is, apparently, not very accurate.
As regards the chrysalis, my illustration is of value only in connection with
the form since the metallic lustre cannot be reproduced.

6. Mazares Moore. (PI. XI, fig. ^a, 5<5).

MooRE, CaL Le/>. yT/. ^. /. C, /, /. 1 27 (1857) . . . Euploea Mazares.
Felder, Novara, Le/>., p. 317,//. 40, /f^. 5, 6 (1864 — 1867). „ Ledereri.

Distant, R/wp. Mai, p. 26, tab. 2, fig. 10(1882—1886)
Snellen, Tijdschr. v. Ent., 34, /. 232(1891) . . . . „ Mazares.

Fruhstorfer, (Seitz, Grossschtn. d. Erdc), p. 252, />/. 83rt'

(1910) ,

Snellen notes the following concerning this species :

" It is possible that this species, so well described by Moore, is the Papilio
TuLLiOLUs of Fabricius {Eni. SysL, III, i, p. 41, 173), but the description, trans-
lated by Godart in Encycl. Meth., IX, p. 181, 18 19, is (]uite insufficient.
E. HoPFEKi Felder, from the K6 Islands is probably a form of Mazares".

W. J. Batavia (3 — 14); D^pok (95) ; Tjampea (160); neighbourhood of the
Pelabouan Ratou or Wijnkoops Bay and of the Tjiletou or Sand Bay on the
south coast (+ 150).

1 1

C. J. Prov. Madioun ; Touban (4) on the north coast.

E. J. Banyoupoutih (45) in the prov. ProboHngo ; Willis mountains in the
prov. Kediri.

This butterfly appears to be common everywhere in the lower lying districts
but less so in the mountains. The shade of the brown of the upper side is
darker in some specimens than in others. Individual differences occur in the
white spots on the upper side of the fore-wings as regards size and number as
well as with respect to the development of some structural blue on these.

At Batavia the larva is found on a climbing plant known locally as Serout
OYOX (Cephalotrophis javanica B1.). Its general colour is brown or dark violet,
sometimes a little lighter and reddish, with many transverse faint white and
numerous wider black lines ; along the ventral margin occurs a yellowish-white
stripe. The ventral side is dark in colour, the legs are black ; the head is
shining black or brown marked with white lines. The very young larva is
evenly shining, partly yellow-brown, partly blackish grey, with a red head.
The most striking feature of the larva consists in its dorsal processes which
resemble those occurring in the species of Danais much more than those noted
by me in any other species of Euploea, being, as is the case with the former,
straight and short, not curved. Only three pairs are present, on the second
and third thoracic and the penultimate, but not on the second, abdominal
segment. Of these three pairs, agreeing in colour with the back, the foremost
is longest and the second shortest; in the very young larva the second pair
has not yet made is appearance.

Before pupating the larvae turn a greenish grey ; the pupae are at first
yellow but on the second day they begin to resemble burnished gold or silver.
In the case of two larvae which pupated on 1 1 "> December the imagines
emerged on the 1 9"^ of the same month, while one which commenced this process
on 4 February and another on 22 March, completed their metamorphosis on
12 February and 29 March respectively.

7. Eleusina Cram. (PI. XI, fig. 6).

Cramer, ///, /. 132, //. 266 D (1782) Papilio Eleusina.

HiiBNER, Satnml. ExoL Sckm., II, //. 3, Jig. 3, 4 9

(1806 — 1827) „

Godart, Enc. Meth., IX, p. 177 (18 19— 1823) .... Danais

Fruhsxorfer, (Seitz, Grossschvi. d. Erde), /. 262 (19 10) . Euploea „

Snellen has the following notes about the present species:

" Godart not only changes the name of this species without cause but he

I 2

confuses it with E. Mazares Moore, for he states that the purplish-white longi-
tudinal stripe above the inner margin of the fore-wings is frequently wanting in
the c? at least, which is incorrect. Cramer's figure is not good but recognizable ".

W. J. Tegal {Lticassai).

C. J. Touban (4) on the north coast.

E. J. Mount Ardjouno [Hekmeyer); Malang (443) in the prov. Pasourouan.

According to Fruhstorker this butterfly is common in Java, but I have
only rarely found it.

8. Leucostictos Gmel. PI. XII, fig. 7^, 7*^, 7c, 7</, 7^).

Gmelin, Sysf. Nat., /, 5,/. 2289, No. 889 (1788— 1 791)
GoDART, E71C. Meth., IX, p. i-JT, No. 2 (18 19— 1823)
Lucas, Lep. Exot., pi. 45^ A'- i (1835) • ' • •

/^ev. Zoo/., 1853,/. 313

Dtstani, /^//o/'. Ma/., p. 26, /ad. 13,/^. 6, 7 (1882— 1886)
Moore, Proc. Zoo/. Sac. London, 1883, /. 301 . .
Fruhstorker, (Seitz, Grossschm. d. Erdc), S. 262 (19 10)

Papilio Leucostictos.
Danais Eunice.
Euploea „

Dehaanii

Vestigiata.

Salphinx Dehaanii.
Euploea Leucostictos.

Snellen has noted in this case :

" In addition to those cited supra, the following probably pertain to the
present species: Viola Butl. from Celebes, Assimilata Felder from the K6
Islands, Kadu Eschholtz from the Marianne and Philippine Archipelagoes,
Depuisei-i Oberthur from Sangir and the Talaut Islands, Nemertes Hbn. from the
Moluccas, Nepos Rober from Goram, Perdita Bud. from New Guinea, and
PoLYMELA Godm. and Salvia from the Solomon Islands".

W. J. Batavia (3 — 14); Depok (95); Tjampea (160); Buitenzorg (275);
mount Karang in the prov. Bantam; Salak mountains (780); mountains of the
prov. Prayangan ; vicinity of Pelabouan Ratou or Wijnkoops Bay and of the
Tjiletou or Sand Bay on the south coast (+ 150).

C. J. Magelang (500); Jokjokarta (113); prov. Madioun.

E. J. Sourabaya on the east coast; prov. Besouki ; Semarou mountains
(740); Tengger mountains (1200) {Jacobson).

This butterfly, of which Fruhstorfer distinguishes the forms Debarbata,
BiocuLATA, Radiata, and Abrupta, is very common ; less so, however, in the
mountains proper than in the somewhat lower lying country. Mr. Ottolander,
however, asserts having observed the species in December 1885 migrating at
an altitude of 1000 Meters in the province of Besouki in East Java.

13'

Individual differences occur in the present species ; the colour of the upper
side, even of the hind-wings, being darker in some specimens than in others;
in the size and number of the white spots on the upper side of the fore-wings
there is also some difference. The structural blue in that region is not equally
developed in all c?, while in the 9 only traces of it, at the most, are present
and frequently it is entirely wanting. In spite of this I have failed to establish
local differences between the numerous specimens I have been able to compare.

The caterpillar lives on Scrouf (Streblus asper Lour), Waringin (Ficus
BENjAMiNEA L.), Ampclos kcbo (Ficus AMPELos Burm.), Gcgareman (Ficus
HETEROPHYLLA L.), Roiikem besar (Flacourtia Rukam Z. and M.). It is black,
the head shining. On the back occur about ten white transverse streaks, the
outermost single, but the others mostly double or even treble; they are so
wide sometimes that nearly the whole back becomes white. At times they are
continued at the sides as single vertical white lines which do not always coincide
with the dorsal streaks or may be entirely wanting. A blood-red spot occurs
at the sides of the third thoracic and the six foremost abdominal segments.
Four pairs of black dorsal processes, about equal in length, are found on the
second and third thoracic and the second and penultimate abdominal segments;
these processes are mostly curved forward a little.

The pupa is yellow with brown streaks at first but on the second day it
becomes like burnished silver or copper with grey and brown streaks radiating
from the caudal extremity. From a pupa of December i^' the perfect insect
developed on December g''' while one of April 6 and one of April 1 4 emerged
on the 14"^ and 24I'' of April respectively.

9. Phoebus Butl. (PI. XII, (fig. 8).

Butler, Proc. Zoo/. Soc. London, 1866, /. 270
Felder, Novara, Lcp., />. 3159 (1864 — 1867). . .
Distant, R/iop. Mai., /. 24, tab. 2, fig. 69 (1882 — 1 886)
Moore, Lep. Ind., /, /. 109, //. 38, fig. i, \a (1890)
Fruhstorfer, (Seitz, Grossschm. d. Erde), p. 259, //.
8o<^ (1910) . . . .

Euploea Phoebus.
„ Castelnaui.

„ Phoebus.

„ Corus.

Fruhstorfer indicates the Java specimens under the name of Pavettae.
Snellen notes the following in connection with the present species :
" Phoebus is probably only a local form of E. Phoenareta Schaller,
Naturf. 21, p. 177, //. 5, fig. I, 2 d" (Midamus Cram. Ill, /. 131, //. 266
A, B) from the Molucciis but I have seen no transition forms".

14

W. J. In the southern part of the prov. Prayangan, in the division of
Soukapoura and in the neighbourhood of the Tjiletou or Sand Bay on the
south coast (+ 1 50).

C.J. ?

E. J. Semarou mountains (740).

As I have never found this butterfly in the northern half of Java, nor
have I ever received it from there, while, on the other hand, I obtained several
from the southern half, I am inclined to think that it is restricted to that
part of the island, including East Java in its range. Fruhstorfer also found
it on the south coast of West Java.

10. Radamanthus F. (PI. XII, fig. 9(7, 9/^).

Fabricius, Ent. Syst., Ill, i, /. 42 d" (1793). . . Papilio Radamanthus.

„ I, /. 40 9 ,. • • ., Diocletianus.

Godart, Enc. Meth., />. 180, lYo. 13 (18 19 — 1823). Danais Alcidice.

HuBNER, Saniml. Exot. Schm., IF, pi. Sc? (1806 — 1822). Trepsichrois Thoosa.
Distant, Rhop. Mai, p. 28, tab. 4, jig. 4, 5, (5 9

(1882 — 1886) Euploea Diocletianus.

Staudinger, Exot. Schm., /. 53, />/. 26 c? (i 884 — 1 888). „ Rhadamanthus.

Snellen, Tijdschr. v. Ent., 42, /. loi (1899) . . „ „

Fruhstorfer, (Seitz, Grosssc/im. d. Eyde), p. 271,

//. 8i<: (1910) „ Diocletianus.

Fruhstorfer distinguishes the Java specimens under the local name of
Alcidice.

Snellen has noted in reference to this species :

"The Javanese specimens belong to the form Alcidice Godart or Thoosa Hbn.,
with only three or four white streaks, shorter than those in the type, near the
inner margin of the hind-wings".

W. J. Depok (95) ; Tjampea (160) ; Sindanglaya (1074) ; Tjibougel (+ 1000) ;
vicinity of Pelabouan Ratou or Wijnkoops Bay (+ 1 50).

C. J. ?

E. J. ?

I have never observed this butterfly at Batavia but it is common at Depok
and many other places. The 6 appear to be commoner than the 9. although
the latter are not rare. The fading to white in this species takes place in a
peculiar manner which likewise appears to occur in Papti.io Caunus Westvv.

15

and in a 9 form of Euripus Hallitherses Dbd., for which reason these are
supposed to mimick the present species. This Papilio does not, however,
appear to occur in Java; the only record of a specimen taken in that island
is of ancient date and, like many such old records, not entirely reliable. In
the present species the structural blue is strongly developed in the c? but litde or
not at all in the 9, while the brown in the 9 is much lighter than the dark .shade
of that colour in the d.

The larva is unknown to me. The pupa was once found by me at Tjampea ;
it very much resembled that of E. Leucostictos Gmel.

II. Eyndhovii Felder. (PI. XII, fig. lo).

Felder, Novara Lep., p. 338 (1864 — 1867) .... Euploea Eyndhovii.
Snellen, Tijdschr. v. Ent., 35, pi. i (1892) .... „ Alcathoe.

Fruhstorfer, (Seitz, Grossschm. d. Erdc, /. 237 (19 10). „ „

Fruhstorfer distinguishes the forms Alcathoe and Geyeri from Java.

Snellen has noted in this connection :

" Felder's description is good, but of the blue sheen on the fore-wings no
trace is to be seen in the specimens in our collection. The description in the
Novara has been drawn up from old specimens from the collections of Eynd-
hoven and van der Capellen ".

W. J. Batavia (3 — 14); Buitenzorg (275); Tjibodas (14 10) ; Patjet (i 114) ;
Tjibougel (+ 1000).

C. J. ?

E. J. Banyouwangy, Semarou mountains (740).

12. HuEBNERi Moore. (PI. XII, fig. 11).

Hubner, Samml. Exot. Sckm., II,pL 9, fig. 1,2(1820—1821). Salpinx Eleusina.

Moore, Cat. Lep., M. E. I. C, I., p. 128 (1857) . . Euploea Hubneri.
Butler, Proc. Zool. Soc London, 1866, /. 273 . . . . „ Janus.

Felder, Novara, Lep., /. 315 (1864 — 1867) „ Moorei.

Moore, Proc. Zool. Soc. London, 1883, />. 280 .... Chastia Haworthi.

Fruhstorfer, (Seitz, Grossschm. d. Erde), p. 237 (1910). Euploea

Snellen has the following notes about the species:

"Moore considers {Proc. Zool. Soc. London, 1883) this species co-specific
with Euploea Haworthi Lucas {Revue Zool., 1853, /. 31 7), but the description

II

i6

is cursory and doubtful ; it mentions no dull longitudinal streak on the fore-
wings, gives the expanse too great — 80 mm. instead of 65 — 70 mm. —
and states that the spots of the inner row along the inner margin of the hind-
wings are oblong. This reminds of a g of E. Menetriesii Felder".

W. J. Batavia (3 — 14); Depok (95); prov. Prayangan (1500).

C. J. Magelang (500) ; Touban on the north coast.

E. J. Malang {443); Tengger mountains (500 — 700) (Fruhstorfer).

This butterfly is not, as Fruhstorfer thinks, rare at Batavia, but is very
common there.

13. Menetriesii Felder. (PI. XII, fig. 12).

Felder, Wicfi, Ent. Monatschr., IV, /. 398 (i860). . Euploea Menetriesii.
Butler, Trans. Linn. Soc. London, 2^^ Scr. I, p. 535,

//. t(), fig. 9 (1879) .. Pinwilli.

Distant, R/iop. Mal.,p. 35, tab. I, fig. 9, 10(1882— 1886). „ „

Snellen, Tijdschi: v. Ent., 33, /. 284 (1890) ... „ Alcathoe.
Fruhstorfer, (Seitz, Grossschm. d. Erdc), p. 238, //.

86a (19 10) „ Deione.

Fruhstorfer separates the Java specimens as the local form Wallengreni.

Snellen notes here the following :

" The Javanese specimens of this species differ from the type from Malacca
and Sumatra in being somewhat smaller and having darker fore-wings, the
former appearing also to be devoid of the white marking. Alcathoe Sn. is
not Godart's species " .

W. J. Depok (95); Buitenzorg (265); Mountains in the prov. Prayangan.

C. J. Prov. Rembang.

E. J. Prov. Kediri ; Prov. Banyouwangy.

14. Rafflesi Moore. (PI. XII, fig. 13a, \2,b, 13^).

Moore, Proc. Zool. Soc. London, 1883, A 3H- • • • Isamia Rafflesi.
Fruhstorfer, (Seitz, Grossschm. d. Erdc), p. 268, //. 84^

(19 10) Euploea Aegyptus.

Fruhstorfer definitely calls the Java form " Rafflesi ".

Snellen notes with references to this species :

"Perhaps Lucas when describing E. Ochsenheimeri {Revue Zoo/., 1853,

17

/. 315) had the present species before him but his description, like most of the
older descriptions of Euploeas without figures, is practically useless, whereas
Moore's may be considered good".

W. J. Batavia (3 — 14); Tjampea (160); Buitenzorg (275); Gadok (+ 550);
vicinity of the Pelabouan Ratou or Wijnkoops Bay (+ 150).

C. J. Touban (6) on the north coast; prov. Madioun.

E. J. Without precise indication of habitat; Malang (500 — 600) {Fnihstor/er).

The species is very common in West Java. In the 6 the colour of the
upper side of the fore-wings is much darker than in the 9 but does not exhibit
any iridescent blue. The white of the upper side increases more strongly in
the 9 than in the c?, in some 9 this increase is particularly striking.

The ^^^ is yellowish white and strongly oval ; it is attached with the apex to
the under side of a young leaf and therefore hangs perpendicularly. The larva
occurs repeatedly on Kcmbang pita (Strophantus dichrotomus B1.) a cultivated
ornamental plant. It is smooth and shining, orange in colour, sometimes
approaching green. The head, the posterior extremity, the stigmata, the lower
extremity of the legs, and the four pairs of processes, except their lower portion,
are black. These processes occur on the first and third thoracic and the second
and penultimate abdominal segments ; the anterior pair is the longest, the third
shortest; they are all somewhat curled in a forward direction. These processes
much resemble those ocurring in the flowers of the plant on which the larva
feeds. A splendid instance of mimicry for a superficial observer; for in this
manner the larva has adapted itself to its environment ! Unfortunately, however,
other EuPLOEA larvae, living on different plants, have similar processes. The
caterpillars are much pestered by Ichneumonidae and Tachiinae. The pupae
are yellow at first but on the second day they begin to resemble burnished
silver or copper, with some ochreous-yellow streaks radiating from the
caudal extremity and provided anteriorly with some black dots. A pupa of
September 8 changed into the imago on September 1 5 and one of December 1 5
emerged on December 23. Once I found a pupa on ^//^/^/as (Ficus Ampelas
Burm.).

15. Gamelia Hiibn, (PI. XIII, fig. 14a, li^b).

HiiBNER, Saturn/. Exot. Schm., II, pi. 10 (1806 — 1827) . Salpinx Gamelia.

Chapman, E71L Monthly Mag., IX, p. 263 9 (1873). . . Euploea Lorraini.
ZiNCKEN, Nova Acta Ac. Nat. Car., XV, p. 186, //. 16,

fig. 18, 19 (188 1) „ Faberi.

Moore, Proc. Zoo/. Soc. London, 1883, /. 3 1 7 . . . . Anodara Gamelia.
Fruhstorfer, (Seitz, Grosssc/ivi. d. Erde), /. 268 (1910) . Euploea „

The following has been noted respecting the species by Snellen:

" E. LoRRAiNi Chapman, is simply a cursory description of the 9. The
illustrations by Hubner and Zincken ars good. Gamelia is hitherto known
from Java only".

W. J. Mount Salak, Megamendoung, Gedeh and other mountains in the
prov. Prayangan.

C. J. ?

E. J. ?

A mountain butterfly common above 1000 meters altitude in the mountains
of West Java.

16. Gloriosa Bud. (PI. XIII, fig. x^a, is^)-

Butler, Pjvc. Zoo/. Soc. London, 1866, /. 293,

/>/. 29, jig. 49 Euploea Gloriosa.

Snellen van Vollenhoven, Tijdsc/ir. v. Enl., 9,

//. 209, //. 10, jig. I d" (1806) „ Superba.

Felder, Novara, Lep., p. 327, //. 41, jig. 5 <3

(1864 — 1867) „ Schlegelii.

Moore, Proc. Zoo/. Soc. London, 1883, /. 321 . . Sticteuploea Lacordairei.
Fruhstorfer, (Seitz, Grossschn. d. Erde), p. 246

(1910) Euploea „

Fruhstorfer is of opinion that among the Java specimens the forms
Defigurata and Magniplaga may be distinguished.

Snellen notes here as follows :

" The first description and illustration of this species are, probably, those of
Snellen van Vollenhoven, but the name Superba was already applied to a
previously described species of Euploea, /. e. E. Superba Herbst from China.
The Javanese specimens do not agree with the type from Celebes, to which
Schlegelii Felder also pertains, but constitute the smaller and less marked form
Lacordairei Moore. Perhaps the species had already previously been described
as Danais Dufresnei from the Philippines by Godart {Enc. Mdth., IX, p. 815,
1 8 19 — 1823) but this description is very superficial and might equally apply
to other species. Gloriosa d* is easily distinguished amongst the Javanese
EuPLOEAs by the two faintly shining, longitudinal streaks in cell i b of the fore-

19

wings, the dark blue lustre of their second half, and the purplish blue spots
along the posterior margin ".

W. J. Neighbourhood of the Pelabouan Ratou or Wijnkoops Bay (+ 150).

C. J. Touban (4) on the north coast.

E. J. Berbek (i 3) in the prov. Kediri ; Tengger mountains (2000) {FitiJislorfei).

Genus HESTIA Hiibn., Dbd., and Hew.

The Hestias are likewise slowly hovering butterflies but they do not fly
as low as most Euploeas; they frequent wooded country but only where the
sunlight is able to penetrate. According to Moore one of the species from
British India is migratory ; this is not the case with the Javanese ones, whereof
the only species which is not actually rare does not, moreover, occur in great
numbers.

I. Belia Westw.

According to Fruhstorfer butterflies of the genus Hestia are called by
the Javanese koupou kertas, i. e. paper butterflies. He must mean that these
butterflies are so designated in Java, for koupou kerias, or rather koupou
ko7tpou karias, is not Javanese but Malay; I have never heard the term.

According to the observation of Captain H. Macpherson, communicated
by Moore, with reference to the Indian species called Hestia Malabarica by
him, the egg is " deposited singly on the under surface of a leaf and is white,
oval, about Vie inch long by V32 broad, attached to the leaf by one of the
small ends and marked with about twenty-two longitudinal rows of hexagonal
indentations". They are said to hatch after 6 or 7 days. The young larva
is stated to remain mostly on the under side of the leaf, eating small holes
into it and only subsequently proceeding to eat the leaf from the margin inwards.
They are said to be full-grown in from 20 to 25 days. According to one of
the figures, given by Moore of this caterpillar, it is evidently closely allied to
that of H. Lynceus Drury.

Westwood, Cab. Or. Entom., p. 1^, pi. 37>

jig. 2 (1848) Hestia Belia.

Snellen van Vollenhoven, Tijdschr. v. Enl.,

Ill, p. 43, //. 3 (i860) Idea Hypermnestra van Jasonia.

20

Butler, Trans. Linn. Soc. London, 2 Ser., I, /. 536,

//. 69, fio. 6 (1879) Hestia Linteata.

Distant, Rhop. Mai., /. 7, tab. \,Jig. i (1882— 1886).
Fruhstorfer, (Seh z, Grossschm. d. Erdc), /. 221,

//. 74<r (1910) „ Hypermnestra.

W. J. Tjampea (160); Gounoung Pantjar, a mountain in the division of
Buitenzorg; vicinity of the Tjiletou or Sand Bay on the south coast; vicinity of
the Pelabouan Ratou or Wijnkoops Bay on the south coast (+ 150) {Frulistorfer).

C.J. ?

E. J. ?

The species appears restricted to West Java, but is not so rare as Fruh-
storfer thinks ; it exhibits many individual differences. Fruhstorfer indicates
particularly the Javanese specimens under the name of Belia.

2. Lynceus Drur)'. (PI. XIII, fig. i6a, \tb\

Drury, ///. Exot. Ent., pi. 7, fig. i (1773) .
Stole, Suppl. Cram., p. 180, //. 42, fig. i (1790)
GoDART, Ene. Meth., IX, p. \<^^, No. 2 (18 19 — 1823)
Doubled AY, Gen. Ditirji. Lep., pi. 13, fig. i (1847)
Moore, Proc. Zool. Soc. London, 1883, /. 218

» J)

,. ,,219
Distant, Rhop. Mai., p. 6, 405, lab. i, fig.

(1882— 1886)

Fruhstorfer, (Setpz, Grossschm. d. Erde),p. 220(1910)

»j »i 11

Papilio Lynceus.

„ Idea.
Idea Lyncea.
Hestia Idea.
Stolli.
„ Reinwardti.

Logani.
„ Donovani.
„ Druryi.

„ Lynceus.

StolU.
„ Logani Mevaria.

W. J. Buitenzorg (265); Tjampea (160); Manondjaya; vicinity of Pela-
bouan Ratou or Wijnkoops Bay (+ 1 50).
C.J. ?
E. J. Banyoupoutih (45) in the prov. Probolingo.

Fruhstorfer distinguishes two species : H. Lynceus Drury, Stolli Moore,
and H. Logani Moore, Mevaria, Fruhst., the former of v^rhich is said to be
rare in Java and the latter extremely common. Snellen considers the two
simply synonymous. The apparent distinction which Fruhstorfer thinks he is

21

able to point out between the two is not to be traced upon investigation ; there
appear to be simply darker or lighter individuals. The alleged difference in
form of the genitalia I am at present unable to admit either. Herr Fruh-
STORFER had the kindness to hand me photographic illustrations of the genitalia
of his two species Stolli and Logan i, that of the former, however, having
been prepared not from a Javanese specimen but from one from Sumatra.
Dr. Henri de Graaf, a specialist in this kind of researches, has investigated,
microscopically 25 specimens from my collection, both light and dark coloured,
and he did not find a single one amongst them whose genitalia agreed with
the figure given of those of Stolli, on the contrary they all agreed with those
given of Logani, and although considerable individual differences were observed
in this respect it was only as regards the form of the valvae and valvulae and
not to such an extent in connection with the structure of the principal organs
that the existence of more than one species could be inferred from it. For
this reason I am of opinion that for the present, at least as regards Java, all
these forms must be referred to a single species. I have nowhere met this
species so abundantly as Fruhstorfer has recorded it to be.

The larva has been observed at Tjampea on a plant which has not been
identified (probably a Gvmnema). It is of the Euploea type, entirely jet-black
with milky-white transverse dorsal bands on the three thoracic segments and
on the third, fourth, fifth, and penultimate abdominal segments. At the sides
of the second and seventh abdominal segments a handsome red spot occurs,
which sometimes is diffused almost into a transverse band.

Four pairs of long black processes are found on the second and third
thoracic and the second and penultimate abdominal segments, the two fore-
most pairs curved forward, the two hindmost somewhat curved backward. The
very handsome pupa, of a golden hue marbled with black, brown, and white,
is not of the Euploea type. From a pupa of February 27"^ the imago appeared
on March 12"'. The illustration of the larva has been made from a not quite
full-grown individual and is somewhat stiff. The larva grows larger.

3. Leuconoe Erichs.

Erichson, Nova Ada Acad. Nat. Cur., XVI, p. 283

(1834) ■ Idea Leuconoe.

DoLTBL. AND Hew., Gen. D. L., pi. 13, fig. 2 (1847). Hestia
Distant, Rhop. Mal.,p. 406, tab. 2,9^ fig- 3 (1882 — 1886).

Snellen, Notes Leyden Mm., 17, />. 119 (1895) • • - van Natunensis.

Tydschr. v. Ent. XXXIX, /. 43 (1896). . „ Leuconoe.

22

Fruhstorfer, Ent. N'ac/ir., 22, /. 65 (1896) .... Hestia Leuconoe.

„ Ber/. Ent. Zeiischr., 42, /. 321 (1897) . . „ „

Snellen, Tijdschr. v. Ent., XLV, /. 75 (1902). ... „ „

Fruhstorfer, Iris, XVI, p. 60 (1903) „ „

„ (Seitz, Grossschiii. d. Erdc), /. 222, //.
74(7 (1910) . . , >, .,

The only specimen, said to have been found in the province of Japara, in
the north of Java, is in the collection of Dr. Pagenstecher at Wiesbaden.
From the figure published by Fruhstorfer, who distinguishes this form under
the name Javana, it pertains, according to Snellen, to the form described by
him as Natunensis.

Perhaps it has been found in the Karimon Djawa islands situate to the north
of the province of Japara and belonging to Java. A specimen, in my collection,
from the Kangean islands, lying east of Java, also belongs to the present form.

Genus IDEOPSIS Horsf. and Moore.

I. Gauka Horsf. (PI. XIII, fig. 17a, \']d, 17c, 17^/).

HoRSFiELD, Cat. Lep. M. E. I. C, pi. t, fig. i (1828). . Idea Gaura.
BoiSDUVAL, Spec. Gen., I, pi. 11, fig. 11 (1836) . . . . „ „

Horsfield and Moore, Cat. Lep. M. E. I. C. I.,/. 133 (1857). Ideopsis
Fruhstorfer, (Seitz, Grossschvi. d. Erdc), p. 216, //. \ba

(i9>o) „ „

W. J. Gounoung Pantjar, a mountain in the division Buitenzorg; in the
vicinity of the Pelabouan Ratou or Wijnkoops bay (+ 1 50) on the south coast.

C. J. Noesa Kambangan {jfacobso7i).

E. J. Semarou mountains (140); prov. Besouki; Blitar; Tengger mountains
(1200) {Jacobsoii).

This species flies amongst the underwood after the manner of the members
of the genus Danais but a little higher, about a man's height or more. The
only specimen obtained by me during the many years I have hunted in the
north of Java is the caterpillar taken on Gounoung Pantjar, at an altitude of
certainly not less than 300 metres ; in the south, especially in the Tengger
mountains in E. J., at an altitude of 1200 metres, it appears to be fairly common.

Lighter or darker specimens occur in both sexes without any apparent

23

connection as to habitat or time of appearance ; the black on the upper side
is not so much diffused in all $ as in the specimen figured. ' The larva taken
at the place just mentioned was found living on a climbing plant which has
been identified as Melodinus Laevigatus B1. It is intermediate between the
EuPLOEA and the Danais type, velvety black with a milky white transverse
band on each segment. It has only two pairs of processes, also black but red
at the base. The anterior pair, projecting from the second thoracic segment,
is considerably longer than the posterior pair which is situate on the penul-
timate segment. The head is shining and furnished with two sharp points.
The pupa is of the Danais type, whitish green with a black caudal hook and
several round black dots. A row of 8 such dots occurs at the boundary ot
the thorax and abdomen, the others being distributed in 6 pairs on the thorax.

Genus DANAIS Latr. ')

Fruhstorfer (Seitz, Grossschvi d. Erde, Danaidae, /. 193) states that
Danais (ANOSfA) Plexippus L., Archippus F., Erippus (Cram.), of American
origin is said to have spread even to Java. The same statement has been
made by various English and American authors ; the first to assert this appears
to be KiRBY. It is, however, probably based on a misconception, due, most
likely, to the fact that D. Genutia Cram, has been referred by some entomo-
logists to D. Flexippus L. The true D. Plexippus L., as has already been
mentioned in the introduction, has spread from America westwards over the
Pacific islands but from Australia only northwards apparently and subsequently
again eastwards. For specimens are known from new Guinea, the Moluccas,
the Talaut islands. North Celebes and Southern China, but I have never seen
any from one of the islands in the sea of Java anymore than from Java,
South Borneo, South Sumatra nor from South Celebes, although I have myself
during five years collected butterflies in that region, and such a large insect —
so conspicuous by its colouring — which, moreover, occurs in the districts
near the coasts, could not have escaped attention.

•) In the Leyden Museum is a specimen of Danais Isimare Cram, which is said to have
been forwarded from Java by the late Reinvvardt. Since no other specimen of this species, to
my knowledge, from Java exists and in view of the unreliable character of many of the old
labels in this museum, I do not propose, for the present, to include the species in the Java fauna.
Fruhstorfer, (Seitz, Grossschm. d. Erde) also mentions a form of D. Affinis F., called Kasirensis
by him, as doubtful for Java, a couple of specimens of which he states to have seen whose
origin, however, he doubts. I do not, therefore, propose to include this species either in the fauna
of Java for the present.

12

24

These insects fly amongst the underwood mostl)' close to the ground ; many
belong to the commonest Rhopalocera of Java. Considerable individual differ-
ence occurs in the size of the white and brown spots and streaks in the same
species. The larvae are furnished with processes similar to those occurring in
the genus Euploea but they are shorter and less numerous. The anterior pair
of these processes is movable, according to Wood Mason ; I have, in fact,
observed this in a couple of species ; if, however, we take into account the
great number of Danias larvae I have reared it must be assumed that this is
of rare occurrence, as has already been stated with respect to Euploeas. The
colouring of the pupae has already been dealt with in the introduction. According
to Moore some of the species, amongst these D. Genutia Cram., very common
in Java, and D. Chrysippus L. in British India, belong to the migratory butter-
flies, but this is not the case in Java.

I. Albata Zinck. (PI. XIII, fig. i8).

ZiNCKEN, Nova Acta Ac. A^at. Cur., X\\ p. i8i, //. i6,

fig. i6 (1831) Euploea Albata.

Moore, Proc. Zoo/. Soc. London, 1883, />. 248. . . . Mangalisa

pRUHsroRFER, (Seitz, Grosssckm. d. Erdc), p. 209 (1910). Danais „

\V. J. Megamendoung, Gedeh, Wajang mountains; (1000 — 2000).

C. J. Oungaran (700 — 1000) {Jacobso7i).

E. J. Tosari (1777); Tengger mountains (i 200) (^af(7i5j£'«) ; prov. Besouky.

A mountain species, very common everywhere in the upper mountains
above 1 000 metres altitude, where it may frequently be observed flying swiftly
close to the ground in the dark forest. Fruhstorfer found this butterfly on
Mount Gedeh as high up as 2600 metres and distinguishes the E. J. specimens
as GiLVA from those of W. J.

2. Melaneus Cram. (PI. XIll, fig. 19).

Cramer, /, /. 48, //. 30 D (1779) Papilio Melaneus.

ZiNCKEN, Nova Acta Ac. Nat. Cur., XV, p. 179 (1831). Euploea Melanea.

Felder, Novara, Lep., p. 349 (1867) Danais Larissa.

Distant, R/iop. MaL, p. 14 (1882 — 1886) , Melaneus.

Moore, Proc. Zool. Soc. London, 1883, /. 250 . . . . Caduga

„ Lep. Lnd., /, /. 60,//. \^,fig- 2, 2d:— <^ d" 9 (1890). „ „

Fruhstorfer, (Seitz, Grosssckm. d. Erde), p. 210, //. 78c

(19 10) Danais

25

The Javanese specimens, according to Snellen belong to the form Larissa
Felder. A still smaller difference in colour evolution originated the form
PsEUDOMELANEUS Moore. Under the latter designation Fruhstorfer specially
indicates the Java form.

W. J. Gedeh and other mountains in the prov. Prayangan (1400 — 1600).

C. J. ?

E.J. ?

A mountain species, not very common.

3. Aglea Cram. (PI. XIII, fig. 20a, 20b).

Cramer, IV, /. 173, //. 377 E (1782) Papilio Aglea.

Distant, R/iop. Rial., p. 14, 408, tab. i,fig. 6 (1880 — 1886). Danais Melaneus.

Moore, Proc. Zool. Soc. London, 1883, /. 251 . . . . Caduga Banksii.

„ Lep. Ind., I, p. 55, //. 12), fig- la — d (1890). . Parantica Aglea.

According to Moore the figure by Cramer represents the form from the
south of British India. The accompanying illustration has been prepared from
a Javanese specimen.

W. J. Meester Cornells (25) ; Depok (95) ; Tjampea (160) ; Buitenzorg (265) ;
Gedeh mountains (1082); vicinity of the Tjiletou or Wijnkoops Bay (+ 150).

C. J. Prov. Madioun.

E. J. Semarou montains (750).

I did not meet with this species in the vicinity of Batavia but at a slightly
higher altitude at Meester Cornelis.

The egg is fairly large, white, strongly ribbed longitudinally. The larva
was found on hanyouivatig (Marsdenia tinctoria R. Br.) at Tjampea. Moore
gives an asclepiad, Tvlophora carnosa, as its food plant in British India and
Calotropis in Ceylon ; it is of the Danais type and shows marbling of milkv
white spots with reticulation of faint lilac lines. On the last three segments,
however, the spots are yellow and in the full-grown larvae a subdorsal row of
yellow spots occurs in addition, except on the i^' and 2°*^ thoracic segments.
A yellow dot invariably occurs on either side of the i*' thoracic segment. The
back is at times of almost a chalky white beneath which a black ground is
scarcely discernable ; in young larvae, on the other hand, the back is less
white and the general colouring, therefore, darker. Two pairs of processes
are present, the anterior pair, on the 2°'^ thoracic segment, being much longer
than the posterior pair which is situate on the penultimate segment ; the former

26

are black at the extremity, white or pearly grey on the outer side and black
or dark lilac on the inner side. The head is black with milky-white spots,
while on each of the pro-legs and on the anal segment occurs a black dot.
The larvae not only nibble the edges of leaves but make holes in them, first
nibbling through the prominent veins on the under side and proceeding thence
to eat awa}'.

The pupa is similar in form to that of the related species, being likewise
green with some white spots on the back of the thorax which subsequently
become partly of a golden lustre. In addition there is a row of black dots at the
boundary of thorax and abdomen, each in turn surrounded by a white ring and
all arranged on a dull gre)- band which is mostly of a golden lustre. The figures
of larva and pupa by Moore do not quite agree with mine. My descriptions
have invariably been taken from the living animal.

4. A.SPASIA F. (PI. XIII, fig. 21a, 2 lb, 2 If).

Fabricius, Mani. Ins., //, /. 1 5 (1787) Papilio Aspasia.

ZiNCKEN, N'ova Ada Ac. Nat. Cur., XV, p. 184,//. 16,

fig- 17 (183 1) Euploea Philomela.

Butler, Proc. Zoo/. Sac. London, 1866, //. 4, //c^. 5 . . Danais Crocea.
Distant, Rhop. Mai., /. 13, lab. i, fig. 7(1882—1886). „ Aspasia.

Moore, Lcp. Ind., /., /. 53, //. 12, fig. \a, b (1890) . Bahora Crocea.

AuRivjLLius, Ent. Fidster, 1897, /. 144 Danais Aspasia.

Fruhstorfer, (Seitz, Grossschm. d. Erdc), /. 205, //.
76er (1910)

Fruhstorfer indicates the Javanese specimens under the name of Philomela.

Snellen noted with reference to this species that, indeed, it differs slightly
from D. Cleona Cram (IV, /. 173, //. 377 F 1782) from the Moluccas,
being somewhat smaller and somewhat longer-winged while the markings are
partly greyish white and the streak in the discoidal cell of the fore-wings is
never entirely interrupted at the middle, but that, nevertheless, it must probably
be looked upon as a form of that species without this being confirmed,
however, by sufficient transition forms.

W. J. Depok (95) ; Buitenzorg (265) ; Tjampea (160) ; Salak mountains (780) ;
Tjibougel (+ 1000); Mount Karang in the prov. Bantam.

C. J. Tegal {Lucasscti) ; Magelang (500); Semarang (25) {Jacobson);
Gounoung Sawou in the prov. Jokjokarta {Jacobson).

E. J. Prov. Kediri ; Tengger mountains (1200) {Jacobsoii).

27

I never encountered this butterfly in the vicinity of Batavia. At Depok
and elsewhere, for instance in the Tengger mountains, it is very common. The
larva feeds on a climber which I was told is called ke/atnbennati, probably a
species of Gymnema. It is entirely of the Danais type. The ground
colour is velvety black while the head is also black. The markings consist of
numerous white streaks and dots, and a subdorsal row of round spots, one on
each segment, those on the median segments being yellow while those on the
outer, especially the posterior ones, are orange. In addition an oblong orange
spot occurs on either side of the last segment. The larva is provided with
two pairs of processes, the anterior pair being fairly long but the posterior
short ; these processes are black except the median part of the anterior pair which
is whitish grey. The anterior processes can to some extent be moved voluntarily.
The pupa is also of the Danais type, light green with black caudal hook
having a couple of black dots in close proximity ; at the boundary between
thorax and abdomen is a row of lo black dots and parallel with it a similar
row of 8 dots on the abdomen ; all these dots are placed within milky-white
rings. The abdomen further has a pair of milky-white dots. A pupa of
30 November produced the imago on 8 December.

5. Agleoides Felder. (PI. XIII, fig. 22a, 22b, 22c).

Felder, Wien. Eut. Motiatschr., IV, p. 398 (i860). . Danais Agleoides.
Butler, Cat. Lep. Fabr., p. -j, p/. i, fig. 2 (1870). . „ Eryx.

Di^TA^r, Rhop. Mai., p. 1 5, /a,^. i, /}V. 5 (1882 — 1886). „ Agleoides.

Staudinger, Exof. Sc/im., p. 49, //. 23 (1884— 1888) . „ Eryx.

Moore, Lep. Ind., I, p. 58,//. 14, //>. la—b (i8go) . Parantica Agleoides.

Concerning this species Snellen noted that it differs from D. Similis
Clerck only by having the light longitudinal lines in the discoidal cell of the
fore-wings uninterupted in both sexes and by the c? possessing a sexual black
spot on the under side of the hind-wings which is not the case in the c? of
D. Similis Clerck.

W. J. Batavia (3—14); Buitenzorg (26s); Tjampea (160); Soukapoura.

C. J. ?

E. J. Jember (98), in the prov. Besouky; Banyoupoutih (45), in the prov.
Probolingo.

The larva which occurs on the same climber as the preceding species and
on mcngkoudou oidan (Lasianihus Firmus Miq.), is entirely of the Danais
type, resembling the larvae of the other species, with a dark ground colour

28

and numerous fairly large yellowish-white spots. The pupa is also of the
Danais type, of a handsome light green with black caudal hook ; a golden
band, containing a row of round black spots, occurs on the boundary between
thorax and abdomen. A few silvery spots in addition on the thorax some of
which contain a black dot. A pupa of 2 March gave the imago on i i March.

6. SiMiLis Clerck. (PI. XIV, fig. 23^', 23*5, 23^).

Clerck, Icones, Ins., I, />/. 16, fig. 3 (1759) Papilio Similis.

Linnaeus, Mus. Lud. U/>'., p. 299 pars (1764). ... „ „

Cramer, /, /. 92, />/. 59, F (1779) „ Aventina.

Butler, Ent. Monthly Mag., X, /. 164 (1874). . . . Danais Vulgaris.

Disfant, R/iop. Mai., p. 10, tab. \, fig. 8 (1882— 1886). Radena

AuRiviLLius, Recensio Critica, p. 99, No. Wja (1882). . Papilio Similis.
Moore, Proc. Zoo/. Soc. London, 1883, /. 223, //. 31,

fig. 4 , , . . . Radena PersimiHs.

Moore, Lcp. hid., I, p. 27, />/. 5, /ig. i, la (1890) . . „ Vulgaris.

Fruhstorfer, (Seitz, Grosssckvi. d. Erde), p. 211, />/.

783 (19 10) Danais Similis.

Snellen noted that Linnaeus and some subsequent authors confused several
allied species with Similis, which caused Butler to apply the name Vulgaris
to the present species. Aventina Cram, is a poor illustration while Distant's
figure of Vulgaris also leaves much to be desired.

W. J. Batavia (3 — 14); D^pok (95); Tjampea (160), vicinity of the Pela-
bouan Ratou or Wijnkoops Bay (+ 1 50).

C. T. Magelang (500).

E. J. ? At an altitude of 300 — 500 metres (Fruhstorfer).

The present species, the Java specimens of which are differentiated by
Fruhstorfer as Vulgaroides, is also very common in the low-lying districts.

The larva, which is found on a climber (Gymnema .?/.), is of the Danais
type and on a very intense black ground colour, has numerous small white
round or rhomboid dots which in young larvae, however, are yellow and not
placed on the dorsal part, or only a small number of them, being, conse-
quently, distributed in two groups. Head black. Two pairs of black processes,
dark orange at the base. The larva can move the anterior processes volun-
tarily and independently of each other. The pupa resembles that of the preceding
species to such an extent that I was unable to discriminate between them. The
spots which in some specimens have a golden lustre are silvery in others.

29

7. JuvENTA Cram. (PI. XIV, fig. 24a, 24^, 24c).

Cramer. //, /. 149, //. 188 B (1779) Papilio Juventa.

HoRSFiELD, Cal Le/>., J\f. E. I. C, pi. 3, //>. 7, 7a (1828). Euploea
Moore, Ibid., i, /. 122, //. 4, //^. 4, \a (1857). . . Danais
Snellen, Tijciscln: v. EnL, XXI, p. 6 (1877), XXXIX,

p. 44 (1896)

Distant, Rhop. Mai., p. 407, tab. 39,./4'. 4 (1882 — 1886). Radena „
Moore, Proc. Zool. Soc. London, 1883, /. 224, //. 39,

Jig. I ,, Exprompta.

Fruhstorfer, (Seitz, Grossschvi. d. Erdc), p. 212,//.

78(5 (i9'o) Danais Juventa.

As regards sexual characters this species agrees with D. Similis ; the form
of wings also is nearly the same, the hind-wings, however, being more rounded.
For ^/3 they are light in colour with dark veins, while in Similis they are dark
with light longitudinal streaks. Cramer's figure is of too deep a blue, while
Distant's is fairly good.

W. J. Batavia (3—14); Depok (95); Buitenzorg (265); Tjampea (160);
vicinity of Felabouan Ratou or Wijnkoops Bay on the south coast (+ 1 50) ;
Soukapoura, in the south of the prov. Prayangan.

C. J. Jokjokarta (113); prov. Semarang (300) {Jacobson); Nousa Kam-
bangan {Jacobson).

E. J. Banyoupoutih (45) in the prov. Frobolingo; Kediri (64) ; Jember (98)
in the prov. Besouky ; in the prov. Banyouwangy and in the prov. Madoura.

This species is extremely common at Batavia and elsewhere in the lower
lying districts of Java. I even found it on the coral islet Edam, situate one
hour's distance north of Batavia. The larva occurs on Ij'abe djawa (Fiper
LoNGUM L.); melati longkeng (Fergularia Odouatissima Sm.) ; on kelambennan
(Gymnema sp.}) ; on a climber known as bcbankcan and on a plant called j^rawa;/
oyot. It is dark red-brown in colour covered with numerous milky-white dots
and transverse streaks. The head is a glossy black. From the back of the
second thoracic segment issues a longer and from that of the penultimate
abdominal segment a shorter pair of filiform processes, the anterior pair being
somewhat bent forward. These processes are black except at the base where they
are orange coloured. Sometimes several larvae are met with on the same plant.

The pupa is of a vegetable green with a black caudal hook; a row of
black dots occurs on the dorsal boundary between thorax and abdomen and a
few similar dots are distributed over the back. Those which pupated in a dark

30

box were of an equally clear green as those found in the open. They are
much pestered by ichneumons. From a pupa of 4 April the imago emerged
on the 1 2'^ of the same month.

Horsfield's figure of the larva is poor, that of the pupa a little better.
As to Semper's figures that of the larva, although likewise scarcely accurate,
is better than that of the pupa.

8. LiMNiACE Cram. (PI. XV, fig. 25^', 2^6, 2y).

Cramer, /, /. 92, //. 59, £>. E. (1779) Papilio Limniace.

IV, p. \12, pi. 377. C. D. (1782) ... „ Melissa.

HuBNER, Samml. Exot. Schm., I, pi. 19, c? (1806 — 1827). Limnas Limniace.
HoRSFiELD, Cat. Lcp., M. E. I. C, pi. 3, fig. 6

(1828) Euploea

Moore, Ibid., i, /. 121, //. A* Jig- 3. 3^ (1857) • • Danais

Distant, R/iop. Mai., p. 16, tab. i,/j^. 9 (1882 — 1886). „ Septentrionis.

Staudinger, Ettf/*. Sc/^w., />. 49,//. 24, d'(i884— 1888) „ Limniace.

Moore, Lep. hid., I, p. 30, //. t, fig. i, \a (1890). Tirumala

Stichel, Berl. Ent. Zeitschr., 1 899, Sitz. Ber. i 7 Mai. Danais

Fruhstorfer, „ „ „ 1 899, /. I 1 5 . . .

Martin, Iris, 1910, /. 21 Tirumala Limniace, var. Makassara.

1 9 10, /. 23 „ Melissa, var. Govana.

Fruhstorfer, (Seitz, Grosssckm. d. Erde), p. 202

(igio) Danais Melissa.

Snellen has noted that of the present species the type as well as the
forms Septentrionis and Melissa occur in Java. The Javanese specimens of
the typical form have been christened Myrsillos by Fruhstorfer.

W.J. Batavia (4 — 14); Tjampea (160); Movmt Karang in the prov.
Bantam ; mountains in the prov. Prayangan.

C. J. Touban (4) and Semarang (6) on the north coast; prov. Tegal;
prov. Madioun; Jokjokarta (113).

E. J. Prov. Kediri ; prov. Besouky ; prov. Banyouwangy.

The forms of this species have been dealt with extensively by Fruhstorfer
in Berliner Etitofu. Zeitschr., 1899 and by Dr. L. Martin in Deutsche
Ent. Zeitschr. Iris, 19 lo, where they have based several species and sub-
species on these various forms ; the latter attributes them to climatic influences.
I suspect that here again we have simply to deal with phenomena of colour
evolution ; the specially pronounced spreading of the white, which plays the

31

chief role here, being doubtless nothing else. In the next species and amongst
EuPLOEAS this peculiar development of the white, although in the latter genus
it manifests itself in a somewhat different manner, arises from the same cause.
The material in my possession has not been labelled with sufficient care, as
regards locality and period of capture, to enable a definite judgment to be
formed in this connection. Two types, one darker and larger (SEinENTRioNis)
and one exhibiting more white (Melissa) are clearly separable in Java. At
Batavia, where I collected during an extensive period, I found, both in the
wet and the dry season, exclusivel)^ butterflies of the Melissa type, a specially
large and fine specimen even in August, therefore at the height of the dry
season ; in the Preanger mountains, however, I invariably took specimens of
the Septentrionis form. At the same time I also possess a specimen of
the Melissa form from Tjampea (i6o) and one supposed to have been captured
on Mount Karang, prov. Bantam, together with one from Touban which evidently
forms a transition to the other type. Moreover, the Melissa specimens are
not constant as regards the white streaks and spots and frequendy show trans-
itions to Septentrionis; the white lines in specimens of the latter are very
unequal in width, the white in this form being of a bluish shade.

From Banyouwangy I received a specimen of the form Septentrionis, most
probably not taken in the mountains, and from Besouky, also from the low
lying districts, a specimen which, according to Snellen, mostly agrees with
the form described by Butler from the Samoa Islands under the name of
Obscurata. From Jokjokarta and from Semarang, on the north coast, of
C. J., I possess specimens which, without a doubt, belong to the form Melissa
and agree with it in size but which at the same time are much darker than
others from W. J. and in this respect, therefore, again approach Septentrionis.
Of the earlier stages nothing is known to me personally. Martin observed
oviposition of Septentrionis. A single egg was deposited on each leaf and
affixed by the truncated end, in a similar manner as was observed by me in
the case of Euploea Midamus L. and E. Rafflesi Moore and recorded by
Macpherson of a species of Hest:a. The egg was hatched on the third day
while the larva 8 or 9 days after turned into a pupa of a shining green with
golden spots and a gold band at the boundary of thorax and abdomen. On
the eighth day the imago emerged.

Moore gives the life history of the larva and pupa as it was observed on
the continent of India by Dr. Forsayeth. According to him the larva lives
on Asclepias, Calotropis, and Hoya. To judge from his figure it strongly
resembles that of D. Chryslppus but the median pair of processes is absent;
the pupa also appears to be of the same type as in that species but is larger.

J3

32

9. Melanippos Cram. (PI. XIV, fig. 2-] a, 2-]^).

Cramer, //, /. 44,//. 127, A. B. (i779) Papilio Melanippus.

//, /. 128, /-/. 180, A. (1779) - Hegesippus.

Distant, R/iof^. Mai, p. 19, tab. 2,/io. i (1882— 1886). Danais Melanippus

var. Hegesippus.
Staudinger, Exo/. 5(7/;;/., /. 49,// 25 c? (1884—1888). Danais Hegesippus.
Moore, Lcp. hid., I, /. 49, //. \\,Jig- i, i«, \b{\'6<^o). Salatura
Fruhstorfer, (Seitz, Grossachm. der Ercie), /. 196,

/-/. 77<: (1910) , , Melanippus.

Snellen noted that in the present species cell \a in the fore-wings is
entirely blackish brown on the upper side, whereas in D. Genutia it invariably
shows a brownish yellow longitudinal band.

The form Hegesippus Cram., which exhibits on the hinds wings a strong
diffusion of white, does not occur in Java, although in some of the specimens
of the type, found there, the under side of the hind-wings has a decided
tendency to become white.

The amount of black on the upper side and that of whitish on the under
side of the hind-wings varies in individuals without any inter-relation, however,
and without this variation, apparently, depending on the period at which they
are captured. It is simply a manifestation ot the irregularity in the process of
colour evolution.

D. Melanippus Cram, is evidendy closely related in form to D. Genutia
Cram., although the larvae distinctly differ; it is a species which has advanced
further in the process of colour evolution than the latter, hence the general
colouring on the upper side is of a much lighter orange than is the case as a rule
in Genutia, a few specimens excepted, and hence also on the hind-wings
much more white occurs which becomes especially diffused in the form Hegesippus.
In other species of Danais, not occurring in Java, the original colour has
remained a much more intense red. Thus the entire evolutional fading process
from red to white is clearly demonstrated by a comparison of many species of
the genus.

W. J. Batavia (3 — 14) confined to the lowest districts, in the Pademangan
Wood and even on the coral islet Edam, situate at an hour's distance north
of Batavia; Buitenzorg (265); Tjampea (160) and in the mountains in the prov.
Prayangan (+1000 — 1700).

C. J. ?

E. J. ?

33

The larva is found on kelambennan (Gymnema 5/.) and ouf/ous a/am
(Tylophora cissoides B1.) and differs from most of the Javanese Danais larvae
in having, not two, but like D. Chrysippus, three pairs of processes which are
filiform, entirely black, and placed on the 2°'^ thoracic and the 2°'^ and penultimate
abdominal segments ; these processes are fairly short, the anterior pair being
the longest whereas the two other pairs are very short. The markings of the
larva consists of pearly grey, transverse lines and spots with a sub-dorsal row
of a golden yellow ; it much resembles the larva of D. Genutia but the ground
colour is darker. The head is black. The pupa has the same shape as in
that species, and is green at first becoming subsequently of a light rose colour
with a few golden spots and a line of black, white, and silver mixed, at the
boundary of thorax and abdomen.

10. Artenice Cram. (PI. XIV, fig. 26).

Cramer, IV, p. 168, //. 375, C. D. (1782) .... PapiHo Artenice.
HuBNER, Sanunl. Exot. Schm., II, pi. 1 6,/}V. i —4(1 806 — 1824) Euploea Chionippe.
EscHHOLTZ in Kotzebue's Reise, III, p. 209, //. 7,

fig. i2rt!, (i (1821) , Idea Abigar.

Distant, Rhop. Mai., p. \OQ, tab. 42, fig. i i (1882^1886). Danais Abigar.
Snellen, Tijcischr. v. Etit., XXXIII, p. 269 (1890);

XXXIX, p. 44 (1896)

Fruhstorfer, (Settz, Grossschn. d. Erde), p. 200(1900). „ Afhnis.

In the case of Artenice, Snellen also noted that cell \a of the fore- wings
is entirely dark on the upper side, but that the under side of the hind-wings
differs considerably both from Melanippus and Genutia.

Besides the type figured by Cramer specimens occur in Java either without
white in the median portion of the upper side of the hind-wings or having
that region strongly suffused with white, the latter being the form Abigar Eschh.
The under side of the hind-wings, moreover, contains more white in some
specimens than in others, this difference, to judge from my specimens, being
more accentuated in the 6 than in the v. The name Artenice is specially
applied by Fruhstorfer to the Java form.

W. J. Batavia (3 — 14); in the lower districts near Pademangan Wood this
butterfly is common, but from the higher regions I have received but a single
specimen.

C. J. The species was described by Cramer from Semarang.

E. J. ?

34

II. GiiNUTiA Cram. (PI. XIV, fig. 28a, 2U, 28c, 28^/).

Ckamer, ///, /. 23, //. 206, C. D. (1782) .... Papilio Genutia.
HiiBNEK, Saijuii/. Exot. Sil/m , I, pi. 21, //>. i, 2

(1806 — 1827) Limnas „

HoRSFiELD and Moore, Cat. Lep. M. E. I. C, I, p. 124,

//. 4, //>. 6, ba (1857) Danais Plexippus.

Distant, R/iop. Mai., p. 18, 408, tab. 2, fig. 2, 3

(1882 — 1886) „ Genutia.

Moore, Prov. Zool. Soc. London, 1883, /. 240 . . . Salatura Intensa.

„ /. 242 Sumatrana.

Lcp. hid., I., p. 45, //. 10, fig. I, irt— ^(1890). „ Genutia.

Fruhstorfek, (Sehz, Grossscl/ni. d. Erdc), p. 194, //.

77r (1910) . . Danais Plexippus.

Fruhstorfer applies the name Intensa Moore to the Java form. According
to Snellen, however, the latter, which is relatively small and dark, is indeed
the common form in Java but Moore's Sumatrana also occurs there in which
the colour of the hind-wings is suffused with white. Evidently, therefore, the
same manifestation of colour evolution occurring in so many of the Danaidae.

W. J. Batavia (3 — 14); Buitenzorg (275); Sindanglaya (1074); m.ount
Salak (780); Gedeh mountains (1425).

C. J. Semarang (10); Srondol in the prov. Semarang (300) [Jacobson);
Magelang (500); prov. Madioun; Jokjokarta (113).

E. J. Kediri (64); Jember (98) in the prov. Besouky; Semarou mountains
(750); Tengger mountains (1200) {Jacobsoti); prov. Banyouwangy; prov.
Madoura.

One of the commonest butterflies in Java, being met with everywhere. In
the Transactions Ent. Soc. London, 1905, />. 92, I find a statement by G. B.
Longstaff to the effect that one evening in December, while at Hourou in
British India, he noticed eight of these butterflies suspended close together
from one of the huge leaf-stalks of a palm, evidently for the purpose of thus
passing the night in company and that about twenty of them were found in
close proximity to each other. Such a gregarious habit I have never observed
either in this or any other species of Danaidae. I find a similar statement,
however, with reference to the American D. (Anosia) Plexippus L.

According to Fruhsiorfer the odoriferous scales in the present species
emit the smell of carrion to an extraordinary degree. At times, however, it
is said not to exhale any odour. I have handled hundreds of these butterflies

35

without ever noticing anything of the sort, and I suspect that this observation
is based on an individual which at the time of capture had just been feedino-
on carrion. 1 once caught a specimen of Euploea IMidamus L. which,
evidently for the same reason, smelt strongly of human faeces ; but I have never
noticed this in other specimens of this species.

The butterfly deposits the minute, round, milky-white eggs each separately
on the under side of a leaf, as has been observed in a few other Danaidae ;
I have, however, found as many as four on the same leaf. The larvae feed on
the leaves of ousous ajam (Tylophora Cissoides B1.) ; wc/i?/'/ /owo/fv;/;' (Pergularia
Odoratissima Sm.) ; kelmnbemian (Gymnema .f/.) and also on an ornamental
plant (Ravisteinia Pulchella) frequently met with at Batavia. Many larvae
are found on the same plant ; a shrub in my garden at Batavia was constantly
covered with them and mostly stripped bare, in spite of my frequently killing
great numbers ; I constantly observed butterflies settle on it in order to deposit
their eggs; they manifestly appear to perceive the presence of this plant
from a distance, although the scent emitted by this flowerless and almost leafless
shrub can only be a faint one.

The larvae are mostly light in colour; the ground colour of the back and
sides is black on which occur numerous pearly-grey spots and transverse lines,
and in addition on each segment, except the first thoracic, two golden-yellow
spots, those on the median segments being largest, while the others decrease
in size towards the head and the tail. A yellow band, interrupted by black
lines, forms the boundary between the sides and the black under surface. The
head is black with a light grey dot enclosed within rings of the same colour
which are open at the base. There are two pairs of processes, the anterior
pair on the second thoracic segment being longer and more filiform than the
posterior pair, which issues from the penultimate segment; alle the processes
are light red near the base the remainder being dark red, almost black. I
have noticed the caterpillar voluntarily moving the posterior but not the
anterior pair.

A dark form, which, however, is not so common, has the back dark purple
or brown with faint black transverse lines and a golden yellow subdorsal dot
on each segment. At the boundary between the .sides and the faint green
under surface is a broad white and yellow band in which the black stigmata
are situate. The processes are black but the anterior pair at the base is of
the same colour as the back. An immature larva, which completely agreed
with the immature larva of D. CiiRvsrppus L., changed into this dark form.

The pupae, which completely agree with those of the species just mentioned,
are light green at first with a double band, half white and half black, at the

36

boundary of thorax and abdomen ; subsequently, however, it turns to a light
rose and the white of the band changes into a row of golden dots or pearls.
Whether the surroundings during pupating are light or dark exercises no
influence upon the colour of the pupa. The caudal hook is black and close to
it four black spots are to be observed. A chrysalis of 2 1 F'ebruary produced
the imago on March i while from pupae of 23 February the perfect insects
emerged on March 2 and 3. The description given of the larva of this species by
Moore does not agree with mine and has reference to a caterpillar with 3 pairs
of processes, like that of D. Melanippus Cram. The same applies to the figure
given by Horsfield, also based on a specimen from British India and this
fact makes it appear probable that the Javanese Genutia is not identical with
that from British India.

12. Chrysippus L. (PI. IV, fig. 29^!, 2C)b, 2qc, 291/, 2g<', 29/).

Linnaeus, Sysl. jVaf., Ed. X, /. 471, A^d. 81 (1758). . Papilio Chrysippus.

Cramer, //, /. 32, />/. 118, B. C. (1779)

Stoll, Stipp/. op Cramer, /. 132, //. 28, //>. 3 (1790) . „ Petilia.

Hubner, Saimnl. Eur. Sckm., pi. 133, fig. 673, 679

(18 1 6) „ Chrysippus.

Hubner, Samtiil. Fxot. Schm. I Limnas

Horsfield, Cat. Lcp., M. E. I. C, pi. 3, /i^. 9, 9a (1828). Euploea
Horsfield and Moore, Ibid., I, p. 126, //. 4, fig. 7, 7a

{1885) Danais

Distani-, Rliop. Mai, p. 20, 408, tab. i, fig. 10, tab.

AO, fig. 13 (1882—1886) ,

Moore, Proc. Zool Soc. London, 1883, /. 238 .... Limnas Bataviana.

„ Lep. Ind., /, /. 36, //. 8, //V. i, \a — e (1890). „ Chrysippus.

Fruhstorker, (Seitz, Grossschm. d. Erde), p. 193 (19 10). Danais „

The specimens from Java belong to the dark form BArAviAN.\ Moore ; the
shade of the principal colour varies, however, some 6 especially being con-
siderably lighter in hue, although not so light as the typical form which occurs
elsewhere. The process of fading to yellow and white in this species generally
manifests itself strongly over its very extensive range and thus produces all
manner of so-called varieties, which, however, have partly been obtained also
by breeding from tj'pical parents. Thus, for instance, the forms Alcippus and
Alcippoidhs, which do not occur in Java, have been distinguished. The form
Petilia Stoll, from the east of the Archipelago and North Australia, is also of

37

a light ground colour. In a consignment, which I received once from Tegal
and which otherwise contained only common Javanese butterflies, amongst them
the form of D. Chrysippus L. common in that island, I found also a fresh
specimen of the form Petilia, agreeing in every respect with those from North
Australia, but which was said to have been captured by Mr. van Bruggen
on the northern slope of Mount Selamat, at an altitude between 1 70 and
1000 metres. Considering, however, that this is the only specimen from Java
known to me and that, moreover, the species most probably is a traveller, I
would deem it advisable for the present not to draw any conclusions from
this fact.

The dark form of Java differs very markedly in the shade of its colour
from Hypolimnas Misippus L., which, it is alleged, mimics this Danais.
Moreover, the flight of the latter, which flies low down over grassy plains, and
of a tree frequenting species like this Hypolimnas, is by no means the same
and for this reason thev do not occur together except accidentally.

W. J. Batavia (3 — ^14); Buitenzorg (275); Bidara Tjina (28); Megamen-
doung mountains (1480); Soukapoura in the south of the prov. Prayangan.
C. J. Prov. Tegal; Magelang (500); Jokjokarta (113).
E. J. Kediri (64) ; prov. Madoura.

According to Moore this butterfly is extremely common on the continent
of India; Mr. Aitken, quoted by him, states "with the exception, perhaps, of
Terias Hecabe, the commonest and most ubiquitous butterfly on the Bombay
Side of India. Up to a level of 7000 feet it may be found anywhere in the
Indian Empire".

This is not so in Java; in the mountains this butterfly occurs indeed but
by no means frequently ; on the plains of the low lying country it is, indeed,
nowhere rare but very common only in some districts and never equally
common as in British India.

According to Moore the egg is cream coloured, sugar-loaf shaped and
deposited singly on the under surface of the leaf. In the caterpillar when
hatched the tentacula are barely visible; but in the full grown one the tentacula
are long and movable ad libihun by the caterpillar. The latter fact I have never
observed in the numerous specimens reared by me. According to my notes
the larva, which I also found in July at the height of the dry season, lives son a
plant designated at Batavia as rembeo^a and bidoura (Calotropis gigantea Ait.),
belonging to the Asclepiadeae. Species of Asclepias are also indicated as its
food plant by Semper in the Philippines, by Trimkn in South Africa, and by

38

Hoffmann in Greece, while Moore mentions Asclepias curassavica for
British India. The caterpillar is pearly grey on the back with many transverse
black lines varying in width, between which on each segment occur two oblong,
sometimes coherent, golden -yellow spots. Along the sides it has a golden-
yellow band across which the foresaid lines partly extend ; a black band divides
the sides from the light grey under-surface. On the pearly-grey head may be
observed black markings resembling those of the caterpillar of Euploea MiDAXtus L.
Three pairs of processes occur on the back, /. e. on the 2"'^ thoracic and on
the 2"'' and penultimate abdominal segments ; they are filiform, the anterior
longest and the middle pair shortest; they are black and, except the anterior
pair, somewhat reddish at the base. In the immature larvae the )'ellow on the
back has not yet formed distinct spots but appears as narrow transverse lines.

The larvae grow very rapidly and change into pupae which very much resemble
those of D. Genutia Cram_ The general colour of many is at first milky
white and becomes subsequently a beautiful green which again fades at times
when the pupae become white. Other pupae are light rose. At the boundary
between thorax and abdomen a double line may be observed which is black
on the side of the thorax but on the side of the abdomen consists of numerous
golden dots placed side by side. On the thorax and the wing-cases some
additional golden dots are found. The caudal hook is black.

As this butterfly inhabits a very extensive area there are many figures of
its earlier stages. Those of Trimen (South Africa) are very good, and the
description he gives of the larva agrees very well with the Java caterpillar. Of
the South European form the larva and pupa have been described and figured
by HiiBNER and by Prof Dr. E. HoFiMANN ; Hubner's description and figures
agree well with the Javanese except that all the processes of the caterpillar
appear to be longer than those of the Java specimens. Hofmann, in his
description, mentions only two pairs of processes but he figures three pairs of
equal length. He also mentions flesh-coloured and green pupae but his figure
of the pupa is very poor. The figures of Semper (Philippines) and of Horsfield
(Java) are very unsatisfactory ; nor are those given by Moore of the^Ceylon
caterpillars and chrysalides beautiful whereas those of the larva from British
India, while not being accurate either, are somewhat better. The illustrations
of the larva and pupae which I now offer are much more successful.

SATYRIDAE.

The Java Satyridae are butterflies which, with a few exceptions, do not
seek strong sunshine but rather frequent shady places and fly also when the
sky is overcast ; some species are actually met with in those parts of the
mountains which are mostly enveloped in clouds and are consequently very
foggy and damp. Some are on the wing during morning and evening twilight,
but in day time onl)' when disturbed and since the latter is the case with
many other species it is very probable that these also are twilight butterflies.
Of the larvae we have but a very incomplete knowledge, but they appear to
live principally on graminaceous plants.

Genus CYLLO Bsd., Herr. Sch.
I. Leda L. (PL XV, fig. ^oa—sor).

Linnaeus, Sys^. Nat., Ed. XII, I, 2, p. 773, No. 151

(1767) Papilio Leda.

Drury, ///. Euf., I, pi. 15,//;.. 5, 6 (1773) ,

Cramer, /, /. 40, //. 26, A. B. (1779) Ismene.

///, „ 5, //. 196, C; IV, p. 8, //. 292, A.

(1779, 1782) „ Leda.

IV, „ 8, //. 291, F. (1782) „ Mycena.

IV, „ 8, //. 292, B. (1782) Phedima.

IV, „ 8, //. 292, C. (1782) „ Arcensia.

HuBNER, Saviinl. Exot. Sc/im. / (1816— 1 8 1 7) . . . . Oreas Leda.
HoRSFiELD, Cat. Lip., M. E. L C, pi. 8, //V. 9(1828). Hipparchia Leda.
Distant, Rhop. Mai., p. 41, tab. 4,//V. 10(1882 — 1886). Melanitis
" .. .. A 42, tab. 4, fig. 9, II, 12,
(1882— 1886) „ Ismene.

14

40

Moore, Lep. Ind., II, p. ii8, />/. 122, 123 (1894) . . Melanitis Ismene.

KoBus, Tijdschr. v. En/., 39, /. 129, //. 6 (1896) . . Cyllo Leda.

BiNGHA^r, Fauna Brit. India, A /. 158 (1905). . . . Melanitis Ismene.
Fruhstorfer, (Seitz, Grossschm. d. Erde), p. 362(1911). „ Leda.

This species is extremely common in Java, where the food of the cater-
pillar is met with everywhere. It is, therefore, unnecessary to enumerate
specific habitats. Moreover, it has a very wide distribution, over the whole
Indo-Australian region as well as elsewhere, to the north as far as north China
and Japan, to Australia in the south, eastward on many of the South Sea islands,
and westward in tropical Africa. The butterfly during morning and evening
twilights flies low near the ground, in the day time only upon being disturbed
when it settles again at a short distance on the ground or on some object.
The peculiar coloration of the under side of the wings, which varies consider-
ably in individuals, makes it difficult to distinguish it from its environment,
which, of course, is considered mimicry although this theory fails to explain
how such great differences in colouring and marking originate. The butterfly
is frequently met with in dwellings, where, after the habit of many Heterocera
it seeks to evade strong daylight. Distant mentions that it greedily sucks
the sweet liquid tapped from palm trees, known as palm wine ; in Java this is
also the case.

In certain individuals, which have been considered a distinct species under
the name of C. Ismene, a remnant of the original universal red colour is to
be observed in the bright orange on the upper side of the fore-wings near
the apex ; frequently this has even faded to yellow or is covered in great
part, if not entirely, by black. The unequal course of the process of colour
evolution, according to the individuals, explains this difference, which manifests
itself still more strongly on the under side of the wings where it causes the
great variation in colouring and marking already alluded to ; this variation has
been attributed to difference in sex or in time of occurrence by authors unac-
quainted with this process, some even considering these differences of specific
value. De Nickville and Moore have in this manner created several species ;
Bingham has, it is true, reduced the number somewhat but not sufficiently.
This is, indeed, unavoidable when one is unacquainted with the process of
colour evolution. For even if it be admitted that existing animal and vegetable
forms are not immutable but may vary through climatic or similar influences
and, therefore, especially in proportion as they inhabit different regions, the
fact that a species, apart from such influences, should be able to become
modified, so to speak of its own accord, appears incomprehensible. Unless it

41

should be conceived as an extraordinary, inexplicable fact — a so-caled mutation —
instead of the result of a normal process of natural evolution. In that case
one is at a loss how to treat these numerous forms except to differentiate
them as numerous species or at least as races, which, however, are by no
ineans well defined but graduating into each other constitute but a confused
entity. It is only through grasping the principle referred to that it becomes
possible to understand these varied forms. It is true that in the case of a
species exhibiting such an amount of variation, like Cyllo Leda L., this is
by no means an easy task; I believe, nevertheless, to have succeeded in this.

By a comparison of more than i 50 specimens from Java, aided by that of
about 70 from other regions I have convinced myself that the Cyllo butterflies,
occurring in Java, with the exception of a small number, forming a distinct
species — Cyllo Suyuuana Moore — all belong to the species Cyllo Leda L.
Neither do I attach any value to the subspecies, which Fruhstorfer recognizes.
With regard to the majority of species specially figured from British India by
Moore, I must also doubt their independent existence; considering, however,
that these do not occur in Java I have been unable to study them and, conse-
quently, I am unable to form a definite judgment in this respect.

Two courses of evolutionary modification are to be observed with regard
to this species in Java. One being the decrease in size of the wings which I
have explained in my paper " lleber die sogenannien Schivdnze der Lepidopterc?i" .
{Deutsche Entomologisclie Zeitschnft Iris 1903), and to which I hope to refer
again when dealing with the Papilionidae. The continuous operation of this
process manifests itself here principally in the gradual disappearance of the
projecting points, the vestiges of a former greater expanse of the wings. In
many individuals of Cyllo Leda L. such appendages are still fairly evident
(pi. XV, fig. T,oo,p,g), but in others, especially in the fore-wings, they become
considerably smaller. Entomologists, who are insufficiently acquainted with this
evolutionary process, are constrained to base distinct species on these different
forms of wing; in this manner Elymnias Caudata Butl., for instance, although
nothing but an old form of E. Undularis Drury, is looked upon as a separate
species. These appendages, however, are nothing but such wing vestiges which
in many Papilios have remained in the form of the so-called tails, sometimes
even — as in the Achates form of Papilio Memnon L. — only in an old
9 form ; an instance which clearly proves that of specific difference there can
be no question here. Thus these appendages are still strongly in evidence in
9 specimens of Cyllo Leda L. having made little evolutional progress in this
respect and still exhibiting, therefore, an old form, which, moreover, like the
old Achates form of Paitlio Memnon L., have also sometimes remained

42

somewhat larger, thereby leading principally to the creation of the so-called
species Cyllo Zithenius Herbst, while others again have regarded it merely $ of
Cyllo Suyudana Moore. The other process, that of colour evolution, manifests
itself at first in the fading of the original red pigment to orange and yellow,
after which this pigment — whether or not first fading to white, of which there
is no evidence — appears to be broken up but in such a manner that the
scales, which contained the pigment, or some of them, become filled with air
and thus acquire an intense white colour. Side by side with this, as usual, the
black pigment at first strongly suffuses itself over the original colour, covering
it completely or nearl}' so but subsequently decreases again sothat either the
black investment becomes lighter in colour or the original colour — mean-
while having faded to yellow — reappears, even though still suffused with some
black. The upper and the under sides of the wings independently pursue their
course in this process so that any specific pattern of marking on the under surface
goes side by side, now with one then with another pattern on the upper surface.
Thus the following combinations may be met with :

upper side, A (PI. XV, fig. 30c) under side, A', E, D, F, H, G, I, K' (PI. V,

fig. 30 d, i, h, k, I, m, 71, p).

„ „ B (PI. XV, „ 30.') „ „ C, E, D, F, H (PI. V, fig. 30 .c.

/, //, k, I).

„ „ C (PI. XV, „ 30/-) „ „ C F, H, K' (PI. V, fig. 30-, k.

It is only in the very pale specimens, mostly V, figured on pl. XV, fig. 30 f, g,
that some connection appears to exist, at least as regards the general tint,
between the upper and under sides which, however, simply indicates an equally
advanced stage in the process of colour evolution. In most cases that is to
say, since there are also specimens with such an upper side, but whose under
side have nothing in common with the latter.

Now first as regards the upper side, it is covered by the black pigments,
either completely or with the exception of a small portion near the apex, to
such an extent that of the original red colour nothing remains or only a spot
faded to orange or yellow, while subsequently this black colour again has
commenced to recede and in many specimens in this manner has become a
light grey. This process is met with especially, but not exclusively, in 9 (PI- XV,
fig. 30/), since I possess also f? in which it occurs, the former, therefore,
having reached apparently a more advanced stage than the majority of
the latter. The reddish, orange, or yellow spot referred to, which in its turn
is margined by intense black containing two clear white dots, varies consider-
ably as to size and shade in individuals, according as to whether the process

43

of diffusion of fhe black pigment has advanced more or less. In specimens
where the black is still on the increase this spot is most pronounced during
the dry season when in general the development has less advanced than in
individuals living in the rainy season, hence the form Ismexe was believed to
be a dry season type. It is, however, invariably of small size ; such large
yellow spots as figured by Moore for many of his alleged species from
British India and which also occur in races of Cyllo in the west of the
Malay Archipelago, are not met with in Java; only in some $ from E. J.
(PI. XV, fig. 305?) the orange shade is likewise fairly diffused. These are,
however, very useful for understanding the existence and history of this spots
in Cyllo Leda L., by means of comparison with them, as has been done in
the introduction. We have here, evidently, to do with what in the introduction
to mv monograph of the Java Pieridae I have termed obstinate spots, in which
the original colour — whether red or, at a more advanced stage, orange or
yellow — is retained for a long period before being covered by the black
pigment. The intense black b)' its side must, as has been explained in the
introduction with reference to the true ocelli, apparently be looked upon as an
accretion of this black pigment at the place where the said obstinate spot caused
an obstruction to its even diffusion in the manner this has taken place in the
remainder of the wing surface. The clear white dots occurring in the spot
referred to must have originated in the same way as has been indicated
in the introduction in the case of the true ocelU. At times similar white
dots occur elsewhere on the upper side and it may happen that these are
not clear white but yellow, in which case the scales are still filled with some
faded and, therefore, yellow pigment. To utilize such white dots as specific
characters simply signalizes a complete want of grasp of the process of colour
evolution.

The course of this process on the under side is much more intricate. Here
also the original red is covered by the black pigment and the latter has subse-
quently decreased step by step, but in so uneven a measure that it occurs
in all manners of blends with the remnants of the original colour, in which the
latter — although sometimes already faded to yellow — is still distincdy
discernable. Two vertical, parallel, curved stripes on the under side ot the
tore-wings are first of all to be regarded as vestiges of the former general
darkening process, the inner one being continued on the hind-wings, while both
occur in various stages of fading and diminution, being wider at one time
(PI. XV, fig. 30//, w) and narrower at another (PI. XV, fig. 30 X-, /, ;/) ; some-
times they are very dark (PI. XV, fig. 30 //, ;//) but not unfrequently the
original red is distincdy recognizable, more so than is shown in the illus-

44

tration (PI. X\^ fig. 30 //, />). These stripes may even disappear entirely or
nearly so, although in that case more or less isolated spots of black pigment remain
(PI. XV, fig. 30rt') in a similar manner as is the case in Pieridae with the
brown spots, produced by the same process, on the under side in species of
Callidryas and Terias. {Pieridae, PI. Ill, fig. 6/, 6/, ^b, and PL IV,/ig. s**).
Frequently the diminution of black at that spot proceeds more evenly giving
rise to the kind of marking termed " /^iese/u/ig " by German Entomologists, where
the black pigment has been retained in so finely divided a condition reminding
one of the minute drops of water of a drizzle, a form of marking which is
also met with in many other butterflies, for instance on the under side of the
hind- wings in Iphias (Hebomoia) Glaucippe Bsd.

If next the black pigment continues to diminish, the original colour no
longer covered and faded to yellow, now reappears, (PI. XV, fig. lOif^), so that
the individuals which are far advanced in this evolutionary process exhibit
yellow on the under side now blended with but little black, and in these the
upper side is faded to a light grey. (PI. XV, fig. 10/).

The ocelli, which have been fully dealt with in the introduction, especially
deserve attention in this connection. Those most highly developed are
found in individuals whose under side has reached an advanced stage of
"drizzling", for the most part where the original colour universally reappears
as yellow. (PI. XV, fig. 10^,'). As a matter of fact all these forms of Cyllo
Leda L. can only be regarded as the same expression of excessive inequality
in the individual manifestion of the evolutionary processes of form and colour
which give rise to the polymorphism in many other butterflies but which in
C. Leda L. presents a greater amount of confusion. Thus, for instance, the
origin of the projecting appendages on the wings already referred to, and at
one time more pronounced than at others, is the same as that of the so-called
tails in the Achates form of Pap. Memnon L. Three circumstances co-operate
to make this less manifest in this case :

1 ^. That in C. Leda L. the colour of the upper surface and that of the under
surface of the wing^s in each individual continues its evolutional modification
without, apparently, any inter-relation; in the cases of polymorphism, as in
Pap. Me.m.n'on L. or Callidryas Pomona L., this holds good likewise but by
no means so universally, since in most cases a definite pattern on the upper
side corresponds to a definite pattern on the under side.

2" That, although in the two species mentioned, for instance, there can
be no (juestion of sexual polymorphism and this difference of form has nothing
to do with sex itself, this difference, nevertheless, manifests itself in the sexes
of these butterflies in various stages of development and thereby facilitates the

45

appreciation of these evolutional modifications; whereas in C. Leda L. no
difference in this respect manifests itself.

3*^. That the difference in colouring in C. Leda L. arises almost entirely
from a greater or lesser diffusion of the black pigment, frequently in a finely
divided condition over a less clear ground colour, making it more difficult to
distinguish a definite colour pattern than is the case with the polymorphic
species, referred to, with their lighter and more distinct colours.

The fact that this solution of the Cyllo problem simply carries us back to
the same evolutionary processes which manifest themselves in other butterflies,
undoubtedly presents a strong presumption in favour of its correctness ; that,
moreover, only an acquaintance with these processes and especially with the
theory of colour evolution can lead to such solution is also incontestably a
strong argument on behalf of its accuracy.

I imagine that the foregoing observations will enable the reader to appre-
ciate my conviction that the numerous species of Cyllo invented by Moore,
de NicKviLLE and others, each provided in addition with a dry-and wet-season
form, do not in reality exist and simply indicate various stages of development
in one and the same species. Together with Cramer's species Phedama,
Mycena, and Arcensia, which simply represent colour forms of Leda, I am
constrained to reject Moore's and Bingham's Melanitis Bela, under which the
latter, moreover, includes Moore's species M. Varaha, M. Gokala, and
M. Fambra ; Hkewise Fruhstorfer's subspecies Simessa, Lacrima, and
I3MEN1DES. Even Cyllo Zithenius Herbst, mentioned by Fruhstorfer under
the forms Gnophodes Btlr. and Gruescens from Java, is simply based on
wrongly understood specimens of C. Leda L. For a long time I have been
in doubt concerning the form K (PI. XV, fig. 300) of which I have received
from the mountains of W. J. numerous specimens differing sVightly />//er se and
invariably exhibiting the same under side K' (PI. XV, fig. 30/). Seeing,
however, that the marking of this under side agrees completely with that of
man)' indubitable Leda 9 and that transition forms occur besides, I was unable
to separate this form either from Cyllo Leda L. Cyllo Suyudana Moore alone,
which is not the same as Phedima Cr. with which Fruhstorfer confuses it,
I recognize as a distinct species since no transitions between this and C. Leda L.
have come to my knowledge, although both are very closely allied in their
colour development. The larger specimens from the Moluccas, however, I
cannot regard otherwise than as a race of the same species, a view which I
am prepared to modify only if, on examination of the generative organs,
specific differences are shown to exist.

Of the different forms of this butterfly occurring in Java I have figured the

46

most divergent; it should, however, be borne in mind that illustrations are
never perfect, and that especially the various shades of colour can scarcely be
reproduced with accuracy.

As regards the larvae I possess about 25 descriptions noted by me at
different times, which I have compared with descriptions by Moqre, de Niceville,
Bingham, and Rainbow, as well as with the figures contained in the works of
these authors and of Horsfield and Kobus and finally with those prepared
under my supervision in Java. From this it is evident that of this species
two larvae are rarely alike. They are long, slender, fusiform in build, the
terminal segment is armed with two divergent setose processes; the head
prominent and armed with two erect obtuse setose processes, like little horns.
The body appears rugose and covered with short microscopic white hairs
which give the appearance of minute white dots on the surface of the body.
The colour is green, sometimes lighter, sometimes darker, on which occasion-
ally the line of the vas dorsalis is visible dorsally while along the back and
sides occur some, occasionally dark, but mostly yellow or white lines, the
principal one of which runs along the ventral boundary and is more or less
raised. All these lines, however, are sometimes only partly visible or even
not at all. The ventral side is dark green covered with fairly long, stiff hairs.
The bristles on the terminal processes are as a rule of the colour of the body
but sometimes they are brownish ; the horns are reddish brown, of a more or
less dark shade, sometimes very light brown; this colour, in most cases, is
continued form the horns along the head in the shape of two bands, uniting
below the head. In addition a milky white band runs along the outer side of
the horns, being also continued on the head and running side by side with
the brown band. It varies, however, exceedingly, sometimes running only along
the head or only along the horns while frequently it is reduced everywhere or
here and there to a narrow line. It may at times, too, be yellow instead of
white. On one occasion I found some black marking on the front of the head.
In young larvae as a rule the horns, and sometimes the head, are black,
which colour is retained until the last moult ; occasionally the horns were pale
rose coloured and the milky-white band was already present.

Sometimes the caterpillars occur gregariously, for instance, on padi (Orvza
Sativa L.) and teboii (Saccharum officinarum L.) but they are also frequently
found solitary on various kinds of grasses, Bambusa, palm leaves and even on
iapak limau (Elephantopus Scaber L.). They are voracious and change into
light green chrysalides with some faint white stripes on the wing cases, without
any processes, and attached to leaves or grass haulms, frequently in numbers

47

close together. From pupae of 27 April and 20 May the imagines emerged
on 6 and 28 May.

Moore's figures of the imago are good but those of Kobus poor. None of
the figures of the larva referred to appears to me a good likeness, nor are those
prepared for me. On the other hand I am able to reproduce here (PI. XV,
fig. 3 If), a very successful figure of the larva of Cyllo Suyudana Moore which is,
indeed, larger than any specimen of C. Leda L. I have ever seen in Java but in
which, otherwise, I can discover no characters differentiating it from that species ;
it is true the white lateral band along the ventral boundary is particularly
distinct and evidently somewhat raised, but similar bands sometimes occur in
the larva of C. Leda L. Like that of the latter it is rugose but this is
difficult of reproduction, which is clearly proved by Kobus' figure.

2. Suyudana Moore. (PI. XV, fig. a, b, c).

Moore, Cat. Lep., M. E. I. C, L, /. 224 (1857) . . Melanitis Suyudana.

Lep. of Ceylon, I,/, i 5, />/. 9,y/V. 2, a— <■ (1880). „ Tambara.

Distant, Rhop. y^a/., /. 41 2, /a^^. 39, /;V. 2 (1882 — 1886). „ Suyudana.

„ /. 413,//. 19, %3 ( „ ). „ Abdulla.

Staudinger, Exot. ScJwi., p. 222, pi. 79(3" (1884 — 1888). „ Suyudana.

Fruhsiorfer, (Seitz, Grossscfwi. d. ^^v/^),/. 364 (191 1). „ Phedima.

Fruhstorfer confuses this species with Phedima of Cramer which represents
a badly coloured 9 of C. Leda L. ; his alleged seasonal forms, consequently,
are of no value. I possess this butterfly from West, Central, and East Java,
the majority of specimens being from the mountains, but one was captured in the
prov. Semarang, very near the north coast, in low lying country, therefore. I
have long been in doubt whether this species should not be considered a form
of Cyllo Leda L. A careful comparison of 19 specimens of the present
species has, however, satisfied me that, even when closely allied to Cyllo
Leda L., it constitutes an independent type, although one specimen may be a
little darker and another somewhat lighter on the upper side, while on the
under side also they differ and the pointed appendage of the fore-wings is very
pronounced in some individuals, whereas in others it is entirely wanting.
Although the manner in which the evolutional modification of its under side
is effected recurs in C. Leda L. no transitions between these two species are
met with. A careful examination of the genitalia of these butterflies conducted
on my behalf by an expert on the subject. Dr. Henri de Graaf of Leyden,
revealed the fact that of these i g specimens only 2 were V and that the latter

15

48

were only a trifle less dark than the c?. The caterpillar found by me on
padi (Oryza Sativa L.), although otherwise greatly resembling that of Cyllo
Leda L. and described with it supra, decidedly differed from it by its greater
size. For these reasons I consider C. Suyudana Moore may be admitted as
a distinct species.

Genus ERITES Westw. Herr. Sch. ')

I. Madura Horsf (PI. XIV, fig. 32).

HoRSFiELD, Cat. Lcp. M. E. I. C.,pl. 5, //>. 8, 8a (1828). Hipparchia Madura.
Westwood, in Doubleday and Hewitson, Gen. D. L.,

p. 392, footnoote (1847) Erites Madura.

DE Nic^.viLLE, Journ. Asiat. Soc. Bengal, LXII, /. 3

(1893) Medura.

Fruhstorfer, (Seitz, Grosssc/nn. der Erde), /. 302 . . „ „

W. J. Buitenzorg (265); Gedeh mountains, in the south of the prov.
Prayangan near Pelabouan Ratou or Wijnkoops Bay and Tjiletou or Sand
bay (+ I 50).

C. J. ?

E. J. Prov. Besouky; Semarou mountains (740); Malang (440) {FnihsL).

The flight of this butterfly is close to the ground, agreeing completely
with that of the genus Ypthima. According to Fruhstorfer it is verj'
common in the south ot Java especially in E. Java, but I have never seen a
specimen from the northern districts of that island. The ocelli on the upper
side of the hind-wings do not occur as regularly as shown in the illustration
nor are they all invariably of the same size.

2. Ocmreana Semp. (PI. XIV, fig. 33).

Semper, Schmett. d. Phil., p. 326, No. 497

(1886 — 1892) Erites Ochreana.

Staudinger, Iris, II, />. 38 (1889) „ Madura var. Ochreana.

') In the Leyden Museum is a specimen of E. Elegans Butl., labelled Java, without any
further indication. As no other specimen from Java is known to me and old labels in that
Museum are not always reliable, I am unable, at present, to accept it as a member of the
Javanese Fauna.

49

DE NiCKViLLE, Jouni. As. Soc. Be7igal, LXII,

/. 5 (1893) Erites Ochreana.

Moore, Lep. liid., II, /. 113,//. 121, Jig. la—b

(1883)

Areentina.

'&

Bingham, Fauna 0/ Brit. India, \,/>. 153 (1905). „ „

Fruhstorfer, (Seitz, Grosssc/im. d. Erdc), p. ^OZ^

pi. 93 (191 1) ., Fruhstorferi.

I only possess a single specimen captured in the southern part ot W. Java
near Pelabouan Ratou or Wynkoops Bay (+ i 50). Mr. Fruhstorfer obtained
it at the same place as well as In the mountains along the south coast of E. Java.

Genus YPTIMA Hb. Westw.

Small butterflies which fly close to the ground and whose larvae, as far as
known, occur on kinds of grass. They are partial to sunshine but they are
on the wing also when the sky is overcast. Characteristic in them is the great
apical ocellus on the fore-wings in which generally two blue dots may be
observed, being the nuclei of the two ocelli whose coalescence has given rise
to the larger eye-like blotches. In some specimens the two, although already
connected, are still clearly distinguishable but where their coalescence is further
advanced the large ocellus acquires consequently an oblong shape and the
greater the size of the butterfly, and with it the apical eye-like blotch, the
rounder does the latter become. In the cf it is usually smaller than in the ?.

I. Jarba de Nic6v. (Fl. XIV, fig. 34)-
Fruhstorfer, (Seitz, Grossschin. d. Erdc), p. 287,//. q(^d, f. Ypthima Jarba.

Fruhstorfer distinguishes two forms of the present species: Gangamela
from E. J. and Eui-eithes from W. J. The former, however, I met with in
both sexes in W. J. and of the latter I received a cf from that entomologist.
I am unable to find any difference of importance between the two. Fruhstorfer
figures a particularly large $ of Eupeithes, not larger, however, than $ speci-
mens of Y. Pandocus Moore, which occurs in W. J.

50

2. HuEBNEKi Kirby. (PI. XVI, fig. 35).

HiiBNER, Ztitrdge, I,/. 17, /'(<;. 83, 84 (1818). . . . Ypthima Philomela.
KiREY, Caf. Diurn. Lep., /. 95, N^. 18 (1871) . . . „ Hiibneri.

Distant, R/iop. Ma/., p. 57, tab. T,/ii;. 5 (1882 — 1886).
Moore, Lep. Ind., II, /. 77,//. \\\,fig. i, la— //(1893). „ „

Elwes and Edwards, Trans. Enl. Soc. London, 1893,

p. % pi. \, fig. <^

BrNGHAM, Fauna Brit. Lidia, I, />. 142 (1905). ... „ „

Fruhstorfer, (Seitz, Grossschm. d. Erde), p. 287,//. 99^

(191 1) „ Philomela.

W. J. Batavia (3 — 14); Meester Cornells (25); Depok (95) ; Buitenzorg (275);
Patjet (11 14) {Jacobson).

C. J. Semarang (6); prov. Jokjokarta {Jacobson); Salatiga (580); (^«<(?(i5(?«).

E. J. Jember, prov. Besouky (98).

The present species is common everywhere in the low lying country ; I
also received specimens, however, from the mountains. Moore and Bingham,
following DE Niceville, recognize definite forms of the dry and wet season.
I am unable to accept the view of these entomologists. I possess many specimens
captured both at the height of the wet and in the middle of the dry season
and can find only individual differences between these. Some specimens are
somewhat darker in colour on the upper side than others, but these occur as
much in the dry as in the wet season. The under side also in some is
more whitish than in others, a feature which appears to manifest itself especially
in $. There is much difference in the size of the oceUi, both as regards the
large apical eye-like blotch and that on the under side of the hind-wings;
some of the ocelli at times are in contact while at others they are separate.

In nine specimes caught at Semarang in August and the beginning of Septem-
ber — therefore at the height of the dry season — the ocelli on the under side of
the hind- wings differ, indeed, somewhat in size but are otherwise quite similar
to those in specimens taken in the middle of the rainy season. I also possess
specimens captured in March — therefore in the wet season — in which these
ocelli are less developed than in the others.

The caterpillar feeds on rounipotit paliit (Paspalum Conjugatum Berg). It
is hairless and dark ochre-yellow, somewhat darker at the sides and with a
faint dark dorsal line. That green individuals occur, as figured by de Nice-
ville and Moore is, however, possible. The chrysalis is green and, according
to DE Nic£vii.LE, sometimes also brown. From a pupa of 29 June the perfect
insect appeared on 10 July.

51

3- Argillosa Snell. (PI. XVI, fig. 36a, ^bd).

Snellen, Tijdscln: v. EnL, XXXV, /. 133 (1892). . . Ypthima Argillosa,
Fruhstorfer, (Seitz, Grossschm. d. Erde), /. 290(1911).

W. J. One specimen, without precise indication of its habitat.
C. J. Touban (2), on the north coast; Dander (30).
E. J. ■>

4. Baldus F. (PI. XVI, fig. na, i^jb, 38a, 38^, 38^-, 380?).

Fabricius, Syst. Eiit., Append., /. 829, (1775) . . . Papilio Baldus.
Donovan, Ins. of India, pi. 36, fig. 2 (1800— 1803) .

Distant, Rhop. Mai., p. 57, tab. 5 (1882 — 1886) . . Ypthima Neuboldi.
„ />. 56, 419, tab. t, fig. 9(1882 —

1886) „ Methora.

Waterhouse, Aid, pi. i-](), fig. 3 (1886 — 1890) . . „ Horsfieldii.

Snellen, Tijdschr. v. Ent., XXXIII, /. 286 (1890). . „ Baldus.

Moore, Lep. Ltd., 11,/. 58,//. \ob, fig. i, i«— ?'(i89o). Thymipa
Elwes and Edwards, Tians. Ent. Soc. London, 1893,

/. 28, //. I, fig. 21, 22 . . Ypthima Leuca.

p/. 1, fig. 1, la II „ Horsfieldii.

Bingham, Fauna Brit. India, I, />. 134 (1905) ... „ Baldus.
Fruhstorfer, (Seitz, Grossschm. der Erdc), p. 289,

//. 99^/ (191 1) » »»

W. J. Batavia (3 — 14); Depok (95); Buitenzorg (265) ; vicinity of Pelabouan
or Wijnkoops Bay (+ 1 50).

C. J. Semarang (6) ; prov. Jokjokarta {Jacobson) ; Gounoung Kidoul
{yacobson).

E. J. Jember, prov. Besouky (98).

Fruhslorfer recognizes two races of this species which he designates
Baldus F. and Horsfieldi Moore and which, in fact, are quite distinct. Of
the latter I possess many specimens from the low-lying country of W. Java,
where it is very common and likewise from Buitenzorg, also some from the
prov. Jokjokarta in C. Java and a single one from Jember, E. Java. Of the
other form I have eleven specimens of both sexes taken at the height of the dry
season at Tjandi, the upper town of Semarang, and one in February — in the

52

middle of the rainy season, therefore — on the Gounoung Kidoul, prov.
Jokjakarta, so that I am unable to admit the separate wet-season race Marshalli
Butl. of Fruhstorfer, at least as far as Java is concerved. Neither can
Baldus and Horsfieldi be regarded as distinct species, since a specimen, taken
by me at the height of the rainy season in the province of Jokjokarta, is
clearly intermediate to a certain extent between the two, the upper side as
well as the pronounced ocelli on the under side agreeing entirely with those in
Bai.dus, while, on the other hand, the strong white powdering of the under
side clearly accords with that of Horsfieldi.

The larva feeds on rouinpout paliit (Paspalum conjugatum Berg) and on
the grass known as giyufyngan (Cynodon Dactylon Pers.). The young cater-
pillar is greenish, but subsequently turns to a faint loam colour or yellowish
white, almost colourless. It has a dark dorsal line and a light band along the
ventral boundary, sometimes with more or less longitudinal lines on the head
which, however, are much more distinct in some specimens than in others.
The tail part terminates in two projections which, like the body, are closely
invested with very short black or white hairs which can only be seen distinctly
with a strong lens. The head is round but with two short projections. The
chrysalis is suspended from a grass stem. A pupa of 3'''^ February produced
the imago on the ii''' of the same month while from one of 25''^ March the
perfect insect emerged on y'^ April. The figures given by Moore of the cater-
pillar and chrysalis are very indistinct.

5. Fasciata Hew. (PI. XVI, fig. 39)-

Hewitson, Trans. Ent. Soc. of London, Ser. 3, 2,/. 287 (1864). Ypthima Fasciata.
Distant, Rkop. Mai., /aiJ. 420, ^^. 122 (1882 — 1886). .
Elwes and Edwards, Trans. Ent. Soc. of London, 1893,/. 44. „ »

Fruhstoreer, (Seitz, Grossschm. d. Erdc), p. 287, //. 99^

(191 1) » »

Only met with in the prov. Besouky in E. J.

6. Pandocus Moore. (PI. XVI, fig. 40^:, 40*^, \oc, 40^^, 40^, 40/).

Moore, Cat. Lep. M. E. I. C, I,/. 235, A^o. 506 (1857). Ypthima Pandocus.
Hewitson, Trans. Ent. Soc. of London, Ser, 3, 2, p. 290,

//. iZ, /ig. 12 (1864) » I.

Butler, Tratis. Ent. Soc. of London, 1879,/. 537 • • » Corticaria.

53

Distant, RIiop. Mai, p. 55, tab. 6, yfo. 8(1882 — 1886). Ypthima Corticaria.

„ „ „ /. 4 1 9 Ypthima Pandocus var Corticaria.

Staudinger, Iris, II, /. 38 (i88g) Ypthima Pandocus.

Elwes and Edwards, Trans. Ent. Soc. ol London, 1893,

/. 22, pi. 2, fig. i^ ,

Fruhstorfer, (Seitz, Grossschm. d. Erdc), p. 293, //. gg/'

(191 1) .. »

W. J. Buitenzorg (165); Salak mountains (780); Tjipanas (11 14); Gedeh
mountains (1480); mountain chain in the prov. Prayangan (+ 1700); mountains
in the prov. Bantam. Vicinity of the Pelabouan Ratou or Wijnkoops Bay on
the south coast (+ 1 50).

C. J. Salatiga (580) {Jacobson).

E. J. Semarou mountains (740); Tosari (1 777) ; Tengger mountains (i 200)
{Jacobson) ; BUtar in the prov. Kediri ; prov. Banyouwangy.

I possess many specimens of this species, not from the low lying country
of northern Java but from the mountains, and likewise from the lower districts
near Wijnkoops Bay on the south coast. They comprise light and dark
coloured, large and small, with more or less pronounced ocelli, all having been
caught either at the height of the dry or during the wet season, so that I find it
impossible to trace in these differences in colour development any relation either
the place or time of capture. Only the light coloured $ specimens from the
Prayangan mountains appear to me to be the largest with the most strongly
developed ocelli. I can, therefore, see nothing but individual differences of
colour evolution in the various races of this species established by Fruhstorfer,
any more than in the species Nigricans proposed by Snellen and Fruhstorfer.
In the numerous specimens of the latter from E. and W. Java, in my possession,
mostly taken during the dry but also in the wet season, I can only recognize
smaller, less developed individuals of Pandocus ; not a single definite feature
characterizing either form can be traced since in large series they may, distinctly
be seen to run into each other. Breeding may ultimately settle this matter.

This butterfly is very common in the mountains flying low near the ground
amongst the underwood, even when there is no sun or during rain. Some
9 attain an expanse of wing of 45 mm. The larva feeds on a//ang a//ang
(Lmperata arundinacea Cyrill), bamboo (Bambusa sp.), and roumpoid pah it
(Paspalum conjugatum Berg). The colour dimorphism, referred to in the
introduction, is met with in these caterpillars, since there are loam- or brick-
coloured as well as light green individuals. They exhibit a dark dorsal stripe

54

and on either side some regular dark dots ; a light line subdorsally and on the
ventral boundary a raised very light dirty white band — in green individuals
it is yellow — above which occur two dark longitudinal lines, al covered with
very short stiff hairs. The body terminates posteriorly in two short projections
and two very short projecting points occur on the head.

The pupa, which is beautifully marbled dark and light brown, is one of the
suspensory kinds.

Genus NEORINA Westw.

I. Crishna Westw. (PI. XVI, fig. 41).

Westwood {Doubl. and Hav. Gen. D. Lep., p. 361) 185 1. Cyllo Chrishna.
Fruhstorfer, Entom. Nachr., 14, /. 337 9 (1893). . . Neorina
„ (Seitz, Grossschm. d. Erde), p. 327, />/. 94a?
(191 1)

W. J. Vicinity of the Pelabouan Ratou or Wijnkoops Bay and Tjiletou or
Sand Bay along the south coast (+ 1 50).

C J. ?

E. J. Semarou mountains (740) and Southern mountains (500).

This butterfly is evidently not rare at the localities enumerated; in the
Southern mountains I found it at rest on the ground and making its way to
hide amongst the tall allano allang grass, exactly in the manner as practised
by Cyllo Leda L., from which I concluded that, like the latter, it flies about
the twilight. I never encountered it, however, in the vicinity of Batavia,
Buitenzorg, Touban, Bodjonegoro, or Kediri, neither have I ever seen specimens
captured in any other districts in Java but those named. On one occasion I
found at Tjibodas in the mountains of the Prov. Prayangan (14 10) on rotian
(Calamus sp.) a caterpillar of the Debts type, but 6.5 cm. in length and of
proportionate bigness. It was light green ; the head somewhat lighter in colour
and having the appearance of being covered all over with white dots, caused
by very short stiff white hairs. It had a dorsal and subdorsal yellow line, the
latter having here and there a pale dot, while on the ventrae boundary occurred
an indistinct yellowish supra-stigmatic and a very distinct brownish-yellow infra-
stigmatic line. The head was furnished with two relatively short projections
which were white spotted with brown on the anterior side and rather yellowish

55

at the base. The hinder portion also terminated in two projections of moderate
length. The stigmata were reddish brown. This caterpillar turned into a suspen-
sory chrysalis of a dirty green sparsely spotted with brown, which, as regards
the general form and the projections on the head, fairly agreed with a Debis
pupa but exhibited two rows of subdorsal projecting, points on the abdomen.
On leaving the mountains, I was obliged to take this pupa with me to Buiten-
zorg but, as frequently happens with pupae carried from the cooler humid
mountain districts to low lying country, it died there. In view of the size of
the caterpillar and its relationship with species of Debts, it can hardly have
been anything but one of N. Chrisna Westw.

Mr. Fruhstorfer is of opinion that the specimens from E. J. and those
from W. J. are distinct, and couples with this view some far-reaching specula-
tions concerning former land connections. I only possess a single specimen
from E. J. but I fail to discover the characters which he indicates as differ-
entiating it from the W. J. specimens — of which I possess many. I am,
therefore, constrained to doubt the justness of his arguments. Moreover, the
same stage in the process of colour evolution may well have been reached by
butterflies of one and the same species occurring in different regions, without
basing theories as to the former existence of land connections on this single
fact, uncorroborated by other evidence.

It appears to me very probable that Neorina Crishna Westw. and its
allies in other regions of the Malayo- Australian fauna are remnants of an ancient
genus, formerly containing many types but now dying out and consequently
represented by few forms. The great size of the present species, as compared
with other closely allied Satyridae, may also have some bearing on the question.
For instance, I possess as pecimen from the Wijnkoops Bay in W. J. with a broad
white transverse band which is particularly large. The genus Hestia probably
owes its large size to the same cause and, amongst Papilios, Ornithopteras
exhibit the same phenomenon. The close association between Neorina and
the tertiary genus Neorinopsis Butl, so aptly referred to in this connection by
Fruhstorfer, also points in the same direction.

Genus DEBIS Westw., Herr. Sch.

The species of Debis appear in general to be mountain butterflies. I have
found only one species, D. Europa F., on a few occasious in low lying country.
Dr. Hagen has had a similar experience with respect to the species of this

i6

56

genus in Sumatra, but according to Dr. Martin, besides D. Europa F.,
D. Manthara Felder (Mekara Butl.) is said to occur also in low lying districts
in Sumatra. The individuals from Java of species occurring also in British India
frequently differ considerably from the latter in the shade of colour, at least in
so far as can be judged from the figures given by Moore. The so-called
seasonal differences, illustrated by Moore, I have been unable to discover in
Java. I must, however, admit that since the majority of Debis species are
fairly rare in Java no large series of these are available for comparison, except
as regards D. Europa ¥. and D. Rohria F., but of these Moore also knows
no seasonal varieties. To judge from Moore's figures two types of larvae appear
to occur among the various species but I have met with only one of them
in Java.

Fruhstorfer considers the species of Debts, except D. Rohria F., to be
twilight butterflies ; as a matter of fact they are never seen on the wing in
day-time except when disturbed.

I. Minerva F. (PI. XVI, fig. 42a, 426).

Fabricius, Etit Sys/., /. 493, No. 216 (1775) .... Papilio Minerva.

Cramer, II, /. 29, //. 116, D. E. (1779) , Arcadia.

Distant, Rhop. Mai., /. 414, tab. 2>^, fig. 8 (1882 — 1886). Lethe Minerva.

Moore, Lep. Ind., I, /. 237, //. 11, fig. i, \a—b (1892). Nemetis
Bingham, Fauna Brit. India, vol. i, p. 90,//. \i,fig. 14 9

(1905) Lethe

Fruhstorfer, (Seitz, Grosssc/im. d. Erde, p. 318,//. 98a

(1911) ' "

W. J. Buitenzorg (265); prov. Prayangan (500) ; vicinity of the Tjiletou or
Sand Bay on the south coast (+150).

C. J. Prov. Jokjokarta {jfacobson).

E. J. Banyouwangy and also without precise indication of habitat.

Common in the lower lying districts. {Fndisf.).

The accompanying figures of the present species are good, only in many 9
the colour of the upper side is much more dull yellowish-grey than is shown
in the illustration. The <S figured by Moore and described by Bingham does
not agree with the Java specimens nor quite with those figured cis from Java
by Fruhstorfer.

The caterpillar feeds on bamboo leaves (Bambusa sp.). It is of slender
build, with two long processes on the head, the hinder part of the body also

57

terminating in two long projections, the latter being practically contiguous. In
colour it is sometimes yellowish green with a double yellow dorsal line, between
which sometimes are found a few yellow and red-brown dots and two yellow
subdorsal lines which are continued on the lower part of the processes on the
head, together with a yellow line on the ventral boundary. The remainder of
the processes on the head is of a very light purple with black extremity, the
apex being slightly yellowish. Over the whole of the body minute white or
yellow dots may be observed which are the extremities of very small hairs.
Some caterpillars, however, are of a dark ochreous yellow or stone colour with
black markings ; in these the processes of the head are also stone coloured with
black extremity.

The pupa is without processes, sometimes of a handsome green at other
times very faintly loam coloured. A pupa of 2y^ April produced the imago
on y^ May.

2. Chandica Moore. (PI. XVI, fig. 43a;, ^^d).

Moore, Ca/. Le/>., M. E. I. C, I,/. 219,^/^.455(1857). Debis Chandica.
Lep. Ind., I, /. 247, //. 1<^, fig. 2, 2a, b (1892) .

Bingham, Fauna Brit. India, I, /. 94 (1905) Lethe „

Fruhstorfer, (Seitz, Grossschm. d. Erde), p. 320, //. 98,
<5 and rt' (1911)

W. J. Prov. Prayangan (+ 1 500).
C. J. ?
E. J. ?

The specimens from Java, of which good figures are now added, differ
considerably, especially as regards the 9, from Moore's figures as well as from
those given by Fruhstorfer of Indian specimens.

3. Samio Dbd. (PI. XVI, fig? 44).

DouBLEDAY, Geu. D. L., p. 360, //. 61, //V. 3 (185 1) . . Debis Samio. •

Felder, Wien, Ent. Mon. Ill, /. 400 (1859) Purana.

Moore, .Lcp. Ind., I, />. 242, //. 78,/^. 2, 2a (1892) . . „ Samio.

Fruhstorfer, (Seitz, Grossschm. d. Erdi), /. 319 (igii). . Lethe „

W. J. Prov. Prayangan (+ 1 500).
C.J. ?

E.J. ?

5«

The present species appears to be known only from Java, where it is
apparently not common. According to Fruhstorfer the 9, which is unknown
to me, has also white transverse lines on the upper side of the fore-wings.

4. Manthara Feld. (PI. XVI, fig. 45^', 45*5, 45^, 45^').

Felder, Novara, Lep., />. 497, No. 861 (1867) .... Debis Manthara.
BuiLER, Anil. Mag. Nat. Hist., so: III, 20, /. 403, //. 9,

fig. 9 c? (1867) Lethe Mekara.

Moore, Lep. Ind., I, />. 244, //. 79, ^V. i, i^, b (1892) . Debis „

Bingham, Fauna Brit. Ind., I /. 95 (1905) Lethe „

Fruhstorfer, (Sehz, Grossschm. d. Erde), p. 320, //. 98*?

(191 1) „ Manthara.

W. J. Buitenzorg (265); Salak mountains (780) ; Megamendoung mountains
(1480) ; prov. Prayangan (+ 1500) ; vicinity of the Pelabouan Ratou or Wijnkoops
Bay (+ 150).

C. J. ?

E. J. Banyouwangy.

The orange colour on the upper side of the hind-wings of the c? is frequently
faded to a dirty yellow and in some specimens the white transverse bands on the
upper side of the fore-wings of the 9 are very indistinct or may be entirely wanting.

From Java I possess two d whose underside is much darker than in the
other specimens and evidently differing from them. Otherwise the individuals
from Java do not deviate much from the Indian specimens figured by Moore.

The larva, which lives on young bamboo leaves, in appearance much
resembles that of D. Minerva F. It is grass-green in colour, covered with
many white or yellowish dots, with a white band at the ventral margin and
two or even four yellow dorsal lines between which, in the full-grown larva,
yellow markings pas.sing into brown, occur. The head is furnished with two
long projections, yellow at the base, passing into rose and with a black apex.
They are very dark in colour on the under side. The hinder portion of the
body also terminates in two long granulated projections which lie closely
appressed or with their apices crossed. On one occasion I found a loam-
coloured caterpillar which, however, sickened and ultimately died. From a
pupa of 2^'"^ April the imago emerged on May 4''', one of 26''^ April produced
the perfect insect on May s'"", while another of 5"" May completed is meta-
morphosis on the 1 2''' of the same month.

59

5. Darena Feld. (PI. XVII, fig. 46^, a,bb).

Felder, Novara, Lep., /. 498, No. 862, //. 68, jig. 4, 5

/i85^N , . • Debis Darena.

Staudinger, Iris, ix, />. 228 (1897). Lethe Darena var. Borneensis, Sumatrensis.

Fruhstorfer, (Seitz, Grosssclim. d. Erde), /. 322, //. 98^

(igii) Lethe Darena.

W. J. Megamendoung, Wayang, and Gedeh mountains (1500).
C.J. ?
E. J. ?

6. Dyrta Feld. (PI. XVII, fig. 4 7«. Mh

Felder, Novara, Lep., p. 497, No. 860 (1867) Debis Dyrta.

Moore, Lep. hid., I, p. 259, //. ^2, fig. 3, Za—c (1892). . Lethe

Staudinger, Exot. ScJim., /. 221, /•/. 78 d* (1884 — 1888)

Bingham, Fmma Brit. India, \, p. 80 (1905) Rohria.

Fruhstorfer, (Seitz, Grossschn. d. Erde), p. 316, //. q^b

(1911) Dyrta.

Bingham considers this species to be the Papilio Rohria of Fabricius,
but Snellen thinks otherwise.

The (3 from Java, now figured, differs considerably from the Indian speci-
mens as figured by Moore as well as from the specimen from Lombok figured
by Fruhsforfer.

W. J. ?

C.J. ?

E. J. Fouspa, Tengger mountains (630) ; Tengger Mountains (i 200) {Jacobson).

Fruhstorfer also obtained this species only from E. J.

7. EuROPA F. (PI. XVII, fig. 48^, ^U).

Fabricius, Syst. Ent., p. 500, A^o. 247 (1775) Papilio Europa.

Cramer, I, p. 125, pi. 79, C D. {\liq) Beroe.

IV, /. 50,//. 313. E. F. (1782) , Arete.

HiiBNER, Samm/. Exot. Sclivi. (1806— 1827) Oreas Europa.

6o

Distant, Rhop. Mai., />. 43, tab. i, fig- 5, 6 (1882—1886). Lethe Europa.
Staudinger, Exot. Schm., />. 221, //. 78 (1884 — 1888) . . „ „

Moore, Lep. hid., I, />. 254, //. ^2, Jig. i, \a (1892) . . „

Bingham, Fauna Brit. India, I, />. 77 (1905) , „

Fruhstorfer, (Seitz, Giosssc/im. d. Erde), p. 315, //. 96^'

( 1 9 1 1 ) »

W. J. Batavia (14); mountains in the prov. Bantam; Sindanglaya (1074) ;
Vicinity of the Pelabouan Ratou or Wijnkoops Bay and the Tjiletou or Sand
Bay on the south coast (+ 150).

C. J. Touban on the north coast (6); prov. Tegal; prov. Madioun.

E. J. Pouspa, Tengger mountains (630) ; Semarou mountains (740) ; prov.
Kediri ; Madoura.

The accompanying figures of this species also do not agree with those of
Moore especially as regards the 9.

Bingham, on the authority of Davidson and Aitken, describes the cater-
pillar as green, paler beneath, fusiform; head with a single short erect horn;
body suddenly attenuated from the 1 1 '^ segment ; the pupa is stated by the
same author to be uniform pale green, stout, smooth, quite regular, except the
head-case which is semi-detached, broad and angular, with two sharp points in
front. Both larva and pupa appear to differ considerably, especially as regards
the processes, from the species of the genus Debis observed by me, but they
agree with the figure of Lethe Drypetes Hew. given by Moore.

8. Rohria F. (PL XVn, fig. 49).

Fabricius, Majit. Ins., II, /. 45, iVb. 446 (i 787) . . . . Papilio Rohria.
KoLLAR in von Hugel's Kaschmir, iv, 2, /. 448, //. 16,

jig. 3, 4 (1844) Satyrus Isana.

Moore, Lep. Lid., i, /. 264, p/. 84, fig. 2, 2a— c (1892) . Lethe Rohria.
Bingham, Fauna Brit. India, I, /. 80 (1905). Lethe Rohria race Nilgeriensis.
Fruhstorfer, (Seitz, Grossschm. d. Erde), /. 315, />/. 97a

(191 1) Lethe Rohria Godana.

W. J. Buitenzorg (265); Sindanglaya (1074); Tjibodas (14 10); Mega
Mendoung mountains (1480); mountains in the prov. Prayangan (1500) ; Salak
mountains (780).

C. J. Oungaran, prov. Semarang (800) {yacobson).

E. J. Tengger mountains (1000); Semarou mountains (740); Blitar.

6i

As shown in the accompanying illustration the colour of Java individuals of
this species is considerably darker on either side than that represented in the
figures of Moore and Fruhstorfer.

The species, although a mountain butterfly, occurs lower, as appears from
the above records, than Fruhstorfer thinks, for he states that it is never met
with below 1200 metres altitude. I can see no difference between specimens
from West and East Java, although Fruhstorfer distinguishes a dry season
E. J. form which he calls Mangala.

Genus ORSOTRIAENA Wallengren.

I. Medus F. (PI. XVII, fig. 50^, sod).

Fabricius, Sys/. Ent., p. 488, No. 198 (1775). . . Papilio Medus.
Cramer, I,/. 816, //. 1 1, C Z?. (i 779) . . . . „ Hesione.

IV, /. 341,//. 362, C (1782) „ Doris.

Wallengren, Kongl. Vet. Akad. Forhandl., 15, /. 79

(1S58) Orsotriaena Hesione.

Distant, Rhop. J/a/., /. 49, /ff<^. 4,/^. 8 (1882 — 1886). Mycalesis Medus.
Staudinger, Exot. Schtn., p. 230, //. 82, fig. 5

(1884 — 1888)

Moore, Lep. hid., I, /. it^, fig. i, la, b, 2 (1892). Orsotriaena „

Martin, Iris, viii, /. 239 (1895)

Bingham, Fauna Brit. India, I, /. 69, fig. 19(1905). „ Meda.

Fruhstorfer, (Seitz, Grossschm. d. Erde), p. i^B.pl.qia

(191 0 „ Medus.

W. J. Batavia (3—14); D^pok (95); Tjampea (160); Buitenzorg (265);
Salak mountains (780); mountains in the provinces of Bantam and Prayangan
(1500); vicinity of the Pelabouan Ratou or Wijnkoops Bay and the Tjiletou or
Sand Bay on the south coast (+ 150).

C. J. Prov. Semarang (300) {Jacobson; Magelang (500); prov. Jokjo-
karta {Jacobson).

E. J. Malang (443); Jember (98) in the prov. Besouky; Banyoupoutih (45)
in the prov. Probolingo ; Madoura.

This species is very common in Java ; amongst the many specimens in my
possession there are, as in the case with many species of Mycalesis, some

62

which are dark on both sides and others which are light brown ; generally
the d* are, indeed, darker, but otherwise I am unable to detect any difference
according to period of capture or habitat. Specimens either with large or
with less developed ocelli are met with in both sexes; amongst these occur
individuals with strongly developed ocelli taken at the height of the dry season
and others with least developed ocelli caught in the wet season. Specimens
such as Moore figures as the dry-season form I have never found in Java.

The caterpillar was repeatedly found on a//ang-a//ang (Imperata Arundi-
NACEA Cyrill.) and according to de NiciviLLE occurs on />adi (Oryza sativa L.).
Fruhstorfer informs us that Martin found the greenish-white globular eggs
on the under side of leaves of various grass species ; he also relates the whole
life histor}^ of the caterpillar according to Martin's observations.

I found them to be of a type such as occurs in some Debts larvae, with
long processes on the head and the hinder portion, the latter being sometimes
closely appressed. Some are greyish-green with a dark dorsal line, a faint
jellow subdorsal line and a clear white band at the ventral margin. With the
aid of a lens it is seen to be covered with short black hairs. The head
resembles ground glass and the horns are red, darker at the base and under
a lens it is also observed to be covered with short stiff hairs. The posterior
processes are faint rose. Another caterpillar was white with a raised white
stripe on the ventral margin, the horns and posterior processes rose. Yet
another caterpillar W'as entirely of a beautiful rose colour, including the horns
and posterior processes, with the stripes already alluded to. This caterpillar
had the head like ground glass but darker. The pupa is attached to a grass
stem and is light brown with longitudinal dark brown stripes ; the head termi-
nates in two sharp points mostly closely appressed in the same manner as is
the case with the posterior processes. According to Martin the imago emerged
after nine days in the fore-noon.

The figures given by Moore of the larva and pupa of O. Man data Moore
from British India and Ceylon closely agree with mine of O. Medus F.

Genus MYCALESIS Hb.

As is the case in the genus Cyllo, in the present genus a universal darkening
of colour appears to occur which subsequently again gradually diminishes.
Vestiges of the original red are, in fact, present in some species. The general
colour in several species as a reddish or brownish grey which in some indi

63

viduals has become very dark, almost black; it cannot be stated definitely
whether this difference is due to the increase of black or to the subsequent
fading process. It is, however, clear that we have to deal here, not with the
result of directly operating influences, but with an evolutional mutation process;
in low lying districts as much as in mountainous regions, in the wet season as
well as at the height of the dry season, light and dark coloured specimens
are equally met with.

Specific distinctions, based purely on some difference in form of wing, as
adopted by Bingham, I am unable to admit; Cyllo Leda L., for instance,
clearly teaches us that when a species is in a state of active evolutional change
this manifests itself in the form of wing as much as in the colouring; for the
same reason the form of wing in some species, such as Danais Agleoides
Felder, differs in the two sexes /. e. when they have reached different stao-es
m the evolution of form. As regards the specific differences which Bingham
and Fruhstorfer infer from the secondary sexual characters in males, I am
not inclined to attach much value to this either. Moreover, Bingham is himself
still doubtful in this respect, while Fruhstorfer observes that these characters
are extremely variable in the genus Debts, even in the seasonal forms of the
same species and the forms which he regards as subspecies.

The form or colour of such secondary characters cannot be considered of
the same weight as attaches to the sexual organs proper. Much accurate
investigation of ample material from different districts and especially much
pedigree breeding are, indeed, required before the acceptance of .specific
differences on these grounds may be considered justifiable.

I. Nala Felder. (PI. XVII, fig. 51^, 51^, 51^, 51^^).

Feeder, Wien, Ent. Mon., p. 403, No. 46 (1859) . . . Mycalesis Nala.
Novara, Le/>., p. 500, No. 868, //. 57, fig. 10

^'867) ," . . „ Sudra.

Fruhstorfer, (Seitz, Grosssclwi. d. Erdc), pi. 936*5 (191 1).

W. J. Tjibodas (14 10); Megamendoung mountains (1480); Salak mountains
(780); mountains in the prov. Prayangan (1500); mount Tjerymay, prov.
Cheribon.

C. J. Oungaran (800) {Jacobson); Magelang (500).

E. J. Tosari (1777); prov. Banyouwangy, Tengger mountains (1200)
iyjacobson).

Feeder, who discovered the species, subsequently described a variety from

17

64

West Java specimens which merely differ in the darlt ochreous yellow margin
on the under side of the wings and which he called Sudra. Fruhstorfer,
however, is of opinion that two species have been mixed up by Felder and
he distinguishes these as Sudra and Nala, but bases them on alleged characters
which in no wise accord with the differentiating characters given by Felder
and to which in any case the latter's names have unjustly been applied, giving
rise to confusion. Fruhstorfer includes in his Sudra the form with the dark
ochreous yellow on the under side, as well as the other form described by
Felder; he designates as Nala some c? individuals found by him particularly
in E. J. which differ principally in having a black shining scent spot on the
upper side of the forevvings, while in addition several small faint ocelli occur
on the under side in the margin of the fore-wings, whereas in Sudra only
two such ocelli are present which, however, are larger. Females in which
several small faint ocelli also occur on the under side of the fore-wings, he
regards as the other sex of this species. Such specimens do, indeed, occur
and the difference in the ocelli appears to be fairly constant ; this character
is, nevertheless, not of great value. As regards the development of the ocelli
in the margin of the under side of the wings, considerable individual difference
exists in many species of Mycalesis ; and this is the case in the present one —
I here add an illustration (PI. XVII, fig. ^06) of a specimen from the Tengger
mountains in E. J., in which these ocelli are considerably larger than usual —
and in these circumstances I consider it too bold to assume specific distinction
merely on account of such difference in the ocelli. The presence or absence
of the forementioned scent spot is in quite another category and if this distinction
were sufficiently confirmed I would not hesitate to admit specific difference on
this ground. But this is not the case. Mr. Fruhstorfer was good enough
to let me have such a c? specimen and Mr. R. van Eecke, Assistant — princi-
pally concerned with Lepidoptera — at the Natural History Museum of Leiden,
who is an expert microscopist besides, had the kindne.ss to examine this black
spot for me. He found, in fact, some scent scales distributed in these spots
but they were likewise observed on the upper side of the forewings in the other
Nala males which are without the black spot, so that the scent scales have
actually no connection with the said black spot which, therefore, is not entitled
to the term scent spot. It appears to consist principally of the ordinary, but very
dark pigmented, scales, agreeing in this respect with similar black spots, as occur
for instance in some specimens of C. Leda L. (c/'. pi. XV, fig. 300), and these are
simply relics of a former particularly intense development of the black pigment at
the place where only after a long period this has spread over an obstinate spot.

65

M. Nala Felder is very common in damp forests of the higher mountain
ranges of Java. A comparison of about lOO specimens, in my possession and
in the L.eiden Museum, not a single one of which shows the black spot referred
to, has convinced me that in general the individuals of both sexes from E. J. —
both of the dry and the wet season — as well as those from C. J., are darker
in colour but I possess one, taken during the wet season in E. J., which is
of a lighter colour. The specimens from W. J. are mostly of a brown shade,
both in the dry and the wet season, the 9 being lightest in colour. Never-
theless, I also possess several c?, of the dry as well as the wet season, from
W. J., whose upper side only is dark. Many specimens, but exclusively those
from W. J. have the border on the under side of a beautiful dark ochreous
yellow (Sudra), mostly of the wet season ; of five specimens taken at Buiten-
zoro- in July — in the dry season, therefore — this margin in only one is of
that colour. In the 9 especially this border on the under side is sometimes
very white and generally various transitions occur between this white and the
deepest ochreous yellow but, as already stated, only in W. J. Amongst these
some 9 are found (PI. XVII, fig. 51^) in which the ocelli, especially on the
under side of the fore-wings, have practically disappeared, agreeing, therefore,
with those which, according to Fruhstorfer, are supposed to be $ of his Nala.
The ochreous yellow in the border on the under side must probably be also
looked upon as a relic of the original red colour ; the brown in most specimens
from E. J. still contains much red.

3. MooREi Felder. (PI. XVII, fig. 52).

Felder, Navara, Lep., p. 502, No. 870,//. 67, //^. 9 (1867). Mycalesis Moorei.
Fruhstorfer, (Seitz, Grosssc/nn. d. Erde), p. iA9^ f^- 9 2^

(191 1) " "

W. J. Tjampea (160); mountains in the district of Buitenzorg; Salak
mountains (780); mountains in the prov. Bantam.

C.J. ?

E. J. Semarou mountains (740); Tengger mountains (1290) {Jacobson).

Like the two preceding, this butterfly is only met with in the forest of the
higher mountain ranges where, however, it is less common than M. Nala
Felder. The figure now given agrees with all the specimens in my possession.
That occurring in Seitz, Grossschm. d. Erde, evidently represents a specimen
with much reduced ocelli as they occur, according to Fruhstorfer, in the dry

66

season. The two specimens I possess from the mountain range in the prov.
Bantam, were, however, taken in September, therefore, at the height of the
dry season.

4. MlNEUS L.

Linn., Sysl. Nat. Ed. XII, I, 2 /. 768, No. 126 (1767)
BuiLER, Cat. Sat. Brit. Miis., p. 1 35, //. 3,//^. 12(1868)
Moore, Lep. of Ceylon, I, p. 22, pi. 11, fig. 4, 4a, b (i 880)
Distant, Rhop. MaL, p. 50, tab. ^, Jig.- 13, 14, 7 {var.)

(1882 — 1886) ,

Staud., Exot. ScJim., p. 230, //. 82 (1884, 1888) .
DE NicEviLLE, Joum. Asiat. Sac. of Bengal, LV, /. 235

//. 12, fig. 3 (1886)

Moore, Lep. Ind., I, />. 183, //. bo, fig. la — /(1892)

Martin, Iris, VIII, /. 241 (1895)

Bingham, Eauna of Brit, hidia, I, /. 58, fig. 8, g (1905)
Fruhstorfer, (Seitz, Grosssehm. d. Erde), p. 343

Papilio Mineus.
Mycalesis Judinella.
„ Mineus.

Calysisma
Mycalesis

Of this kind, which in Java is almost entirely replaced by the next species,
I received, however, a typical 6 taken by Mrs. Lechner at + 1000 metres in
the province of Prayangan in W. J.

5. HoRSFiELDi Moore. (PI. XVII, fig. 53a, 53<5, 53^, 53^^).

Moore, Lep. Lid., I, /. 197, pi. 66, fig. 2, 2a — ^(1892). Calysisma Horsfieldi.

Martin, Iris, VIII, /. 242 (1895) Mycalesis (Cal.)

DE Niceville and Martin, Joum. Asiat. Soc. Bengal, 64,

A 379 (1895) Mycalesis

Fruhstorfer, (Seitz, Grosssclnn. d. Erde), p. 345 . . „ „

W. J. Batavia (3—14); Depok (95); Buitenzorg (275); mountains in the
prov. Bantam ; vicinity of the Pclabouan Ratou or Wijnkoops Bay on the south
coast (+ 150).

C. J. Semarang (6) ; Jokjokarta and Nousa Kambangan {Jacobson).

E. J. Semarou mountains (740); prov. Banyouwangy; mountains in the
south of the prov. Passourouan [Eru/istorfer) ; Jember (98) in the prov. Besouky.

The numerous specimens collected by me in Java were erroneously indenti-

67

fied as MiNEUs L. by Snellen, but after consulting Mr. Fruhstorfer and his
collection I am convinced that they must be referred to Moore's later species
HoRSFiELDL The clear white patch of specialized scales, duly observed by
Snellen, but incorrectly termed ochreous yellow by Moore, near the costa
on the upper side of the hind-wings in the d, distinctly separates this species
from Mineus L. In Mineus such a patch is indeed present but is covered
by pencils of long hairs, while in Horsfieldi also a similar patch occurs
which, however, is larger and more distinct and is placed at some distance in
front of these pencils of hairs. The figures given by Moore are, however,
poor and do not represent the species well ; the seasonal forms figured by him
are not to be traced, at least in Java. The d are generally darker in colour
than the 9, but there are some lighter ones also. Difference in colour or size
according to the dry or wet season is on the whole not to be observed but
considerable difference in the stage of development of the ocelli on the under
side does occur. Those most developed I find, the same as in M. Perseus F,,
in specimens captured in April near Wijnkoops Bay; moreover, these ocelli
are generally better developed during the wet than in the dry season ; I possess
several specimens with specially small ocelli, all taken during the dry season.
Amongst these even considerable difference in the degree of development exists
but in those least developed the ocelli are never reduced to white dots, and
I have individuals taken at the height of the dry season, both in the mountains
and in the lowlands, whose ocelli are equally developed as is usually the case
during the wet season. The individual unequal advance in colour development,
already referred to under Cyllo Leda L., therefore, exists here also, but a
separation into seasonal races I am unable to recognize. On the upper side
of the hind-wings of the 9 sometimes one and sometimes two ocelli are visible
or they may even be absent altogether.

I have observed the larva on a//a/io allang (Imperata arundinacea Cyrill.)
and on a bamboo like grass. Some larvae are dark green with a pale green
lateral line, while in addition a dorsal and several dark and light longitudinal
lines are to be seen. The body terminates in two short, spinous, faint red
points. The head is reddish brown and furnished with two short stout points.
Another larva was reddish-brown with a faint dark dorsal line and a lateral
band, between which similar bands ran obliquely backwards; it was also strongly
chagreened. Yet another larva, likewise strongly chagreened, was light greyish
brown with a light line on the ventral margin and indistinct dark brown dorsal
bands, and with faint marking, resembling a linked row of rhombs, at the sides.

The reddish brown larva before pupating turned to pale green. The pale
green pupa has yellowish white stigmata and is devoid of processes. It was

68

firmly attached by the tail to the box in which it was contained and, like the
pupa of Orsotriaena Medus F., is suspended horizontally. A pupa of
April 2 2'"' produced the imago on April 30"*.

5. Perseus F. (PI. XVII, fig. 54a, 54*^, 54^).

Fabricius, Sys/. Ent., />. 488, No. i99 (i775)- • • • Papilio Perseus.

„ Ent. SysL, Suppl. /. 426 (1778) „ Blasius.

Donovan, Ins. N'ew. Holland, pi. 2b, fig. 3 (1805) . „ Perseus.

Hewitson, Exof. Buttcrf., iv, Mycal., fig. 35 (1804) . . Mycalesis Lalassis.

Distant, Rliop. Mai, />. 52, tab. 7, lig. 7 (1882 — 1886). „ Blasius.
Moore-, Lep. Ind., I,/>. 1 74,//. h^,fig- i, la—d, fig. 2, 2a— c

(1892) Mycalesis Calysisma Perseus.

Bingham, Fauna Brit. India, I, /. 57 (1905) . „ >,
Fruhstorfer, (Seitz, Grossschm. d. Erde),p. ^42

(191 1) ., 1.

W. J. Batavia (3 — 14); Depok (95); Tjampea (160); vicinity ofthePela-
bouan Ratou or Wijnkoops Bay on the south coast (+ 1 50).
C. J. Semarang (6).
E. J. Pouspa (630) ; Jember, prov. Besouky (98).

The species differs particularly, as Bingham quite correctly observes, from
its allies in Java — /. c. M. Horsfielpi Moore — by the peculiar bending
of three of the ocelli on the under side of the hind-wings. In the figure in
Seitz, die Grossschm. d. Erde, 92a this feature has been completely overlooked.
In my figure I have indicated it (PI. XVII, fig. 54a).

The species is very common in Java. As in the case of Horsfieldi
Moore, darker and lighter individuals occur, the d* being usually somewhat
darker but this has no reference to the seasons. The same applies to the
ocelli. Specimens with small ocelli occur, smaller in some than in others ;
these have been described by Fabricius as M. Blasius. In a specimen from
Tjampea I find the ocelli most reduced, being partly resolved into white dots.
I possess such specimens with small ocelli caught in may at Pouspa in E. J.
and in August — therefore, in the middle of the dry season — at Semarang,
but likewise some captured about the same time at Batavia and Semarang,
with well developed ocelli, and a 9 taken at Semarang in September where the
stage of development is, so to speak, intermediate. The development of the

69

ocelli I found most advanced, here as in Horsfieldi, in specimens caught in
April near Wijnkoops Bay.

I have also taken at Batavia on the same day in the rainy season two c?,
in one of which the eye-spot on the upper side of the fore-wings is very
distinct while in the other no trace is to be seen of it. Some of my other
specimens agree with the first and some with the second. As regards the row
of ocelli on the under side of the fore-wings I found amongst 43 specimens
many with only two, some with three or four, but only one with five ocelli
as in the accompanying figure. The small lower eye-spot is mostly wanting.
From this it will be seen to what extent the number of ocelli varies in the
same species — which is also the case with the European species of Satykus —
and that, consequently, there is no justification for basing forms or species on
the number of these. The attendant circumstances in this and the preceding
kind are doubtless indentical with those pointed out under Cyllo Leda L. ;
the rainy season indeed greatly promotes this development but only in the
individuals which have acquired susceptibility for it through evolution, while in
the case where the stage of evolutional change has progressed very considerably
this will manifest itself even in individuals making their appearance in the dry
season. Definite wet-and dry-season forms or races do not, therefore occur.
Much light is still required on the subject, not only as regards the genus
Mycalesis but in connection with Debts and Ypthima as well and we must await
further observation and breeding. Although I have numerous specimens at my
disposal, by far the great majority of them have been carefully labelled only as
regards the habitat but not as to the time of capture ; it was only during the
latter part of my stay in Java the importance of these data became manifest to me.
On one occasion I bred the imago from a larva living on allang allang
(Imperata arundinacea Cyrill).

6. Janardana Moore. (PI. XVIII, fig. 55).

Moore, Cat. Lep. M. E. I. C, I, /. 234, No. 502 (1857). Mycalesis Janardana.

Distant, Rhop. Mai., p. 54, /«<5. 5, //;«-. 2 (1882 — 1886).

Fruhstorfer, (Seitz, Grossschtn. d. Erde), p. t^/^i {iqii). „ „

W. J. Batavia (3 — 14) ; mountains in the prov. Bantam ; Depok (95) ; Buiten-
zorg (265); vicinity of the Tjiletou or Sand Bay on the south coast (+ 150).
C. J. Nousa Kambangan {Jacobson).
E. J. Prov. Banyouwangy.
Of this species I possess many specimens from W. Java but none exhibiting

70

great difference, and positively none of the little ocellated form as it occurs in
M. Perseus F. Both light and dark individuals are, however, met with in the
present species in the dry as well as in the wet season.

This butterfly is very common at Batavia and elsewhere in W. J., and
according to Fruiistorfer also in the mountains of E. J. The upper surface of
the fore-wings of the c? is black for the greatest part but it is incorrect to call
this a scent-spot for although some sent-scales are, indeed, to be found there,
as in the c? of M. Nala Felder, by far the greatest part of the scales in the
area referred to are simply ordinary scales coloured an intense black by a dark
pigment. The 9 are more yellowish brown but some resemble the c? in colour.
Wet or dry season makes no difference in this respect. Even in 9 taken at
the height of the wet season black dots replace the two ocelli of the under side
of the hind-wings. The form SAGnriFERA of Fruhstorfer is perhaps simply a
somewhat unusually developed 9. Fruhstorfer, on the authority of Dr. Martin,
describes the caterpillar which is said much to resemble that of M. Mineus L.,
and to be produced from similar greenish white eggs deposited singly. The
larva is supposed to be entirely nocturnal, during the day hiding itself deep
in the stems of grass, feeding only at might and on being touched drops,
feigning death. His description does not completely accord with mine. I
found the larva on roumpout benggala (Paspalum mollicomum Kunth). It
was of the same type as that of Ypthima Pandocus Moore, only slightly
larger, brown marbled with dark grey or loam-coloured with a dark dorsal
line and small blood-red spots ; stigmata black. The two processes on
the head and the posterior appendages as in Y. Pandocus. The pupa was
of a uniform brown. The perfect insect emerged on March 4''' from a pupa
of February 23"^"^.

Martin notes that this butterfly emerges, not like its allies between 8 and
10 o'clock in the forenoon, but only between 2 and 3 o'clock in the afternoon,
and does not begin to fly until twilight, so that he regards it as a twilight
butterfly, which I consider very probable from the fact that, like other
allied kinds, it is only seen on being disturbed by accident when it speedily settles
down again.

7. Fusca Feld. (PI. XVIII, fig. 56).

Felder, Wien, Enlom. Mon., iv, /. 401, A^o. 24 (i860). Dasyomma Fuscum.

Hewitson, Exot. Buttcrfl., iii, Mycal. fig. 26 (1862). . Mycalesis Diniche.

V, J/j'^fl/., y/^. 61, 62 (1874). .. Margites.

Distant, Rhop. Mai., /. '^i, tab. S,/ig. i (1882 — 1886). „ Fusca.

71

W. J. Tjampea (i6o); Buitenzorg (265); vicinity of the Pelabouan Ratou
or Wijnkoops Bay and the Tjiletou or Sand Bay on the south coast (+ 150).
C.J. ?
E. J. ?

Fourteen specimens of this species exhibit no difference except that the 9 are
much more strongly developed than the d".

8. Oroaxis Hew. (PI. XVIII, fig. 57).

Hewitson, Exot. Biitterfl., Ill, Mycal., fig. 38, 39(1864). Mycalesis Oroatis.

Distant, EntomologisI, 18, /. 259 (1885) „ Ustulata.

Rhop.Mal.,p.\\^,tab.\\,fio.\b{\^%2—\%Z(i).

W. J. Mountains in the district of Buitenzorg and mountains in the prov.
Prayangan.
C. J. ?
E. J. Mountains along the south coast. {FruhstorJ'er).

18

RAGADIDAE.

These butterflies are now usually considered as belonging to the Satyridae.

Genus RAGADIA Westw., Hew. Sch.

I. Crista Hb.

HiiBNER, Zutrdge, III, /. 2\, fig. 675, 676 (1825 — 1832). Euptychia Crisia.

HoRSFiELD, Cat. Lep. M. E. I. C, pi. '^,Jig. 9, 9^(1828).
Distant, /?/iop. Ala/., p. 420, /ad. ig,/!£: 7 {1882— 1886).
Fawcet, A?t?i. Mao. Nat. Hist., Scr. 6, 20,/. 11 i (1897).
Fruhstorfer, (Seitz, Grosschm. d. Erde), p. 261,//. 90^
(1811)

Hippaichia Makuta.
Ragadia Crisia.
„ Simplex.

„ Crisia.

W. J. Mountains in the prov. Prayangan; vicinity of the Pelabouan Ratou
or Wijnkoops Bay on the south coast (+ 1 50). At the latter place also the
form Simplex Fawcett.

C. J. Nousa Kambangan. {yacobsoii).

E. J. ?

I have received many specimens but all exclusively from the localities
mentioned, never from the northern parts of the island, Mr. Fruhstorfer
also appears to possess it from South Java only. According to Martin this
species is very common everywhere in Sumatra, even in the lowlands.

ELYMNIADAE.

Genus ELYMNIAS Hb. Heir. Sch.

The genus Elymnias, which is regarded as pertaining to the Satyridae by
most entomologists, exhibits in its larvae a great resemblance to those of Cyllo
not only as regards the slender build but particularly in respect of the peculiar
horns on the head and the spiny processes at the hinder end of the body.

Of E. Undulakis Drury and E. Lais F., with which I am best acquainted,
the former loves sunshine and is, therefore, principally met with in gardens
and other spots not densely covered with vegetation while the latter is to be
found by preference in plantations of Cocos nucifera L. which does not
generally occur in forests. It is, therefore, incorrect to call the members of
this genus in general forest butterflies, as is done by Wallace for instance.
All the larvae known in the genus likewise live on palm trees.

Whereas E. Undularis Drury, as has already been stated, seeks the
sunshine I have only once observed E. Lals F. flying about in the evening
twilight but otherwise, when disturbed, as frequently happens in the plantations
referred to, I have always seen it settle immediately on a tree trunk, in exactly
the same manner as Cyllo Leda L. It appears to me, therefore, that
E. Lais F. is likewise one whose members only fl}' in the twilight; this
may also be the case with other species of the genus. In the island of Saleyer
I noticed the same habit in the species which there replaces E. Lais F.
Dr. Martin, it is true, appears to have observed an individual of this species
flying in broad daylight, being engaged in the act of ovipositing, but under
stress of similar instinctive functions an insect may well deviate from its normal
habits, just as when disturbed it seeks safety in flight. I am, therefore, unable
to agree with Martin's assertion that Elymniads only fly in the full glare of
the sun. On the other hand 1 consider his suggestion, that Elymnias is an

74

ancient genus in process of dying out, more probable. It is, indeed, a fact
that several species of the genus are by no means common, some being even
very rare, while it is extremely doubtful that this should be due to the scarcity
of the food plant.

The pupae, as far as they are known to me, have not the form of those
of Cyllo Leda L., but rather resemble the chr)'salides of some species of
Debis, such as Debis manthara Felder.

The species of this genus furnish ample material for theorizing about
mimicr)'. As a matter of fact they follow the various lines of colour evolution —
as is the case with so many MalayoAustralian butterflies, and which, as has
already been explained above, finds its strongest expression in the genera
Euploea and Dana is — and in such a manner that whereas one species follows
on the lines of coloration in Euploeas others follow on those of Danaids. In
some even, such as E. Undulahis Drury, in a similar manner as occurs in
Euploea Midamus L., /. e. the c? following on the lines of coloration in Euploas
while the 9 moves in the direction of Danais where, as for instance in D. Genutia
Cram., a considerable amount of red has been retained. For, that it is a
question here of colour evolution is clearly demonstrated from a comparison
between the races E. Caudata Bull., in the d* of which much more of the
ancient Danais colour has been retained than in that of E. Undularis Drury
and E. Nigrescens Butl., both sexes of which have practically become entirely
obfuscated. For, as Martin justly assumes, these forms are nothing but races,
while climatic influences or the fantastic conception of insular melanism, any
more than former land connections, have nothing to do with this phenomenon.
It simply exhibits the various stages of transition as they arise from the uneven
advance of the evolutionary darkening process. Al these cases of colour
agreement are loosely accepted as instances of mimicry and the particulair
species thus imitated are indicated without hesitation, although unprejudiced
observation would clearly show that of such accurate copying there is never
any question and moreover, on account of the somewhat concealing habit of
most of the Elvmnia imagines, this mimetic protection would be of little use
to them.

I. Undularis Drury. (PI. XVIII, fig. 58^, 58<J).

Druky, ///. Exo/. Ell/., II, //. 10, //V. 2 (1773) . Papilio Undularis.

Cramer, II, />. 141, //. 189, E. G. (1779). . . „ Protogenia.

Ill, /. I 10, />/. 256, A. D. (1782) . . „ Undularis.

HiiBNER, Ztilmgc, I, /. 12, //. 37, 38 (181 8) . . Elymnias Jynx.

75

HoRSFiELD, Cat. Lif. M. E. I. C, //. 8, //V. 8a, b

(1828) Melanites Undularis.

HoRSFiELD and Moore, Ibid., I, /. 237, //. 6,

fig. 7, la (1857)

Staudinger, Exot. Sclnn., /. 237, //. 86 (1884 —

1888) Elymnias „

Moore, Lep. Ind., II, /. 145,//. 133, Ai,'. i, \a~c

('894) „ Undularis.

Bingham, Fauna Brit. India, I, /. 171 (1905). . „ „

Fruhstorfer, Iris, XX, /. 174 (1907) .... „ Hypermnestra.

„ (Seitz, Grossschni. d. Erde), /. 374,

//. 8;^ (191 I)

W. J. Batavia (3—14); Depok (95); Buitenzorg (265); Tjampea (160);
vicinity of the Pelabouan Ratou or Wijnkoops Bay and the Tjiletou or Sand
Bay (+ 1 50) on the south coast.

C J. Semarang (6); prov. Tegal; Touban on the north coast (6);
Magelang (500).

E. J. Sourabaya (6); Banyoupoutih, prov. Probolingo (45); Madoura.

This is one of the commonest butterflies in Java ; but not so abundant in
the mountains as in the lowlands where the principal, food of the larva, Cocos
Nucifera L., is found in plenty. It is everywhere met with in gardens and
the r? may at times be seen frolicking together in the sun so that any one,
not acquainted with the striking difference in colour between the male and
female, would doubtless conclude he was witnessing the preliminary play of the
sexes before pairing. For this is no fighting but manifest play ; I have, indeed,
witnessed it on one occasion for half an hour before they separated and went
their own way. Bates mentions the same fact with regard to 6 of small
Heliconidae. Once I noticed a c? of the present species diffusing a very
unpleasant odour on being strongly compressed. In the introduction I have
already dealt with the sexual colour dimorphism in this species and the inter-
pretation of mimicry in connection with it.

The reddish brown band along the lower margin on the upper side of the
hind-wings in the cf, as well as the general orange colour and black on the
upper side of the 9, is darker in some individuals and lighter in others without
seasonal differences being thereby indicated.

The caterpillar occurs on the leaves of ke/apa (Cocos Nuciffra L.), of
pinang (Areca Catechu L.), arcn (Arenga saccharifera Lab.), kiray (Metroxy-
i.oN Sagus Rottb.), and other species of palms. Young individuals are green

76

with black head, horns, and tail processes; in the full-grown larvae on the
general pale green colour occurs a double golden-yellow dorsal line and in
addition on either side tree similar longitudinal lines, the upper one being
widest and dilating at several of the rings to yellow spots which in turn contain
some orange, and sometimes even dark green, smaller spots. Ihe colour varies
somewhat in different individuals. The head is of a handsome pale brown or
sometimes yellow and has two setose horns of the same colour, between which
occur two vertical white or yellow stripes which curve downward, each behind
either horn. The hinder portion of the body terminates in two long setose
processes which may be either green, bright orange, or pale brown and which
receive the forementioned wide longitudinal lines on either side of the double
dorsal line. The larva does not, therefore, differ essentially from the Indian one
figured by Moore and probably pertains to the same species. The pupa possesses
the form, already alluded to, peculiar to the genus, and in colour resembles
the larva in a remarkable manner. It is suspended so close against leaves
that one would imagine it was attached by means a girdle thread. A chrysalis
of March 12''^ produced the butterfly on the 20"* of the same month. My
figures are very successful and fairly agree with those of Moore.

2. EsAKA Hew. (PI. XVIII, fig. 64a, 64(J).

Hewitson, Exo/. Bidterfl., Ill, Melaniiis,/ig. 5 (1863). Melanitis Esaka.
Moore, Joum. Lmn. Soc. of London, XXI, /. ZTi'^pl- 3.

fiff. 5 (1885) Dyctis Andersonii.

Fruhstorfer, Ent. Nadir., XX, /. 21 (1894) ■ • Elymnias Maheswara.
Stett. Ent. Zci/., 1894, /. 124, //. 4,

fig- 1 "i

Moore, Lep. hid., 11,/. xtc^, pi. 143,//^. 2, 2a (1894). Agrusia Andersonii.

Bingham, Fauna Brit. India, 1, />. 179 (1905) . . Elymnias Esaca.

Fruhstorfer, Iris, XX, /. 248 (1907) Mimadelias Maheswara.

„ (Seitz, Grossschm. d. Erdc), p. 392,

//. 90<5 (191 1) Elymnias

W. J. Gedeh mountains (i 250) ; mountains in the prov. Prayangan (+ 1 500).

C. J. ?

E. J. ?

My collection contains three cf of this species. A specimen identified by
Snellen as Vasudeva Moore is doubtless the 9 of the present species, the
Maheswara of Fruhsuorfer.

77

3. Panthera F.

Fabricus, Mant. Ins., Ill, /. 39, A^o. 40 (1787) . . . Papilio Panthera.

HoRSFiELD, Caf. Lep. M. E. I. C, pi. ^, fig. 7, 7r(i828). Melanitis Dusara.
Butler, An7i. Mag. N'af. Hist., Scr. 3, 20, p. 404,//. 9,

fig. 10 (1867) Elymnias Lutescens.

Distant, Rhop. Mai, p. 62, tab. t,fig. 4, 5 (1882— 1886).

Moore, Lep. Ind., II, /. 152,/'/. \s^,fig. 2, 2a — (5(1894). .. Mimus.

Fruhstorfer, Iris, XX, /. 216 (1907) „ Panthera.

„ (Seitz, Grossschm. d. Erdc), p. 371, //.

883 (191 1) ,

W. J. Tjampea (160); mount Pantjar, district Buitenzorg; mountains in

the prov. Prayangan (1500).

C.J. ?

E. J. Prov. Passourouan.

This species is not rare at Tjampea but elsewhere, apparently, it is not at
all common. In the lowlands I never met with it. The specimens in my
possession from W. Java, especially those from Tjampea, belong to the form
named Dusara by Fruhstorfer ; the colour of the cf is not always equally
dark brown, some approaching the 9 in shade. A d from the prov. Passou-
rouan completely agrees with the latter but not with the form Dulcibella
figured by him, which is unknown to me.

The larvae were repeatedly observed on aren oiitan (Arenga Obtusifolia
Mart.), pinang rendeh (Pinanga Kuhlii B1.), and kiray Metroxylon Sagus
Rottb.), I was unable to distinguish it from the larva of E. Undularis Drury.
The same was the case with the pupae which, however, were particularly
brightly coloured ; I have notes of one chrysalis being ornamented on the
beauiful green colour with longitudinal yellow lines mixed with handsome red,
containing here and there oblong black spots. Dr. L. Martin, in the Deutsche
Entomologische National- Bibliotlick 19 10, A^o. 12, gave a description of the larva
and pupa bred by him .at Sintang in West Borneo. From a .spherical milky-
white ^^% deposited on the leaf of a dwarf species of palm the caterpillar was
hatched after four days, being yellowish white at first but which after feeding
became yellowish green with black head, when is gradually changed into the
full-grown larva whose description on the whole answers to that given by me
of the larva of E. Undularis Drury, but in which besides the double dorsal
line only a single lateral line on either side is said to occur and whose horns
are said to be black and to terminate in three points. The pupa also has the

78

same form as mine and a system of coloration practically conforming with that
observed by me. Dr. Martin also specifically refers to the bright colour of
the pupa and to the fact — likewise observed by me in the chrysalis of
E. Undularis Drury — that its ventral surface is firmly appressed to the leaf
to which it is attached by the caudal extremity.

4. Dara Dist. (PI. XVIII, fig. 59).

Distant, Ajui. Mao. A^af. Hist. Ser. 5, 19,/. 50(1884). Elymnias Dara.

Staudinger, Iris, II, /. 39 (1889) „ Albofasciata.

DE Nic^viLLE, Journ. Bombay Soc. Nat. Hist. V, /. 202,

//. D, fig. 4 (1890) Dyctis Daedalion.

Moore, Lep. Ind., 11,/. 154,//. 137, /ig. 2, 2^(189/)). Elymnias Daedalion.
Bingham, Fauna Brit. India, I, /. 178 (1905). . . „ Dara.

Fruhstorfer, Iris, XX, /. 214 (1907) „

„ (Seitz, Grossschm. d. Erde), p. 373,

//. 88^ (191 1)

W. J. Tjibougel (+ 1000) in the prov. Prayangan ; vicinity of the Pela-
bouan Ratou or Wijnkoops Bay (+ 1 50).

C. J. ?

E. J. Banyouwangy (+ 500).

Fruhstorfer calls the figured form Diminuta and distinguishes somewhat
larger forms from W. Java under the name of Bengena.

From W. J. I only possess cT and from E. J. a single 9- I was, therefore,
unable to form a judgment, although I doubt this difference in size. These cf are
however, much darker in colour than this 9 and the white on the upper side
of the former is tinted with beautiful violet.

5. Casiphone Hb. (PI. XVIII, fig. 60a, bob).

HuBNER, Samml. Exot. Sclim., Ill (1806 — 1827). . . Elymnias Casiphone.
Distant, /?//o/!>. J/fl/.,/. 64, 7a(5. 6,yfo^. 10 (1882 — 1886).

Fruhstorfer, Iris, XX, /. 207 (1907) „ „

„ (Seitz, Grossschm. d. Erde), p. 2>^i, pi.
Sye (191 1) ,, »

W. J. Sindanglaya (1074) ; Gedeh mountains (i 250) ; Salak mountains (780).

C. J. ?

E. J. Without precise indication of habitat.

79

At Tjampea I once found, attached to a Musa leaf, a pupa, resembling that
of E. Undularis Drury, from which a butterfly belonging to the present
species emerged. I am quite unable to agree with the observations concerning
this species in the well-known work of Seitz. The upper side of the d* has,
it is true, a great resemblance to that in Euploea Midamus L., and the 9 less
so; nevertheless, the form and flight differ so much in the two species that
they can scarcely be confused. Although in some o' the same metallic gloss
occurs on the upper side of the fore-wings as in the Euploea referred to,
nothing of this is to be seen in several d in my possession. I only possess
a single cf from E. J. — a very good specimen by the way — and I can detect
no difference between this and my W. J. specimens. The upper side of the
fore-wings is equally dark as in the most glossy <d from W. J. and the blue
metallic gloss, although not pronounced, is, nevertheless, distinctly in evidence.
Consequently it does not quite agree with the figure given by Fkuhstorfer
of his E. J. form Alumna.

6. Lais Cram. (PI. XVIII, fig. bia, 6i/;, 6ir, tie/, die, bia, til).

Cramer, II, /. 21, //. iio A. B. (1779) Papilio Lais.

Distant, Rhop. Mai, p. 62, iab. c,, fig. 2 (1S82 — 1886). Elymnias Lais.
Staudinger, E.xot. Sc/un., p. 237, //. 86 cT (1884 — 1888). „ „

Hagen, Ja/ird. Nass. Vcr. Naiurk., XLIX, /. 184,//. 4,

Jig. 6 (1896) „ Baweana.

Fruhstorfer, Iris, XX, /. 200 (1907) „ Nesaea.

„ (Seitz, Grossschm. d. Erdc), p. 379, />/.

87r, rt' (1911) ,.,... „ „

W. J. Batavia (3 — 14); Buitenzorg (265); Tjibougel (+ 1000); mountains
in the prov. Prayangan; vicinity of the Pelabouan Ratou or Wijnkoops Bay
on the south coast (+ 150).

C. J. Prov. Tegal; prov. Rembang; Magelang (500).

E. J. Malang (443).

This butterfly is common in Java wherever the coco-nut palm grows,
especially in plantations of that tree ; it flies, as has already been noted above,
in the twilight. My cT specimen from E. J. is not shot with bluish green on
the upper side like the cT from W. J., especially from Batavia, but does not
otherwise differ. Fruhstorfer separates the E. J. specimens from those of
W. J. as the form Herml\. The form from the island of Bavvean differs

'9

So

distinctly from the Java form in both sexes having the peculiar marking on the
upper surface of the fore-wings mostly covered by reddish brown or rather in
having retained the former obfuscation in that portion of the fore-wings. My
specimens, however, from the mountains of the prov. Prayangan — not those
from Wijnkoops Bay — show a decided increase of this brown, especially near
the apex of the fore-wings, while the Leiden Museum possesses a 9 from
Malang in E. J. in which this feature is almost ecpally pronounced as in the
Bawean individuals. This difference of colouring again is surely due, not to
local influence, but to the unequal advance in the same process of colour
evolution. For this reason I have considered it desirable to figure these different
forms. The larva and pupa are of the same type as in E. Undularis Drury
and occur likewise on leaves of the kclapa (Cocos Nucifera L.) and /^iiiaiig
(Areca Catechu L ). The caterpillar attains a length of 6.5 centimetre, is
green with eight golden-yellow longitudinal lines as in E. Undularis Drury,
but the subdorsal pair, which is similarly continued on the head and the tail
spines, while it has been also united to a single stripe does not show the small
spots peculiar to that species. The head terminates in two not very long,
setose, horns which, together with the head, are pale flesh-coloured. The tail
processes are long, spiny, and yellow. In the young larvae, however, the
horns as well as the tail spines are black. The pupa has the same rigidity as
that of the species just referred to, is of a delicate green with four streaks com-
mencing from the tail end, consisting of non-amalgamated lines, partly hand-
some red, partly yellow ; the process on the heatl is similarly coloured. The
larva pupated against the wall of the bottle in which it had been reared, but
it had previously covered that portion to which its ventral side was attached
with a fine web.

7. Kamara Moore. (PL XVIII, fig. 62).

Moore, CaL Lep. M. E. I. C, I, /. 239, No. 516

(1857) Elymnias Kamara.

de Nicevillk, Jotiru. Bombay Soc. Nat. His/., X, /. 19,

//. R, /^. 9. 10. II (1895) .. Erinyes.

Fruhstorfer, (Seuz, Grossschvi. d. Erdc), p. 382, //. 87^?

(igii) „ Kamara.

W. J. Buitenzorg (265); Sindanglaya (1074); Prayangan mountains.

C. J. ?

E. J. Near Lake Klakah (230) in the prov. Probolingo.

8i

8. Ceryx Bsd. (PI. XVIII, fig. 63).

BoisDUVAi., Spec. Gen., I, //. 9> A?- 38 (1836) Melanitis Ceryx.

Fruhstorfer, /ri's, XX, /. 212 (1907) Elymnias „

„ (Seitz, Grossse/im. d. Eide), /. 383 (191 i) . „ „

W. J. Tjibodas (14 10); mountains in the prov. Prayangan.

C. J. ?

E. J. ?

Fruhstorp'er also only knows this species from the mountains of W. J.,
where, however, it is not rare, the type and the dark form called Hestonia
by him, occuring together. In the colour of the sexes I can see no noteworthy
difference. According to Hagen this species is also a mountains butterfly in
Sumatra.

9. KiiNSTiERi Honr.

Fruhstorfer, Ent. A'ac/tr., XX, No. 3, /. 43 (1894) • Elymnias Gauroides.
Berl. Ent. Zeitschr., XXXIX, /. 2^^^ pl-

18. A- 4 (1894)

Fruhstorfer, (Settz, Grossschvi. d. Erdc), p. 2,%a^ {\<^\\). „ Ktinstieri.

Mr. Fruhstorfer captured a 9 of this species at Tjisewou in the prov.
Prayangan at an altitude of about 700 Metres. This is the only specimen
known from Java.

INDEX

OF

Java DANAIDAE, SATYRIDAE, RAGADIDAE and ELYMNIADAE.

Page.

1. EuPLOEA Climena, Cram 4

2. „ Malayica, Bull 5

3. „ Deheeri. Doherty .... 6

4. „ Crameri, Luc 7

5. „ MiDAMUS L 7

6. „ Mazare';, Moore .... 10

7. „ Eleusina, Cram 11

8. „ Leucostictos, Grael. ... 12

9. „ Phoebus, ButI 13

10. „ Radaiiaxthus, Felder . . 14

11. „ EijNDHOVii, Felder. ... 15

12. „ Huebneri, Moore .... 15

13. „ Menetriesii, Pelder ... 16

14. „ Rafflesii, Moore .... 16

15. „ Gameua, Hb 17

16. „ Gloriosa, Bull 18

17. Hestia Belta, Westw 19

18. „ Lynceus, Drury .20

19. „ Leuconoe, Erichs 21

20. Ideopsis Gaura, Horsf. 22

21. Danais Albata, Zinck 24

22. „ Melaneus, Cram 24

23. „ Aglea, Cram 25

24. „ Aspasia F 26

25. „ Agleoides, Felder .... 27

26. „ SiMiLis, Clerck 28

27. „ JuvENTA, Cram 29

28. „ LiMNiACE, Cram 33

29. „ Melanu'Pus, Cram 32

30. „ Artenice, Cram 33

31. „ Gknutia, Cram 34

32. „ Chrysippus L 36

33. Cyllo Leda L 39

34. „ Suyudana, Moore 47

35. PYRITES Madura, Horsf. 48

36
37
38
39

40
41
42,
43
44

45
46,

47
4S

49
50
51
52
53
54
55
56
57
58

59
60
61

62

63
64

65
66

67
68
69

Page.

Erites Ochreana, Semp 48

Ypthima Jarba, de Nic 49

HOuNERi, Kirby .... 50

Argillosa, Snell 51

Bai.dus, F 51

Fasciata, Hew 52

Pandocus, Moore .... 54

Neorina Chrisna, Westw 54

Debis Minerva F 56

Chandica, Moore 57

Samio, Dbd 57

Manthara, Felder 58

Darena, Felder 59

Dyrta, Felder 59

Europa F 59

Rohria, F 60

Orsotriaena Medus F 61

Mycalesis Nala, Felder 63

Moorei Felder 65

„ Mineus L 66

„ Horsfieldi, Moore . . 66

„ Perseus F 68

„ Janardana, Moore . . 69

„ FuscA, Felder 70

„ Oroatjs, Hew 71

Ragadia Crisia, Hb 72

Elymnias Undularis, Drury ... 74

„ EsAKA, Hew 76

Panthera F 77

Dara, Dist 78

Casiphone, Hbn. . .

Lais, Cram

Kamara, Mooie . . .

Cery-x, Bsd 8 1

Kunstieri, Honr. ... 81

78

79
80

EXPLANATION OF PLATE XL

Fig. I. EuPLOEA Climena, Cram, form Sepulchralis But). V- a (Batavia^; b id. (/ (C. Java);
c cT (Amboyna); d form Eurypon Hew. $ (Ke islands).

„ 2. Euploea Malayica Bull, a imago cT; b larva.

„ 3. Euploea Crameri Luc. a ^■, b ^.

„ 4. Euploea Midamus L. form Basilissa Cram, a imago cf ; 5 id $; r pupa; (only the

form of the pupa is sufficient; its strong metallic lustre does not show in the illustration);

d, e caterpillars.
„ 5. Euploea Mazares Moore, a imago cT; b larva.
„ 6. Euploea Eleusina Cram, cf-

riKl'KRS & Snfxlf.N, The Rhop. of Java.

riatc XI

Printed by P. JV. M. Trap^ Leiden.

EXPLANATION OF PLATE XII.

Fig. 7. EuPLOEA Leucostictos Gmel. a imago cT; b id. form Nemesthes Hb. (Saparoua) ;
c id. form Assimilata Felder (Ke islands); d id. 9 form Viola Bull. (Celebes),
e larva.
8. EupLOEA Phoebus Bull. cT-
„ 9. Euploea Radamanthus F. form Alcidice. a cf; * $.
„ 10. Euploea Eijndhovii Felder. (j\
„ II. Euploea Hubneri Moore, cf.

„ 12. Euploea Menetriesii Felder cT form Wallengreni. Fruhst. cf.
„ 13. Euploea Rafflesi Moore, a imago cf ; '' '<'■ 9; ^ larva.

PiEPF.RS & Snellen, The Rhop. of Java.

Plate XII

r rimed by F. W. M. Trap, Leiden.

EXPLANATION OF PLATE XIII.

Fig. 14. Eupi.oEA Gamki.ta Hb. a c? (on the lower part of the upper side of the fore-wings
occurs a dull-lustrous spot while the lower part of the under side of the fore-wings
as well as the fore part of the upper side of the hind- wings are also dull-lustrous;
this is not shown in the figures) ; d 9-

15. EuPLOEA Gloriosa Butl. a form Lacordairei Moore cf (Java); 6 form Schlegelii
Felder 9- (Celebes).

16. Hestia Lynceus Drury. a larva (an immature specimen); 6 pupa.

17. Ideopsis Gaura Horsf. a imago cf; 6 id. 9; f larva (immature); d pupa.

18. Danais Albata Zinck cf.
iq. Danai.s Melaneus Cram., form Larisca Felder cT-

20. Danais Aglea Cram, a imago cf ; ^ larva.

21. Danais Aspasia F. a imago cf; 6 larva; c pupa. (The pupa is not occurately reproduced).

22. Danais Agleoides Felder. a imago cT; f> larva; c pupa. (The figura of the pupa is
not very accurate).

20

PiEPERs & Snellen, The Rhop. of Java.

riate XIII

Prinleii hy J'. IV. M. T/afi., Lcidm.

EXPLANATION OF PLATE XIV.

Fig. 23. Danais Similis L. a imago; b larva; c pupa.
„ 24. Danais Juventa Cram, a imago ^ ; b larva; c pupa.
„ 25. Danais Limniace Cram, a form Melissa Cram. (The figure while reproducing the

general impress is nevertheless not very accurate); b form Septentrionis Dist.;

c form Obscurata Butl.

26. Danais Artenice Cram, cf .

27. Danais Melanippus Cram, a imago d^; b larva.

28. Danais Genuiia Cram, a imago (f; b larva; f larva (dark form); d pupa.

29. Danais Chrvsippus L. a imago cf; 1^ id. form Petilia Stoll (Tegal, C. J.); c larva;
d id. immature ; e, f pupae.

32. Erites Madura Horsf. cf-

33. Erites Ochreana Semp. $.

34. Ypthima Jarba de Nic. form Gangamala Fruhst. $.

PiEPF.RS & Snellen, The Rliop. of Java.

Plate XIV

PrinUJ by P. W. M. Trap, Laden.

EXPLANATION OF PLATE XV.

Fig. 30. Cyllo Leda L. a Representation of the ocelli on the underside; b ocellus magnified;
c upperside form A ; d underside form A' ; e upperside form B ; / upperside form C
(to dark in colour); g underside form C (to dark in colour); h underside form D;
i underside form E ; k underside form F ; / underside form H ; m underside form G ;
« underside form I ; 0 upperside form K ; p underside form K' ; q upperside of a
specimen from E. J.; r Cyllo Constantia Cram. (Moluccas), to demonstrate how
the black is gradually diffused over the original colour.

„ 31. Cyllo Suvudana Moore, a (f; b id.; c larva; (/pupa.

PlEPERS & Snki.len, The Rhop. of Java

30*

Printed by P. IV. M. Trap^ Leiden.

EXPLANATION OF PLATE XVI.

F'g- 35- Ypthima Hubneri Kirby.

„ 36. Ypthima Argillosa Snell. a cT; ^ 9-

„ 37. Ypthima Baldus F. <j cf ; i $.

„ 38. Ypthima Baldus F. form Horspieldi Moore, a imago c? ; ^ id 9- The colour is
much too light and does not differ much from that of the cf ; it is suffused with a
yellow brown sheen; c larva; (/ pupa.

39. Ypthima Fasciata Hew. 9-

40. Ypthima Pandocus Moore, a imago cT. (The colour is too dark) ; 3 id. 9 ; c id.
form Nigricans Snell. d'; d, e larvae; / pupa.

41. Neorina Crishna Westw. cf-

42. Debis Minerva F. a cf ; ^ 9-

43. Debis Chandica Moore, a c^; b '^.

44. Debis Samio Dbd. cf-

45. Debis Manthara Felder. a imago cf; '!' id. 9; ^ larva; d pupa.

PiKPKRS & Snkllen, The R/iop. of Java.

Plate XVI

Printed iy P. W. M. Trap, Leiden.

EXPLANATION OF PLATE XVIL

Fig. 46. Debis Darena Felder. a cf ; ^ 5.
„ 47. Debis Dyrta Felder. a (f; 6 '$.
„ 48. Debis Europa F. a c?; * 9-
„ 49. Debis Rohria F.

„ 50. Oksotriaena Medus F. a larva; 6 pupa.
„ 51. Mycalesis Nala Felder. a o'; b id. with large ocelli; c id. 9; id.; d id. loimSuDRA

Felder.
„ 52. Mycalesis Moorei Felder. cf.
„ 53. Mycalesis Horsfiei.di Moore, a imago ci'; l> id. V; c id. specimen with small ocelli;

d larva.
„ 54. Mycalesis Perseus F. « cT; ^ 9; ^ specimen with small ocelli.

21

PiEr'F.RS & Snellen, The Rlwp. of Java.

463

riate XVII

Prin/eJ by P. IV. M. Trap, Leiden.

EXPLANATION OF PLATE XVIII.

Fig. 55. Mycalesis Janardana Moore, cf-

„ 56. Mycalesis Fusca Felder cT-

„ 57. Mycalesis Oroatis Hew. c?.

„ 58. Elymnias Undularis F. a larva; b pupa.

„ 59. Elymnias Dara Distant, (j". (The centre of the ocelli on the hind-wings is darker).

„ 60. Elymnias Casiphone Hb. a iS; b ^.

„ 61. Elymnias Lais Cram, a imago cf (Batavia); 6 9 (Batavia); c $ (Prayangan); (/9 Ambarawa.

(C. J.); e (marked also c in error) 9 (island Bawean); / larva; g pupa.

„ 62. Elymnias Kamara Moore. 9-

„ 63. Elymnias Cery.x Bsd. (f.

„ 64. Elymnias Esaka Hew. a (j^; b ^.

I'li-.i'KRS & Snki.lkn, Till- Rliop. of Java.

Plate XVIII

Prinleii by P. W. M. Trap, Laden.

/Vv^-<-'l

The complete series of

THE RHOPALOCERA OF JAVA

will consist of:

PIERIDAE, With 4 coloured plates Gld. 18.—

HESPERIDAE, With 6 coloured plates .... Gld. 25.—

DANAIDAE, SATYRIDAE, RAGADIDAE,

ELYMNIADAE, With 8 coloured plates. . . Gld. 37.50

In preparation :

LIBYTHEIDAE, ERYCINIDAE. LYCAENIDAE.

PAPILIONIDAE.

HELICONIDAE, EURYTELIDAE, NYMPHALIDAE.