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[_From the_ TRANSACTIONS _of the_ LINNEAN SOCIETY, vol. xxv.]
I. _On the Phenomena of Variation and Geographical Distribution as
illustrated by the_ Papilionidæ _of the Malayan Region. By_ ALFRED R.
WALLACE, _Esq._
(Plates I.–VIII.)
Read March 17, 1864.
When the naturalist studies the habits, the structure, or the affinities
of animals, it matters little to which group he especially devotes
himself; all alike offer him endless materials for observation and
research. But, for the purpose of investigating the phenomena of
geographical distribution and of local or general variation, the several
groups differ greatly in their value and importance. Some have too
limited a range, others are not sufficiently varied in specific forms,
while, what is of most importance, many groups have not received that
amount of attention over the whole region they inhabit, which could
furnish materials sufficiently approaching to completeness to enable us
to arrive at any accurate conclusions as to the phenomena they present
as a whole. It is in those groups which are and have long been
favourites with collectors that the student of distribution and
variation will find his materials the most satisfactory, from their
comparative completeness.
Preeminent among such groups are the diurnal Lepidoptera or Butterflies,
whose extreme beauty and endless diversity have led to their having been
assiduously collected in all parts of the world, and to the numerous
species and varieties having been figured in a series of magnificent
works, from those of Cramer, the contemporary of Linnæus, down to the
inimitable productions of our own Hewitson. But, besides their
abundance, their universal distribution, and the great attention that
has been paid to them, these insects have other qualities that
especially adapt them to elucidate the branches of inquiry already
alluded to. These are the immense development and peculiar structure of
the wings, which not only vary in form more than those of any other
insects, but offer on both surfaces an endless variety of pattern,
colouring, and texture. The scales with which they are more or less
completely covered imitate the rich hues and delicate surfaces of satin
or of velvet, glitter with metallic lustre, or glow with the changeable
tints of the opal. This delicately painted surface acts as a register of
the minutest differences of organization,—a shade of colour, an
additional streak or spot, a slight modification of outline continually
recurring with the greatest regularity and fixity, while the body and
all its other members exhibit no appreciable change. The wings of
Butterflies, as Mr. Bates has well put it[1], “serve as a tablet on
which Nature writes the story of the modifications of species;” they
enable us to perceive changes that would otherwise be uncertain and
difficult of observation, and exhibit to us on an enlarged scale the
effects of the climatal and other physical conditions which influence
more or less profoundly the organization of every living thing.
Footnote 1:
See ‘The Naturalist on the Amazons,’ 2nd edit. p. 412.
A proof that this greater sensibility to modifying causes is not
imaginary may, I think, be drawn from the consideration that while the
Lepidoptera as a whole are of all insects the least essentially varied
in form, structure, or habits, yet in the number of their specific forms
they are not much inferior to those orders which range over a much wider
field of nature, and exhibit more deeply seated structural
modifications. The Lepidoptera are all vegetable-feeders in their
larva-state, and suckers of juices or other liquids in their perfect
form. In their most widely separated groups they differ but little from
a common type, and offer comparatively unimportant modifications of
structure or of habits. The Coleoptera, the Diptera, or the Hymenoptera,
on the other hand, present far greater and more essential variations. In
either of these orders we have both vegetable- and animal-feeders,
aquatic, and terrestrial, and parasitic groups. Whole families are
devoted to special departments in the economy of nature. Seeds, fruits,
bones, carcases, excrement, bark, have each their special and dependent
insect tribes from among them; whereas the Lepidoptera are, with but few
exceptions, confined to the one function of devouring the foliage of
living vegetation. We might therefore anticipate that their population
would be only equal to those of the sections of the other orders that
have a similar uniform mode of existence; and the fact that their
numbers are at all comparable with those of entire orders, so much more
varied in organization and habits, is, I think, a proof that they are in
general highly susceptible of specific modification.
The Papilionidæ are a family of diurnal Lepidoptera which have hitherto,
by almost universal consent, held the first rank in the order; and
though this position has recently been denied them, I cannot altogether
acquiesce in the reasoning by which it has been proposed to degrade them
to a lower rank. In Mr. Bates’s most excellent paper on the
Heliconidæ[2], he claims for that family the highest position, chiefly
because of the imperfect structure of the fore legs, which is there
carried to an extreme degree of abortion, and thus removes them further
than any other family from the Hesperidæ and Heterocera, which all have
perfect legs. Now it is a question whether any amount of difference
which is exhibited merely in the imperfection or abortion of certain
organs, can establish in the group exhibiting it a claim to a high grade
of organization; still less can this be allowed when another group,
along with perfection of structure in the same organs, exhibits
modifications peculiar to it, together with the possession of an organ
which in the remainder of the order is altogether wanting. This is,
however, the position of the Papilionidæ. The perfect insects possess
two characters quite peculiar to them. Mr. Edward Doubleday, in his
‘Genera of Diurnal Lepidoptera,’ says, “The Papilionidæ may be known by
the apparently four-branched median nervule and the spur on the anterior
tibiæ, characters found in no other family.” The four-branched median
nervule is a character so constant, so peculiar, and so well marked, as
to enable a person to tell, at a glance at the wings only of a
butterfly, whether it does or does not belong to this family; and I am
not aware that any other group of Butterflies, at all comparable to this
in extent and modifications of form, possesses a character in its
neuration to which the same degree of certainty can be attached. The
spur on the anterior tibiæ is also found in some of the Hesperidæ, and
is therefore supposed to show a direct affinity between the two groups;
but I do not imagine it can counterbalance the differences in neuration
and in every other part of their organization. The most characteristic
feature of the Papilionidæ, however, and that on which I think
insufficient stress has been laid, is undoubtedly the peculiar structure
of the larvæ. These all possess an extraordinary organ situated on the
neck, the well-known =Y=-shaped tentacle, which is entirely concealed in
a state of repose, but which is capable of being suddenly thrown out by
the insect when alarmed. When we consider this singular apparatus, which
in some species is nearly half an inch long, the arrangement of muscles
for its protrusion and retraction, its perfect concealment during
repose, its blood-red colour, and the suddenness with which it can be
thrown out, we must, I think, be led to the conclusion that it serves as
a protection to the larva by startling and frightening away some enemy
when about to seize it, and is thus one of the causes which has led to
the wide extension and maintained the permanence of this now dominant
group. Those who believe that such peculiar structures can only have
arisen by very minute successive variations, each one advantageous to
its possessor, must see, in the possession of such an organ by one
group, and its complete absence in every other, a proof of a very
ancient origin and of very long-continued modification. And such a
positive structural addition to the organization of the family,
subserving an important function, seems to me alone sufficient to
warrant us in considering the Papilionidæ as the most highly developed
portion of the whole order, and thus in retaining it in the position
which the size, strength, beauty, and general structure of the perfect
insects have been generally thought to deserve.
Footnote 2:
Transactions of the Linnean Society, vol. xxiii. p. 495.
The Papilionidæ are pretty widely distributed over the earth, but are
especially abundant in the tropics, where they attain their maximum of
size and beauty and the greatest variety of form and colouring. South
America, North India, and the Malay Islands are the regions where these
fine insects occur in the greatest profusion, and where they actually
become a not unimportant feature in the scenery. In the Malay Islands in
particular the giant Ornithopteræ may be frequently seen about the
borders of the cultivated and forest districts, their large size,
stately flight, and gorgeous colouring rendering them even more
conspicuous than the generality of birds. In the shady suburbs of the
town of Malacca two large and handsome Papilios (_Memnon_ and
_Nephelus_) are not uncommon, flapping with irregular flight along the
roadway, or, in the early morning, expanding their wings to the
invigorating rays of the sun. In Amboyna and other towns of the
Moluccas, the magnificent _Deiphobus_ and _Severus_, and occasionally
even the azure-winged _Ulysses_, frequent similar situations, fluttering
about the orange-trees and flower-beds, or sometimes even straying into
the narrow bazaars or covered markets of the city. In Java the
golden-dusted _Arjuna_ may often be seen at damp places on the roadside
in the mountain districts, in company with _Sarpedon_, _Bathycles_, and
_Agamemnon_, and less frequently the beautiful swallow-tailed
_Antiphates_. In the more luxuriant parts of these islands one can
hardly take a morning’s walk in the neighbourhood of a town or village
without seeing three or four species of _Papilio_, and often twice that
number. No less than 120 species of the family are now known to inhabit
the Archipelago, and of these ninety-six were collected by myself.
Twenty-nine species are found in Borneo, being the largest number in any
one island, twenty-three species having been obtained by myself in the
vicinity of Sarawak; Java has twenty-seven species; Celebes and the
Peninsula of Malacca twenty-three each. Further east the numbers
decrease, Batchian producing seventeen, and New Guinea only thirteen,
though this number is certainly too small, owing to our present
imperfect knowledge of that great island.
In estimating these numbers I have had the usual difficulty to
encounter, of determining what to consider species and what varieties.
The Malayan region, consisting of a large number of islands of generally
great antiquity, possesses, compared to its actual area, a great number
of distinct forms, often indeed distinguished by very slight characters,
but in most cases so constant in large series of specimens, and so
easily separable from each other, that I know not on what principle we
can refuse to give them the name and rank of species. One of the best
and most orthodox definitions is that of Pritchard, the great
ethnologist, who says, that “_separate origin and distinctness of race,
evinced by a constant transmission of some characteristic peculiarity of
organization_,” constitutes a species. Now leaving out the question of
“origin,” which we cannot determine, and taking only the proof of
separate origin, “_the constant transmission of some characteristic
peculiarity of organization_,” we have a definition which will compel us
to neglect altogether the _amount_ of difference between any two forms,
and to consider only whether the differences that present themselves are
_permanent_. The rule, therefore, I have endeavoured to adopt is, that
when the difference between two forms inhabiting separate areas seems
quite constant, when it can be defined in words, and when it is not
confined to a single peculiarity only, I have considered such forms to
be species. When, however, the individuals of each locality vary among
themselves, so as to cause the distinctions between the two forms to
become inconsiderable and indefinite, or where the differences, though
constant, are confined to one particular only, such as size, tint, or a
single point of difference in marking or in outline, I class one of the
forms as a variety of the other.
I find as a general rule that the constancy of species is in an inverse
ratio to their range. Those which are confined to one or two islands are
generally very constant. When they extend to many islands, considerable
variability appears; and when they have an extensive range over a large
part of the Archipelago, the amount of unstable variation is very large.
These facts are explicable on Mr. Darwin’s principles. When a species
exists over a wide area, it must have had, and probably still possesses,
great powers of dispersion. Under the different conditions of existence
in various portions of its area, different variations from the type
would be selected, and, were they completely isolated, would soon become
distinctly modified forms; but this process is checked by the dispersive
powers of the whole species, which leads to the more or less frequent
intermixture of the incipient varieties, which thus become irregular and
unstable. Where, however, a species has a limited range, it indicates
less active powers of dispersion, and the process of modification under
changed conditions is less interfered with. The species will therefore
exist under one or more permanent forms according as portions of it have
been isolated at a more or less remote period.
What is commonly called variation consists of several distinct phenomena
which have been too often confounded. I shall proceed to consider these
under the heads of—1st, simple variability; 2nd, polymorphism; 3rd,
local forms; 4th, coexisting varieties; 5th, races or subspecies; and
6th, true species.
1. _Simple variability._—Under this head I include all those cases in
which the specific form is to some extent unstable. Throughout the whole
range of the species, and even in the progeny of individuals, there
occur continual and uncertain differences of form, analogous to that
variability which is so characteristic of domestic breeds. It is
impossible usefully to define any of these forms, because there are
indefinite gradations to each other form. Species which possess these
characteristics have always a wide range, and are more frequently the
inhabitants of continents than of islands, though such cases are always
exceptional, it being far more common for specific forms to be fixed
within very narrow limits of variation. The only good example of this
kind of variability which occurs among the Malayan Papilionidæ is in
_Papilio Severus_, a species inhabiting all the islands of the Moluccas
and New Guinea, and exhibiting in each of them a greater amount of
individual difference than often serves to distinguish well-marked
species. Almost equally remarkable are the variations exhibited in most
of the species of _Ornithoptera_, which I have found in some cases to
extend even to the form of the wing and the arrangement of the nervures.
Closely allied, however, to these variable species are others which,
though differing slightly from them, are constant and confined to
limited areas. After satisfying oneself, by the examination of numerous
specimens captured in their native countries, that the one set of
individuals are variable and the others are not, it becomes evident that
by classing all alike as varieties of one species we shall be obscuring
an important fact in nature, and that the only way to exhibit that fact
in its true light is to treat the invariable local form as a distinct
species, even though it does not offer better distinguishing characters
than do the extreme forms of the variable species. Cases of this kind
are the _Ornithoptera Priamus_, which is confined to the islands of
Ceram and Amboyna, and is very constant in both sexes, while the allied
species inhabiting New Guinea and the Papuan Islands is exceedingly
variable; and in the island of Celebes is a species closely allied to
the variable _P. Severus_, but which, being exceedingly constant, I have
described as a distinct species under the name of _Papilio Pertinax_.
2. _Polymorphism or dimorphism._—By this term I understand the
coexistence in the same locality of two or more distinct forms, not
connected by intermediate gradations, and all of which are occasionally
produced from common parents. These distinct forms generally occur in
the female sex only, and the intercrossing of two of these forms does
not generate an intermediate race, but reproduces the same forms in
varying proportions. I believe it will be found that a considerable
number of what have been classed as _varieties_ are really cases of
_polymorphism_. Albinoism and melanism are of this character, as well as
most of those cases in which well-marked varieties occur in company with
the parent species, but without any intermediate forms. Under these
circumstances, if the two forms breed separately, and are never
reproduced from a common parent, they must be considered as distinct
species, contact without intermixture being a good test of specific
difference. On the other hand, intercrossing without producing an
intermediate race is a test of dimorphism. I consider, therefore, that
under any circumstances the term ‘variety’ is wrongly applied to such
cases.
The Malayan Papilionidæ exhibit some very curious instances of
polymorphism, some of which have been recorded as varieties, others as
distinct species; and they all occur in the female sex. _Papilio
Memnon_, L., is one of the most striking, as it exhibits the mixture of
simple variability, local and polymorphic forms, all hitherto classed
under the common title of varieties. The polymorphism is strikingly
exhibited by the females, one set of which resemble the males in form,
with a variable paler colouring; the others have a large spatulate tail
to the hinder wings and a distinct style of colouring, which causes them
closely to resemble _P. Coon_, a species of which the sexes are alike
and inhabiting the same countries, but with which they have no direct
affinity. The tailless females exhibit simple variability, scarcely two
being found exactly alike even in the same locality. The males of the
island of Borneo exhibit constant differences of the under surface, and
may therefore be distinguished as a local form, while the continental
specimens, as a whole, offer such large and constant differences from
those of the islands that I am inclined to separate them as a distinct
species—_P. Androgeus_, Cr. We have here, therefore, distinct species,
local forms, polymorphism, and simple variability, which seem to me to
be distinct phenomena, but which have been hitherto all classed together
as varieties. I may mention that the fact of these distinct forms being
one species is doubly proved. The males, the tailed and tailless
females, have all been bred from a single group of the larvæ, by Messrs.
Payen and Bocarmé, in Java, and I myself captured in Sumatra a male _P.
Memnon_, L., and a tailed female _P. Achates_, Cr., “in copulâ.”
_Papilio Pammon_, L., offers a somewhat similar case. The female was
described by Linnæus as _P. Polytes_, and was considered to be a
distinct species till Westermann bred the two from the same larvæ (see
Boisduval, ‘Species Générales des Lépidoptères,’ p. 272). They were
therefore classed as sexes of one species by Mr. Edward Doubleday, in
his ‘Genera of Diurnal Lepidoptera,’ in 1846. Later, female specimens
were received from India closely resembling the male insect, and this
was held to overthrow the authority of M. Westermann’s observation, and
to reestablish _P. Polytes_ as a distinct species; and as such it
accordingly appears in the British Museum List of Papilionidæ in 1856,
and in the Catalogue of the East India Museum in 1857. This discrepancy
is explained by the fact of _P. Pammon_ having two females, one closely
resembling the male, while the other is totally different from it. A
long familiarity with this insect (which, replaced by local forms or by
closely allied species, occurs in every island of the Archipelago) has
convinced me of the correctness of this statement; for in every place
where a male allied to _P. Pammon_ is found, a female resembling _P.
Polytes_ also occurs, and sometimes, though less frequently than on the
continent, another female closely resembling the male; while not only
has no male specimen of _P. Polytes_ yet been found, but the female
(_Polytes_) has never yet been found in localities to which the male
(_Pammon_) does not extend. In this case, as in the last, distinct
species, local forms, and dimorphic specimens have been confounded under
the common appellation of varieties.
But, besides the true _P. Polytes_, there are several allied forms of
females to be considered, namely, _P. Theseus_, Cr., _P. Melanides_, De
Haan, _P. Elyros_, G. R. G., and _P. Romulus_, L. The dark female
figured by Cramer as _P. Theseus_ seems to be the common and perhaps the
only form in Sumatra, whereas in Java, Borneo, and Timor, along with
males quite identical with those of Sumatra, occur females of the
_Polytes_ form, although a single specimen of the true _P. Theseus_,
Cr., taken at Lombock would seem to show that the two forms do occur
together. In the allied species found in the Philippine ♀Islands (_P.
Alphenor_, Cr., _P. Ledebouria_, Eschsch., ♀ _P. Elyros_, G. R. G.)
forms corresponding to these extremes occur along with a number of
intermediate varieties, as shown by a fine series in the British Museum.
We have here an indication of how dimorphism may be produced; for let
the extreme Philippine forms be better suited to their conditions of
existence than the intermediate connecting links, and the latter will
gradually die out, leaving two distinct forms of the same insect, each
adapted to some special conditions. As these conditions are sure to vary
in different districts, it will often happen, as in Sumatra and Java,
that the one form will predominate in the one island, the other in the
adjacent one. In the island of Borneo there seems to be a third form;
for _P. Melanides_, De Haan, evidently belongs to this group, and has
all the chief characteristics of _P. Theseus_, with a modified
coloration of the hind wings. I now come to an insect which, if I am
correct, offers one of the most interesting cases of variation yet
adduced. _Papilio Romulus_, L., a butterfly found over a large part of
India and Ceylon, and not uncommon in collections, has always been
considered a true and independent species, and no suspicions have been
expressed regarding it. But a male of this form does not, I believe,
exist. I have examined the fine series in the British Museum, in the
East India Company’s Museum, in the Hope Museum at Oxford, in Mr.
Hewitson’s and several other private collections, and can find nothing
but females; and for this common butterfly no male partner can be found
except the equally common _P. Pammon_, a species already provided with
two wives, and yet to whom we shall be forced, I believe, to assign a
third. On carefully examining _P. Romulus_, I find that in all essential
characters,—the form and texture of the wings, the length of the
antennæ, the spotting of the head and thorax, and even the peculiar
tints and shades with which it is ornamented,—it corresponds exactly
with the other females of the _Pammon_ group; and though, from the
peculiar marking of the fore wings, it has at first sight a very
different aspect, yet a closer examination shows that every one of its
markings could be produced by slight and almost imperceptible
modifications of the various allied forms. I fully believe, therefore,
that I shall be correct in placing _P. Romulus_ as a third Indian form
of the female _P. Pammon_, corresponding to _P. Melanides_, the third
form of the Malayan _P. Theseus_. I may mention here that the females of
this group have a superficial resemblance to the _Polydorus_ group, as
shown by _P. Theseus_ having been considered to be the female of _P.
Antiphus_, and by _P. Romulus_ being arranged next to _P. Hector_. There
is no close affinity between these two groups of _Papilio_, and I am
disposed to believe that we have here a case of mimicry, brought about
by the same causes which Mr. Bates has so well explained in his account
of Heliconidæ, and which thus led to the singular exuberance of
polymorphic forms in this and allied groups of the genus _Papilio._ I
shall have to devote a section of my paper to the consideration of this
subject.
The third example of polymorphism I have to bring forward is _Papilio
Ormenus_, Guér., which is closely allied to the well-known _P.
Erechtheus_, Don., of Australia. The most common form of the female also
resembles that of _P. Erechtheus_; but a totally different-looking
insect was found by myself in the Aru Islands, and figured by Mr.
Hewitson under the name of _P. Onesimus_, which subsequent observation
has convinced me is a second form of the female of _P. Ormenus_.
Comparison of this with Boisduval’s description of _P. Amanga_, a
specimen of which from New Guinea is in the Paris Museum, shows the
latter to be a closely similar form; and two other specimens were
obtained by myself, one in the island of Goram and the other in Waigiou,
all evidently local modifications of the same form. In each of these
localities males and ordinary females of _P. Ormenus_ were also found.
So far there is no evidence that these light-coloured insects are not
females of a distinct species, the males of which have not been
discovered. But two facts have convinced me this is not the case. At
Dorey, in New Guinea, where males and ordinary females closely allied to
_P. Ormenus_ occur (but which seem to me worthy of being separated as a
distinct species), I found one of these light-coloured females closely
followed in her flight by three males, exactly in the same manner as
occurs (and, I believe, occurs only) with the sexes of the same species.
After watching them a considerable time, I captured the whole of them,
and became satisfied that I had discovered the true relations of this
anomalous form. The next year I had corroborative proof of the
correctness of this opinion by the discovery in the island of Batchian
of a new species allied to _P. Ormenus_, all the females of which,
either seen or captured by me, were of one form, and much more closely
resembling the abnormal light-coloured females of _P. Ormenus_ and _P.
Pandion_ than the ordinary specimens of that sex. Every naturalist will,
I think, agree that this is strongly confirmative of the supposition
that both forms of female are of one species; and when we consider,
further, that in four separate islands, in each of which I resided for
several months, the two forms of female were obtained and only one form
of male ever seen, and that about the same time M. Montrouzier in
Woodlark Island, at the other extremity of New Guinea (where he resided
several years, and must have obtained all the large Lepidoptera of the
island), obtained females closely resembling mine, which, in despair at
finding no appropriate partners for them, he mates with a widely
different species,—it becomes, I think, sufficiently evident that this
is another case of polymorphism of the same nature as those already
pointed out in _P. Pammon_ and _P. Memnon_. This species, however, is
not only _dimorphic_, but _trimorphic_; for, in the island of Waigiou, I
obtained a third female quite distinct from either of the others, and in
some degree intermediate between the ordinary female and the male. The
specimen is particularly interesting to those who believe, with Mr.
Darwin, that extreme difference of the sexes has been gradually produced
by what he terms sexual selection, since it may be supposed to exhibit
one of the intermediate steps in that process which has been
accidentally preserved in company with its more favoured rivals, though
its extreme rarity (only one specimen having been seen to many hundreds
of the other form) would indicate that it may soon become extinct.
The only other case of polymorphism in the genus _Papilio_, at all equal
in interest to those I have now brought forward, occurs in America; and
we have, fortunately, accurate information about it. _Papilio Turnus_,
L., is common over almost the whole of temperate North America; and the
female resembles the male very closely. A totally different-looking
insect both in form and colour, _Papilio Glaucus_, L., inhabits the same
region; and though, down to the time when Boisduval published his
‘Species Général,’ no connexion was supposed to exist between the two
species, it is now well ascertained that _P. Glaucus_ is a second female
form of _P. Turnus_. In the ‘Proceedings of the Entomological Society of
Philadelphia,’ Jan. 1863, Mr. Walsh gives a very interesting account of
the distribution of this species. He tells us that in the New England
States and in New York all the females are yellow, while in Illinois and
further south all are black; in the intermediate region both black and
yellow females occur in varying proportions. Lat. 37° is approximately
the southern limit of the yellow form, and 42° the northern limit of the
black form; and, to render the proof complete, both black and yellow
insects have been bred from a single batch of eggs. He further states
that, out of thousands of specimens, he has never seen or heard of
intermediate varieties between these forms. In this interesting example
we see the effects of latitude in determining the proportions in which
the individuals of each form should exist. The conditions are _here_
favourable to the one form, _there_ to the other; but we are by no means
to suppose that these conditions consist in climate alone. It is highly
probable that the existence of enemies, and of competing forms of life,
may be the main determining influences; and it is much to be wished that
such a competent observer as Mr. Walsh would endeavour to ascertain what
are the adverse causes which are most efficient in keeping down the
numbers of each of these contrasted forms.
Dimorphism of this kind in the animal kingdom does not seem to have any
direct relations to the reproductive powers, as Mr. Darwin has shown to
be the case in plants, nor does it appear to be very general. One other
case only is known to me in another family of my eastern Lepidoptera,
the _Pierulæ_; and but few occur in the Lepidoptera of other countries.
The spring and autumn broods of some European species differ very
remarkably; and this must be considered as a phenomenon of an analogous
though not of an identical nature[3]. _Araschnia prorsa_, of Central
Europe, is a striking example of this alternate or seasonal dimorphism.
Mr. Pascoe has pointed out two forms of the male sex in some species of
Coleoptera belonging to the family Anthribidæ, in seven species of the
two genera _Xenocerus_ and _Mecocerus_ (Proc. Ent. Soc. Lond., 1862, p.
71); and no less than six European Water-beetles, of the genus
_Dytiscus_, have females of two forms, the most common having the elytra
deeply sulcate, the rarer smooth as in the males. The three, and
sometimes four or more, forms under which many Hymenopterous insects
(especially Ants) occur must be considered as a related phenomenon,
though here each form is specialized to a distinct function in the
economy of the species. Among the higher animals, albinoism and melanism
may, as I have already stated, be considered as analogous facts; and I
met with one case of a bird, a species of Lory (_Eos fuscata_, Blyth),
clearly existing under two forms, since I obtained both sexes of each
from a single flock.
Footnote 3:
Among our nocturnal Lepidoptera, I am informed, many analogous cases
occur; and as the whole history of many of these has been investigated
by breeding successive generations from the egg, it is to be hoped
that some of our British Lepidopterists will give us a connected
account of all the abnormal phenomena which they present.
The fact of the two sexes of one species differing very considerably is
so common, that it attracted but little attention till Mr. Darwin showed
how it could in many cases be explained by what he termed sexual
selection. For instance, in most polygamous animals the males fight for
the possession of the females, and the victors, always becoming the
progenitors of the succeeding generation, impress upon their male
offspring their own superior size, strength, or unusually developed
offensive weapons. It is thus that we can account for the spurs and the
superior strength and size of the males in Gallinaceous birds, and also
for the large canine tusks in the males of fruit-eating Apes. So the
superior beauty of plumage and special adornments of the males of so
many birds can be explained by supposing (what there are many facts to
prove) that the females prefer the most beautiful and perfect-plumaged
males, and that thus slight accidental variations of form and colour
have been accumulated till they have produced the wonderful train of the
Peacock and the gorgeous plumage of the Bird of Paradise. Both these
causes have no doubt acted partially in insects, so many species
possessing horns and powerful jaws in the male sex only, and still more
frequently the males alone rejoicing in rich colours or sparkling
lustre. But there is here another cause which has led to sexual
differences, viz. a special adaptation of the sexes to diverse habits or
modes of life. This is well seen in female Butterflies (which are
generally weaker and of slower flight), often having colours better
adapted to concealment; and in certain South American species (_Papilio
torquatus_) the females, which inhabit the forests, resemble the _Æneas_
group, which abound in similar localities, while the males, which
frequent the sunny open riverbanks, have a totally different coloration.
In these cases, therefore, natural selection seems to have acted
independently of sexual selection; and all such cases may be considered
as examples of the simplest dimorphism, since the offspring never offer
intermediate varieties between the parent forms.
The distinctive character therefore of dimorphism is this, that the
union of these distinct forms does not produce intermediate varieties,
but reproduces them unchanged. In simple varieties, on the other hand,
as well as when distinct local forms or distinct species are crossed,
the offspring never resembles either parent exactly, but is more or less
intermediate between them. Dimorphism is thus seen to be a specialized
result of variation, by which new physiological phenomena have been
developed; the two should therefore, whenever possible, be kept
separate[4].
Footnote 4:
The phenomena of _dimorphism_ and _polymorphism_ may be well
illustrated by supposing that a blue-eyed, flaxen-haired Saxon man had
two wives, one a black-haired, red-skinned Indian squaw, the other a
woolly-headed, sooty-skinned negress—and that instead of the children
being mulattoes of brown or dusky tints, mingling the separate
characteristics of their parents in varying degrees, all the boys
should be pure Saxon boys like their father, while the girls should
altogether resemble their mothers. This would be thought a
sufficiently wonderful fact; yet the phenomena here brought forward as
existing in the insect-world are still more extraordinary; for each
mother is capable not only of producing male offspring like the
father, and female like herself, but also of producing other females
exactly like her fellow-wife, and altogether differing from herself.
If an island could be stocked with a colony of human beings having
similar physiological idiosyncrasies with _Papilio Pammon_ or _Papilio
Ormenus_, we should see white men living with yellow, red, and black
women, and their offspring always reproducing the same types; so that
at the end of many generations the men would remain pure white, and
the women of the same well-marked races as at the commencement.
3. _Local form, or variety._—This is the first step in the transition
from variety to species. It occurs in species of wide range, when groups
of individuals have become partially isolated in several points of its
area of distribution, in each of which a characteristic form has become
segregated more or less completely. Such forms are very common in all
parts of the world, and have often been classed as varieties or species
alternately. I restrict the term to those cases where the difference of
the forms is very slight, or where the segregation is more or less
imperfect. The best example in the present group is _Papilio Agamemnon_,
L., a species which ranges over the greater part of tropical Asia, the
whole of the Malay archipelago, and a portion of the Australian and
Pacific regions. The modifications are principally of size and form,
and, though slight, are tolerably constant in each locality. The steps,
however, are so numerous and gradual that it would be impossible to
define many of them, though the extreme forms are sufficiently distinct.
_Papilio Sarpedon_, L., presents somewhat similar but less numerous
variations.
4. _Coexisting variety._—This is a somewhat doubtful case. It is when a
slight but permanent and hereditary modification of form exists in
company with the parent or typical form, without presenting those
intermediate gradations which would constitute it a case of simple
variability. It is evidently only by direct evidence of the two forms
breeding separately that this can be distinguished from _dimorphism_.
The difficulty occurs in _Papilio Jason_, Esp., and _P. Evemon_, Bd.,
which inhabit the same localities, and are almost exactly alike in form,
size, and coloration, except that the latter always wants a very
conspicuous red spot on the under surface, which is found not only in
_P. Jason_, but in all the allied species. It is only by breeding the
two insects that it can be determined whether this is a case of a
coexisting variety or of _dimorphism_. In the former case, however, the
difference being constant and so very conspicuous and easily defined, I
see not how we could escape considering it as a distinct species. A true
case of coexisting forms would, I consider, be produced, if a slight
variety had become fixed as a local form, and afterwards been brought
into contact with the parent species with little or no intermixture of
the two; and such instances do very probably occur.
5. _Race, or subspecies._—These are local forms completely fixed and
isolated; and there is no possible test but individual opinion to
determine which of them shall be considered as species and which
varieties. If stability of form and “_the constant transmission of some
characteristic peculiarity of organization_” is the test of a species
(and I can find no other test that is more certain than individual
opinion), then every one of these fixed races, confined as they almost
always are to distinct and limited areas, must be regarded as a species;
and as such I have in most cases treated them. The various modifications
of _Papilio Ulysses_, _P. Peranthus_, _P. Codrus_, _P. Eurypilus_, _P.
Helenus_, &c., are excellent examples; for while some present great and
well-marked, others offer slight and inconspicuous differences, yet in
all cases these differences seem equally fixed and permanent. If,
therefore, we call some of these forms species, and others varieties, we
introduce a purely arbitrary distinction, and shall never be able to
decide where to draw the line. The races of _Papilio Ulysses_, L., for
example, vary in amount of modification from the scarcely differing New
Guinea form to those of Woodlark Island and New Caledonia, but all seem
equally constant; and as most of these had already been named and
described as species, I have added the New Guinea form under the name of
_P. Penelope_. We thus get a little group of Ulyssine Papilios, the
whole comprised within a very limited area, each one confined to a
separate portion of that area, and, though differing in various amounts,
each apparently constant. Few naturalists will doubt that all these may
and probably have been derived from a common stock; and therefore it
seems desirable that there should be a unity in our method of treating
them: either call them all _varieties_ or all _species_. Varieties,
however, continually get overlooked; in lists of species they are often
altogether unrecorded; and thus we are in danger of neglecting the
interesting phenomena of variation and distribution which they present.
I think it advisable, therefore, to name all such forms; and those who
will not accept them as species may consider them as subspecies or
races.
6. _Species._—Species are merely those strongly marked races or local
forms which, when in contact, do not intermix, and when inhabiting
distinct areas are generally believed to have had a separate origin, and
to be incapable of producing a fertile hybrid offspring. But as the test
of hybridity cannot be applied in one case in ten thousand, and even if
it could be applied, would prove nothing, since it is founded on an
assumption of the very question to be decided—and as the test of
separate origin is in every case inapplicable—and as, further, the test
of non-intermixture is useless, except in those rare cases where the
most closely allied species are found inhabiting the same area, it will
be evident that we have no means whatever of distinguishing so-called
“true species” from the several modes of variation here pointed out, and
into which they so often pass by an insensible gradation. It is quite
true that, in the great majority of cases, what we term “species” are so
well marked and definite that there is no difference of opinion about
them; but as the test of a true theory is, that it accounts for, or at
the very least is not inconsistent with, the whole of the phenomena and
apparent anomalies of the problem to be solved, it is reasonable to ask
that those who deny the origin of species by variation and selection
should grapple with the facts in detail, and show how the doctrine of
the distinct origin and permanence of species will explain and harmonize
them. It has been recently asserted by a high authority that the
difficulty of limiting species is in proportion to our ignorance, and
that just as groups or countries are more accurately known and studied
in greater detail the limits of species become settled[5]. This
statement has, like many other general assertions, its portion of both
truth and error. There is no doubt that many uncertain species, founded
on few or isolated specimens, have had their true nature determined by
the study of a good series of examples: they have been thereby
established as species or as varieties; and the number of times this has
occurred is doubtless very great. But there are other and equally
trustworthy cases in which, not single species, but whole groups have,
by the study of a vast accumulation of materials, been proved to have no
definite specific limits. A few of these must be adduced. In Dr.
Carpenter’s ‘Introduction to the Study of the Foraminifera,’ he states
that “_there is not a single specimen of plant or animal of which the
range of variation has been studied by the collocation and comparison of
so large a number of specimens as have passed under the review of
Messrs. Williamson, Parker, Rupert Jones, and myself, in our studies of
the types of this group_;” and the result of this extended comparison of
specimens is stated to be, “_The range of variation is so great among
the Foraminifera as to include not merely those differential characters
which have been usually accounted_ SPECIFIC, _but also those upon which
the greater part of the_ GENERA _of this group have been founded, and
even in some instances those of its_ ORDERS” (Foraminifera, Preface, x).
Yet this same group had been divided by D’Orbigny and other authors into
a number of clearly defined _families_, _genera_, and _species_, which
these careful and conscientious researches have shown to have been
almost all founded on incomplete knowledge.
Footnote 5:
See Dr. J. E. Gray “On the Species of Lemuroids,” Proc. Zool. Soc.
1863, p. 134.
Professor DeCandolle has recently given the results of an extensive
review of the species of _Cupuliferæ_. He finds that the best-known
species of oaks are those which produce most varieties and subvarieties,
that they are often surrounded by provisional species; and, with the
fullest materials at his command, two-thirds of the species he considers
more or less doubtful. His general conclusion is, that “_in botany the
lowest series of groups_, SUBVARIETIES, VARIETIES, _and_ RACES _are very
badly limited; these can be grouped into_ SPECIES _a little less vaguely
limited, which again can be formed into sufficiently precise_ GENERA.”
This general conclusion is entirely objected to by the writer of the
article in the ‘Natural History Review,’ who, however, does not deny its
applicability to the particular order under discussion, while this very
difference of opinion is another proof that difficulties in the
determination of species do not, any more than in the higher groups,
vanish with increasing materials and more accurate research.
Another striking example of the same kind is seen in the genera _Rubus_
and _Rosa_, adduced by Mr. Darwin himself; for though the amplest
materials exist for a knowledge of these groups, and the most careful
research has been bestowed upon them, yet the various species have not
thereby been accurately limited and defined so as to satisfy the
majority of botanists.
Dr. Hooker seems to have found the same thing in his study of the Arctic
flora. For though he has had much of the accumulated materials of his
predecessors to work upon, he continually expresses himself as unable to
do more than group the numerous and apparently fluctuating forms into
more or less imperfectly defined species[6].
Footnote 6:
In his paper on the “Distribution of Arctic Plants,” Trans. Linn. Soc.
xxiii. p. 310, Dr. Hooker says:—
“The most able and experienced descriptive botanists vary in their
estimate of the value of the ‘specific term’ to a much greater
extent than is generally supposed.”
“I think I may safely affirm that the ‘specific term’ has three
different standard values, all current in descriptive botany, but
each more or less confined to one class of observers.”
“This is no question of what is right or wrong as to the real value
of the specific term; I believe each is right according to the
standard he assumes as the specific.”
Lastly, I will adduce Mr. Bates’s researches on the Amazons. During
eleven years he accumulated vast materials, and carefully studied the
variation and distribution of insects. Yet he has shown that many
species of Lepidoptera, which before offered no special difficulties,
are in reality most intricately combined in a tangled web of affinities,
leading by such gradual steps from the slightest and least stable
variations to fixed races and well-marked species, that it is very often
impossible to draw those sharp dividing-lines which it is supposed that
a careful study and full materials will always enable us to do.
These few examples show, I think, that in every department of nature
there occur instances of the instability of specific form, which the
increase of materials aggravates rather than diminishes. And it must be
remembered that the naturalist is rarely likely to err on the side of
imputing greater indefiniteness to species than really exists. There is
a completeness and satisfaction to the mind in defining and limiting and
naming a species, which leads us all to do so whenever we
conscientiously can, and which we know has led many collectors to reject
vague intermediate forms as destroying the symmetry of their cabinets.
We must therefore consider these cases of excessive variation and
instability as being thoroughly well established; and to the objection
that, after all, these cases are but few compared with those in which
species can be limited and defined, and are therefore merely exceptions
to a general rule, I reply that a true law embraces all apparent
exceptions, and that to the great laws of nature there are no real
exceptions—that what appear to be such are equally results of law, and
are often (perhaps indeed always) those very results which are most
important as revealing the true nature and action of the law. It is for
such reasons that naturalists now look upon the study of _varieties_ as
more important than that of well-fixed species. It is in the former that
we see nature still at work, in the very act of producing those
wonderful modifications of form, that endless variety of colour, and
that complicated harmony of relations, which gratify every sense and
give occupation to every faculty of the true lover of nature.
_Variation as specially influenced by Locality._
The phenomena of variation as influenced by locality have not hitherto
received much attention. Botanists, it is true, are acquainted with the
influences of climate, altitude, and other physical conditions in
modifying the forms and external characteristics of plants; but I am not
aware that any peculiar influence has been traced to locality,
independent of climate. Almost the only case I can find recorded is
mentioned in that repertory of natural-history facts, ‘The Origin of
Species,’ viz. that herbaceous groups have a tendency to become arboreal
in islands. In the animal world, I cannot find that any facts have been
pointed out as showing the special influence of locality in giving a
peculiar _facies_ to the several disconnected species that inhabit it.
What I have to adduce on this matter will therefore, I hope, possess
some interest and novelty.
On examining the closely allied species, local forms, and varieties
distributed over the Indian and Malayan regions, I find that larger or
smaller districts, or even single islands, give a special character to
the majority of their Papilionidæ. For instance: 1. The species of the
Indian region (Sumatra, Java, and Borneo) are almost invariably smaller
than the allied species inhabiting Celebes and the Moluccas; 2. The
species of New Guinea and Australia are also, though in a less degree,
smaller than the nearest species or varieties of the Moluccas; 3. In the
Moluccas themselves the species of Amboyna are the largest; 4. The
species of Celebes equal or even surpass in size those of Amboyna; 5.
The species and varieties of Celebes possess a striking character in the
form of the anterior wings, different from that of the allied species
and varieties of all the surrounding islands; 6. Tailed species in India
or the Indian region become tailless as they spread eastward through the
archipelago.
Having preserved the finest and largest specimens of Butterflies in my
own collection, and having always taken for comparison the largest
specimens of the same sex, I believe that the tables I now give are
sufficiently exact. The differences of expanse of wings are in most
cases very great, and are much more conspicuous in the specimens
themselves than on paper. It will be seen that no less than fourteen
Papilionidæ inhabiting Celebes and the Moluccas are from one-third to
one-half greater in extent of wing than the allied species representing
them in Java, Sumatra, and Borneo. Six species inhabiting Amboyna are
larger than the closely allied forms of the northern Moluccas and New
Guinea by about one-sixth. These include almost every case in which
closely allied species can be compared.
PAPILIONIDÆ.
Species of the Moluccas and Celebes│Closely allied species of Java and
(large). │ the Indian region (small).
Expanse. │ Expanse.
inches. │ inches.
Ornithoptera Helena 7·6│O. Pompeus 5·8
(Amboyna) │
„ „ │O. Amphrisius 6·0
───────────────────────────────────┼───────────────────────────────────
Papilio Macedon 5·8│P. Peranthus 3·8
(Celebes) │
P. Philippus (Moluccas) 4·8│ „ „
───────────────────────────────────┼───────────────────────────────────
P. Blumei (Celebes) 5·4│P. Brama 4·0
P. Alphenor (Celebes) 4·8│P. Theseus 3·6
P. Gigon (Celebes) 5·4│P. Demolion 4·0
P. Deucalion (Celebes) 4·6│P. Macareus 3·7
P. Agamemnon, var. 4·4│P. Agamemnon, var. 3·8
(Celebes) │
───────────────────────────────────┼───────────────────────────────────
P. Eurypilus (Moluccas) 4·0│P. Jason 3·4
P. Telephus (Celebes) 4·3│ „ „
───────────────────────────────────┼───────────────────────────────────
P. Ægisthus (Moluccas) 4·4│P. Rama 3·2
P. Miletus (Celebes) 4·4│P. Sarpedon 3·8
P. Androcles (Celebes) 4·8│P. Antiphates 3·7
P. Polyphontes 4·6│P. Diphilus 3·9
(Celebes) │
Leptocircus Curtius 2·0│L. Meges 1·8
(Celebes) │
│
Species inhabiting Amboyna (large).│ Allied species of New Guinea and
│ the North Moluccas (smaller).
Papilio Ulysses 6·1│P. Penelope 5·2
„ „ │P. Telegonus 4·0
───────────────────────────────────┼───────────────────────────────────
P. Polydorus 4·9│P. Leodamas 4·0
P. Deiphobus 6·8│P. Deiphontes 5·8
───────────────────────────────────┼───────────────────────────────────
P. Gambrisius 6·4│P. Ormenus 5·6
„ „ │P. Tydeus 6·0
───────────────────────────────────┼───────────────────────────────────
P. Codrus 5·1│P. Codrus, var. 4·3
│ _papuensis_
Ornithoptera Priamus, ♂ 8·0│Orn. Poseidon, ♂ 7·0
The differences of form are equally clear.
_Papilio Pammon_ everywhere on the continent is tailed in both sexes. In
Java, Sumatra, and Borneo, the closely allied _P. Theseus_ has a very
short tail, or tooth only, in the male, while in the females the tail is
retained. Further east, in Celebes and the South Moluccas, the hardly
separable _P. Alphenor_ has quite lost the tail in the male, while the
female retains it, but in a narrower and less spatulate form. A little
further, in Gilolo, _P. Nicanor_ has completely lost the tail in both
sexes.
_Papilio Agamemnon_ exhibits a somewhat similar series of changes. In
India it is always tailed; in the greater part of the archipelago it has
a very short tail; while far east, in New Guinea and the adjacent
islands, the tail has almost entirely disappeared.
In the _Polydorus_-group two species, _P. Antiphus_ and _P. Diphilus_,
inhabiting India and the Indian region, are tailed, while the two which
take their place in the Moluccas, New Guinea, and Australia, _P.
Polydorus_ and _P. Leodamas_, are destitute of tail, the species
furthest east having lost this ornament the most completely.
Western species, tailed. Eastern species (closely allied),
less tailed.
Papilio Pammon (India) tailed. P. Thesus (islands) very short
tail.
P. Agamemnon, var. tailed. P. Agamemnon, var. not tailed.
(India) (islands)
P. Antiphus (India, tailed. P. Polydorus (Moluccas) not tailed.
Java)
P. Diphilus (India, tailed. P. Leodamas (New not tailed.
Java) Guinea)
The most conspicuous instance of local modification of form, however, is
exhibited in the island of Celebes, which in this respect, as in some
others, stands alone and isolated in the whole archipelago. Almost every
species of _Papilio_ inhabiting Celebes has the wings of a peculiar
shape, which distinguishes them at a glance from the allied species of
every other island. This peculiarity consists, first, in the upper wings
being generally more elongate and falcate; and secondly, in the costa or
anterior margin being much more curved, and in most instances exhibiting
near the base an abrupt bend or elbow, which in some species is very
conspicuous. This peculiarity is visible, not only when the Celebesian
species are compared with their small-sized allies of Java and Borneo,
but also, and in an almost equal degree, when the large forms of Amboyna
and the Moluccas are the objects of comparison, showing that this is
quite a distinct phenomenon from the difference of size which has just
been pointed out.
In the following Table I have arranged the chief Papilios of Celebes in
the order in which they exhibit this characteristic form most
prominently. (See Plate VIII.)
Papilios of Celebes, having the Closely allied Papilios of the
wings falcate or with abruptly surrounding islands, with less
curved costa. falcate wings and slightly curved
costa.
1. P. Gigon, n. s. P. Demolion (Java).
2. P. Telephus, n. s. P. Jason (Sumatra).
3. P. Miletus, n. s. P. Sarpedon (Moluccas, Java).
4. P. Agamemnon, var. P. Agamemnon, var. (Borneo).
5. P. Macedon, n. s. P. Peranthus (Java).
6. P. Ascalaphus. P. Deiphontes, n. s. (Gilolo).
7. P. Hecuba, n. s. P. Helenus (Java).
8. P. Blumei. P. Brama (Sumatra).
9. P. Androcles. P. Antiphates (Borneo).
10. P. Rhesus. P. Aristæus (Moluccas).
11. P. Theseus, var., ♂. P. Thesus, ♂ (Java).
12. P. Codrus, var. P. Codrus (Moluccas).
13. P. Encelades. P. Leucothoë (Malacca).
It thus appears that every species of _Papilio_ exhibits this peculiar
form in a greater or less degree, except one, _P. Polyphontes_, Bd.,
allied to _P. Diphilus_ of India and _P. Polydorus_ of the Moluccas.
This fact I shall recur to again, as I think it helps us to understand
something of the causes that may have brought about the phenomenon we
are considering. Neither do the genera _Ornithoptera_ and _Leptocircus_
exhibit any traces of this peculiar form. In several other families of
Butterflies this characteristic form reappears in a few species. In the
Pieridæ the following species exhibit it distinctly:—
1. Eronia tritæa │compared with Eronia Valeria (Java).
2. Iphias Glaucippe, var.│ „ „ Iphias Glaucippe (Java).
3. Pieris Zebuda │ „ „ Pieris Descombesi (India).
4. P. Zarinda │ „ „ P. Nero (Malacca).
5. P., n. s. │ „ „ P. Hyparete (Java).
──────────────────────────┬────────────────────────────────────────────
6. P. Hombronii │ have the same form, but are isolated
│ species.
7. P. Ithome │ „
──────────────────────────┼────────────────────────────────────────────
8. P. Eperia, _Bd._ │compared with P. Coronis (Java).
9. P. Polisma │ „ „ P., n. s. (Malacca).
10. Terias, n. s. │ „ „ P. Tilaha (Java).
The other species of _Terias_, one or two _Pieris_, and the genus
_Callidryas_ do not exhibit any perceptible change of form.
In the other families there are but few similar examples. The following
are all that I can find in my collection:—
Cethosia Æole compared with Cethosia Biblis (Java).
Junonia, n. s. „ „ Junonia Polynice (Borneo).
Limenitis Limire „ „ Limenitis Procris (Java).
Cynthia Arsinoë, var. „ „ Cynthia Arsinoë (Java, Sum.,
Born.).
All these belong to the family of the Nymphalidæ. Many other genera of
this family, as _Diadema_, _Adolias_, _Charaxes_, and _Cyrestis_, as
well as the entire families of the Danaidæ, Satyridæ, Lycænidæ, and
Hesperidæ, present no examples of this peculiar form of the upper wing
in the Celebesian species.
The facts now brought forward seem to me of the highest interest. We see
that almost all the species in two important families of the Lepidoptera
(Papilionidæ and Pieridæ) acquire, in a single island, a characteristic
modification of form distinguishing them from the allied species and
varieties of all the surrounding islands. In other equally extensive
families no such change occurs, except in one or two isolated species.
However we may account for these phenomena, or whether we may be quite
unable to account for them, they furnish, in my opinion, a strong
corroborative testimony in favour of the doctrine of the origin of
species by successive small variations; for we have here slight
varieties, local races, and undoubted species, all modified in exactly
the same manner, indicating plainly a common cause producing identical
results. On the generally received theory of the original distinctness
and permanence of species, we are met by this difficulty: one portion of
these curiously modified forms are admitted to have been produced by
variation and some natural action of local conditions; whilst the other
portion, differing from the former only in degree, and connected with
them by insensible gradations, are said to have possessed this
peculiarity of form at their first creation, or to have derived it from
unknown causes of a totally distinct nature. Is not the _à priori_
evidence in favour of the assumption of an identity of the causes that
have produced such similar results? and have we not a right to call upon
our opponents for some proofs of their own doctrine, and for an
explanation of its difficulties, instead of their assuming that they are
right, and laying upon us the burthen of disproof?
Let us now see if the facts in question do not themselves furnish some
clue to their own explanation. Mr. Bates has shown that certain groups
of butterflies have a defence against insectivorous animals,
independent of swiftness of motion. These are generally very abundant,
slow, and weak fliers, and are more or less the objects of mimicry by
other groups, which thus gain an advantage in a freedom from
persecution similar to that enjoyed by those they resemble. Now the
only Papilios which have not in Celebes acquired the peculiar form of
wing belong to a group which is imitated both by other species of
_Papilio_ and by Moths of the genus _Epicopeia_, West. This group is
of weak and slow flight; and we may therefore fairly conclude that it
possesses some means of defence (probably in a peculiar odour or
taste) which saves it from attack. Now the arched costa and falcate
form of wing is generally supposed to give increased powers of flight,
or, as seems to me more probable, greater facility in making sudden
turnings, and thus baffling a pursuer. But the members of the
_Polydorus_-group (to which belongs the only unchanged Celebesian
_Papilio_), being already guarded against attack, have no need of this
increased power of wing; and “natural selection” would therefore have
no tendency to produce it. The whole family of Danaidæ are in the same
position: they are slow and weak fliers; yet they abound in species
and individuals, and are the objects of mimicry. The Satyridæ have
also probably a means of protection—perhaps their keeping always near
the ground and their generally obscure colours; while the Lycænidæ and
Hesperidæ may find security in their small size and rapid motions. In
the extensive family of the Nymphalidæ, however, we find that several
of the larger species, of comparatively feeble structure, have their
wings modified (_Cethosia_, _Limenitis_, _Junonia_, _Cynthia_), while
the large-bodied powerful species, which have all an excessively rapid
flight, have exactly the same form of wing in Celebes as in the other
islands. On the whole, therefore, we may say that all the butterflies
of rather large size, conspicuous colours, and not very swift flight
have been affected in the manner described, while the smaller-sized
and obscure groups, as well as those which are the objects of mimicry,
and also those of exceedingly swift flight, have remained unaffected.
It would thus appear as if there must be (or once have been) in the
island of Celebes, some peculiar enemy to these larger-sized butterflies
which does not exist, or is less abundant, in the surrounding islands.
Increased powers of flight, or rapidity of turning, was advantageous in
baffling this enemy; and the peculiar form of wing necessary to give
this would be readily acquired by the action of “natural selection” on
the slight variations of form that are continually occurring. Such an
enemy one would naturally suppose to be an insectivorous bird; but it is
a remarkable fact that most of the genera of Fly-catchers of Borneo and
Java on the one side (_Muscipeta_, _Philentoma_), and of the Moluccas on
the other (_Monarcha_, _Rhipidura_), are almost entirely absent from
Celebes. Their place seems to be supplied by the Caterpillar-catchers
(_Graucalus_, _Campephaga_), of which six or seven species are known
from Celebes and are very numerous in individuals. We have no positive
evidence that these birds pursue butterflies on the wing, but it is
highly probable that they do so when other food is scarce[7]. However
this may be, the fauna of Celebes is undoubtedly highly peculiar in
every department of which we have any knowledge; and though we may not
be able to trace it satisfactorily, there can, I think, be little doubt
that the singular modification in the wings of so many of the
butterflies of that island is an effect of that complicated action and
reaction of all living things upon each other in the struggle for
existence, which continually tends to readjust disturbed relations, and
to bring every species into harmony with the varying conditions of the
surrounding universe.
Footnote 7:
Mr. Bates has suggested that the larger Dragon-flies (_Æshna_, &c.)
prey upon butterflies; but I did not notice that they were more
abundant in Celebes than elsewhere.
But even the conjectural explanation now given fails us in the other
cases of local modification. Why the species of the western islands
should be smaller than those further east,—why those of Amboyna should
exceed in size those of Gilolo and New Guinea—why the tailed species of
India should begin to lose that appendage in the islands, and retain no
trace of it on the borders of the Pacific, are questions which we cannot
at present attempt to answer. That they depend, however, on some general
principle is certain, because analogous facts have been observed in
other parts of the world. Mr. Bates informs me that, in three distinct
groups, Papilios which on the Upper Amazon and in most other parts of
South America have spotless upper wings obtain pale or white spots at
Para and on the Lower Amazon; and also that the _Æneas_-group of
Papilios never have tails in the equatorial regions and the Amazons
valley, but gradually acquire tails in many cases as they range towards
the northern or southern tropic. Even in Europe we have somewhat similar
facts; for the species and varieties of butterflies peculiar to the
island of Sardinia are generally smaller and more deeply coloured than
those of the mainland, and _Papilio Hospiton_ has lost the tail, which
is a prominent feature of the closely allied _P. Machaon_.
Facts of a similar nature to those now brought forward would no doubt be
found to occur in other groups of insects, were local faunas carefully
studied in relation to those of the surrounding countries; and they seem
to indicate that climate and other physical causes have, in some cases,
a very powerful effect in modifying specific form, and thus directly aid
in producing the endless variety of nature.
I may state that I can adduce facts perfectly analogous to these from
other families of Lepidoptera, especially the Danaidæ; but as the
greater part of the species are still undescribed, I can only now assert
that similar phenomena do occur there.
_Mimicry._
I need scarcely say that I entirely agree with Mr. Bates’s explanation
of the causes which have led to one group of insects mimicking another
(Trans. Linn. Soc. vol. xxiii. p. 495). I have, therefore, only now to
adduce such illustrations of this curious phenomenon as are furnished by
the Eastern Papilionidæ, and to show their hearing upon the phenomena of
variation already mentioned. As in America, so in the Old World, species
of Danaidæ are the objects which the other families most often imitate.
But, besides these, some genera of Morphidæ and one section of the genus
_Papilio_ are also less frequently copied. Many species of _Papilio_
mimic other species of these three groups so closely that they are
undistinguishable when on the wing; and in every case the pairs which
resemble each other inhabit the same locality.
The following list exhibits the most important and best-marked cases of
mimicry which occur among the Papilionidæ of the Malayan region and
India:—
Mimickers[8]. Species mimicked. │Common habitat.
DANAIDÆ.
1. Papilio paradoxa, Euplœa Midamus, _Cr._, ♂│Sumatra, &c.
_Zink._, ♂ │
—— ——, ♀ —— ——, ♀ │ „
──────────────────────────────────────────────────────┼────────────────
2. —— ——, _West._ E. Rhadamanthus │Sumatra, &c.
3. P. Caunus, E., sp. │Borneo.
4. P. Thule, _Wall._ Danais sobrina, _Bd._ │New Guinea.
5. P. Macareus, _Godt._ D. Aglaia, _Cr._ │Malacca, Java.
6. P. Agestor, _G. R. G._ D. Tytia, _G. R. G._ │Northern India.
7. P. idæoides, _Hewits._ Hestia Leuconoë, │Philippines.
_Erichs._ │
8. P. Delessertii, _Guér._ Hestia, sp. │Penang.
│
MORPHIDÆ.
9. P. Pandion, _Wall._, ♀ Drusilla bioculata, │New Guinea.
_Guér._ │
│
PAPILIO (POLYDORUS- and COON-groups).
10. P. Pammon, _L._ (Romulus, Papilio Hector, _L._ │India.
_L._), ♀ │
11. P. Theseus, _Cr._, var., P. Antiphus, _Fab._ │Sumatra, Borneo.
♀ │
12. P. Theseus, _Cr._, var., P. Diphilus, _Esp._ │Sumatra, Java.
♀ │
13. P. Memnon, var. P. Coon, _Fab._ │Sumatra.
_Achates_, ♀ │
14. P. Androgeus, var. P. Doubledayi, _Wall._ │Northern India.
_Achates_, ♀ │
15. P. Œnomaus, _God._, ♀ P. Liris, _God._ │Timor.
Footnote 8:
The terms “_mimicry_” and “_mimickers_” have been objected to on the
ground that they imply _voluntary action_ on the part of the insects.
This appears to me of little importance compared with the advantages
of convenience, flexibility, and expressiveness which they undoubtedly
possess, especially as the whole theory propounded by the originator
of the term in this sense excludes all idea of voluntary action. The
only approximately synonymous words, not implying will, are
_resemblance_, _similarity_, and _likeness_; and it is evident that
none of these can be applied intelligibly under the variety of forms
required, and to which Mr. Bates’s expression so readily lends itself
in the terms _mimic_, _mimickers_, _mimicry_, _mimicked_. Add to this
the inconvenience of changing a term which, from the interest and wide
discussion of the subject, must be already very generally understood,
and I think it will be admitted that nothing would be gained by
altering it, even if a better word were pointed out, which has not yet
been done.
We have therefore fifteen species or marked varieties of _Papilio_ which
so closely resemble species of other groups in their respective
localities, that it is not possible to impute the resemblance to
accident. The first two in the list (_Papilio paradoxa_ and _P. Caunus_)
are so exactly like _Euplœa Midamus_ and _E. Rhadamanthus_ on the wing,
that, although they fly very slowly, I was quite unable to distinguish
them. The first is a very interesting case, because the male and female
differ considerably, and each mimics the corresponding sex of the
_Euplœa_. A new species of _Papilio_ which I discovered in New Guinea
resembles _Danais sobrina_, Bd., from the same country, just as _Papilio
Macareus_ resembles _Danais Aglaia_ in Malacca, and (according to Dr.
Horsfield’s figure) still more closely in Java. The Indian _Papilio
Agestor_ closely imitates _Danais Tytia_, which has quite a different
style of colouring from the preceding; and the extraordinary _Papilio
idæoides_ from the Philippine Islands must, when on the wing, perfectly
resemble the _Hestia Leuconoë_ of the same region, as also does the _P.
Delessertii_, Guér., imitate an undescribed species of _Hestia_ from
Penang. Now in every one of these cases the Papilios are very scarce,
while the Danaidæ which they resemble are exceedingly abundant—most of
them swarming so as to be a positive nuisance to the collecting
entomologist by continually hovering before him when he is in search of
newer and more varied captures. Every garden, every roadside, the
suburbs of every village are full of them, indicating very clearly that
their life is an easy one, and that they are free from persecution by
the foes which keep down the population of less favoured races. This
superabundant population has been shown by Mr. Bates to be a general
characteristic of all American groups and species which are objects of
mimicry; and it is interesting to find his observations confirmed by
examples on the other side of the globe.
The remarkable genus _Drusilla_, a group of pale-coloured butterflies,
more or less adorned with ocellate spots, is also the object of mimicry
by three distinct genera (_Melanitis_, _Hyantis_, and _Papilio_). These
insects, like the _Danaidæ_, are abundant in individuals, have a very
weak and slow flight, and do not seek concealment, or appear to have any
means of protection from insectivorous creatures. It is natural to
conclude, therefore, that they have some hidden property which saves
them from attack; and it is easy to see that when any other insects, by
what we call accidental variation, come more or less remotely to
resemble them, the latter will share to some extent in their immunity.
An extraordinary dimorphic form of a female _Papilio_ has come to
resemble the Drusillas sufficiently to be taken for one of that group at
a little distance; and it is curious that I captured one of these
Papilios in the Aru Islands hovering along the ground, and settling on
it occasionally, just as it is the habit of the Drusillas to do. The
resemblance in this case is only general; but this form of _Papilio_
varies much, and there is therefore material for natural selection to
act upon so as ultimately to produce a copy as exact as in the other
cases.
The eastern Papilios allied to _Polydorus Coon_ and _P. Philoxenus_,
form a natural section of the genus resembling, in many respects, the
_Æneas_-group of South America, which they may be said to represent in
the East. Like them, they are forest insects, have a low and weak
flight, and in their favourite localities are rather abundant in
individuals; and like them, too, they are the objects of mimicry. We may
conclude, therefore, that they possess some hidden means of protection,
which makes it useful to other insects to be mistaken for them.
The Papilios which resemble them belong to a very distinct section of
the genus, in which the sexes differ greatly; and it is those females
only which differ most from the males, and which have already been
alluded to as exhibiting instances of dimorphism, which resemble species
of the other group.
The resemblance of _P. Romulus_ to _P. Hector_ is, in some specimens,
very considerable, and has led to the two species being placed to follow
each other in the British Museum Catalogues and by Mr. E. Doubleday. I
have shown, however, that _P. Romulus_ is probably a dimorphic form of
the female _P. Pammon_, and belongs to a distinct section of the
genus[9].
Footnote 9:
See Plate II. fig. 6.
The next pair, _P. Theseus_, Cr., and _P. Antiphus_, Fab., have been
united as one species both by De Haan and in the British Museum
Catalogues. The ordinary variety of _P. Theseus_ found in Java almost as
nearly resembles _P. Diphilus_, Esp., of the same country. The most
interesting case, however, is the extreme female form of _P. Memnon_
(_P. Achates_, Cr.)[10], which has acquired the general form and
markings of _P. Coon_, an insect which differs from the ordinary male
_P. Memnon_, as much as any two species differ which can be chosen in
this extensive and highly varied genus; and, as if to show that this
resemblance is not accidental, but is the result of law, when in India
we find a species closely allied to _P. Coon_, but with red instead of
yellow spots (_P. Doubledayi_, Wall.), the corresponding variety of _P.
Androgeus_ (_P. Achates_, Cram., 182, A, B,) has acquired exactly the
same peculiarity of having red spots instead of yellow. Lastly, in the
island of Timor, the female of _P. Œnomaus_ (a species allied to _P.
Memnon_) resembles so closely _P. Liris_ (one of the _Polydorus_-group),
that the two, which were often seen flying together, could only be
distinguished by a minute comparison after being captured.
Footnote 10:
See Plate I. fig. 4.
The last six cases of mimicry are especially instructive, because they
seem to indicate one of the processes by which dimorphic forms have been
produced. When, as in these cases, one sex differs much from the other,
and varies greatly itself, it may happen that occasionally individual
variations will occur having a distant resemblance to groups which are
the objects of mimicry, and which it is therefore advantageous to
resemble. Such a variety will have a better chance of preservation; the
individuals possessing it will be multiplied; and their accidental
likeness to the favoured group will be rendered permanent by hereditary
transmission, and, each successive variation which increases the
resemblance being preserved, and all variations departing from the
favoured type having less chance of preservation, there will in time
result those singular cases of two or more isolated and fixed forms
bound together by that intimate relationship which constitutes them the
sexes of a single species. The reason why the females are more subject
to this kind of modification than the males is, probably, that their
slower flight, when laden with eggs, and their exposure to attack while
in the act of depositing their eggs upon leaves, render it especially
advantageous for them to have some additional protection. This they at
once obtain by acquiring a resemblance to other species which, from
whatever cause, enjoy a comparative immunity from persecution.
This summary of the more interesting phenomena of variation presented by
the eastern Papilionidæ is, I think, sufficient to substantiate my
position, that the Lepidoptera are a group that offer especial
facilities for such inquiries; and it will also show that they have
undergone an amount of special adaptive modification rarely equalled
among the more highly organized animals. And, among the Lepidoptera, the
great and pre-eminently tropical families of Papilionidæ and Danaidæ
seem to be those in which complicated adaptations to the surrounding
organic and inorganic universe have been most completely developed,
offering in this respect a striking analogy to the equally
extraordinary, though totally different, adaptations which present
themselves in the _Orchideæ_, the only family of plants in which mimicry
of other organisms appears to play any important part, and the only one
in which striking cases of polymorphism occur; for such we must consider
to be the male, female, and hermaphrodite forms of _Catasetum
tridentatum_, which differ so greatly in form and structure that they
were long considered to belong to three distinct genera.
_Arrangement and Geographical Distribution of the Malayan_ Papilionidæ.
Although the species of Papilionidæ inhabiting the Malayan region are
very numerous, they all belong to three out of the nine genera into
which the family is divided. One of the remaining genera (_Eurycus_) is
restricted to Australia, and another (_Teinopalpus_) to the Himalayan
Mountains, while no less than four (_Parnassius_, _Doritis_, _Thais_,
and _Sericinus_) are confined to Southern Europe and to the
mountain-ranges of the Palæarctic region.
The genera _Ornithoptera_ and _Leptocircus_ are highly characteristic of
Malayan entomology, but are uniform in character and of small extent.
The genus _Papilio_, on the other hand, presents a great variety of
forms, and is so richly represented in the Malay islands, that more than
one-fourth of all the known species are found there. It becomes
necessary, therefore, to divide this genus into natural groups before we
can successfully study its geographical distribution.
Owing principally to Dr. Horsfield’s observations in Java, we are
acquainted with a considerable number of the larvæ of Papilios; and
these furnish good characters for the primary division of the genus into
natural groups. The manner in which the hinder wings are plaited or
folded back at the abdominal margin, the size of the anal valves, the
structure of the antennæ, and the form of the wings are also of much
service, as well as the character of the flight and the style of
coloration. Using these characters, I divide the Malayan Papilios into
four sections, and seventeen groups, as follows:—
Genus ORNITHOPTERA.
a. _Priamus_-group. Black and green.
b. _Pompeus_-group. Black and yellow.
c. _Brookeanus_-group.
Genus PAPILIO.
A. Larvæ short, thick, with numerous fleshy tubercles; purplish.
a. _Nox_-group. Abdominal fold in ♂ very large; anal valves small,
but swollen; antennæ moderate; wings entire, or tailed:
includes the Indian _Philoxenus_-group.
b. _Coon_-group. Abdominal fold in ♂ small; anal valves small, but
swollen; antennæ moderate; wings tailed.
c. _Polydorus_-group. Abdominal fold in ♂ small, or none; anal
valves small or obsolete, hairy; wings tailed or entire.
B. Larvæ with third segment swollen, transversely or obliquely
banded; pupa much bent. Imago with abdominal margin in ♂
plaited, but not reflexed; body weak; antennæ long; wings much
dilated, often tailed.
d. _Ulysses_-group.
e. _Peranthus_-group.
_Protenor_-group (Indian) is somewhat intermediate between
these, and is nearest to the _Nox_-group.
f. _Memnon_-group.
g. _Helenus_-group.
h. _Erectheus_-group.
i. _Pammon_-group.
k. _Demolion_-group.
C. Larvæ subcylindrical, variously coloured. Imago with abdominal
margin in ♂ plaited, but not reflexed; body weak; antennæ short,
with a thick curved club; wings entire.
l. _Erithonius_-group. Sexes alike, larva and pupa something like
those of _P. Demolion_.
m. _Paradoxa_-group. Sexes different.
n. _Dissimilis_-group. Sexes alike; larva bright-coloured; pupa
straight, cylindric.
D. Larvæ elongate, attenuate behind, and often bifid, with lateral
and oblique pale stripes, green. Imago with the abdominal margin
in ♂ reflexed, woolly or hairy within; anal valves small, hairy;
antennæ short, stout; body stout.
o. _Macareus_-group. Hind wings entire.
p. _Antiphates_-group. Hind wings much tailed (swallow-tails).
q. _Eurypylus_-group. Hind wings elongate or tailed.
Genus LEPTOCIRCUS.
making, in all, twenty distinct groups of Malayan Papilionidæ.
The first section of the genus _Papilio_ (A) comprises insects which,
though differing considerably in structure, have much general
resemblance. They all have a weak, low flight, frequent the most
luxuriant forest districts, seem to love the shade, and are the objects
of mimicry by other Papilios.
Section B consists of weak-bodied, large-winged insects, with an
irregular wavering flight, and which, when resting on foliage, often
expand the wings, which the species of the other sections rarely or
never do. They are the most conspicuous and striking of eastern
Butterflies.
Section C consists of much weaker and slower-flying insects, often
resembling in their flight, as well as in their colours, species of
Danaidæ.
Section D contains the strongest-bodied and most swift-flying of the
genus. They love sunlight, and frequent the borders of streams and the
edges of puddles, where they gather together in swarms consisting of
several species, greedily sucking up the moisture, and, when disturbed,
circling round in the air, or flying high and with great strength and
rapidity.
In the following Table I have arranged all the Malayan Papilionidæ in
what appears to me their most natural succession, and have exhibited
their distribution in twenty-one columns of localities, extending from
the Malay peninsula, on the north-west, to Woodlark Island, near New
Guinea, on the south-east. The double line divides the Indo-Malayan from
the Austro-Malayan region; and those islands which form natural
zoological groups are connected by brackets.
_Table showing the Distribution of the Malayan_ Papilionidæ.
┌───┬──┬───────────────────╥─────────────────────────────────────────────┐
│ │ │ ║ INDO-MALAYAN REGION. │
├───┼──┼───────────────────╫────────┬────────┬────────┬─────╥────────────┤
│ │ │ =Ornithoptera.= ║Malacca.│Sumatra.│Borneo. │Java.║Philippines.│
│ │ │ ║ │ │ │ ║ │
├───┼──┼───────────────────╫────────┬────────┬────────┬─────╥────────────┤
│ │a.│_Priamus_-group. ║ │ │ │ ║ │
│ 1│ │Priamus, _L._ ║ │ │ │ ║ │
│ 2│ │Poseidon, _Db._ ║ │ │ │ ║ │
│ 3│ │Crœsus, _Feld._ ║ │ │ │ ║ │
│ 4│ │Tithonus, _De ║ │ │ │ ║ │
│ │ │ Haan._ ║ │ │ │ ║ │
│ 5│ │Urvilliana, _Guér._║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │b.│_Pompeus_-group. ║ │ │ │ ║ │
│ 6│ │Remus, _Cr._ ║ │ │ │ ║ │
│ 7│ │Helena, _L._ ║ │ │ │ ║ │
│ 8│ │Leda, _Wall._ ║ │ │ │ ║ │
│ 9│ │Pompeus, _Cr._ ║ 1│ 1│ 1│ 1║ │
│ 10│ │Nephereus, _G. R. ║ │ │ │ ║ 1│
│ │ │ G._ ║ │ │ │ ║ │
│ 11│ │Magellanus, _Feld._║ │ │ │ ║ 1│
│ 12│ │Criton, _Feld._ ║ │ │ │ ║ │
│ 13│ │Plato, _Wall._ ║ │ │ │ ║ │
│ 14│ │Haliphron, _Bd._ ║ │ │ │ ║ │
│ 15│ │Amphrisius, _Cr._ ║ 1│ │ 1│ 1║ │
│ │ │ ║ │ │ │ ║ │
│ │c.│_Brookeana_-group. ║ │ │ │ ║ │
│ 16│ │Brookeana, _Wall._ ║ │ 1│ 1│ ║ │
│ │ │ ║ │ │ │ ║ │
│ │ │ =Papilio.= ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ A.│a.│_Nox_-group. ║ │ │ │ ║ │
│ 17│ │Nox, _Sw._ ║ 1│ │ │ 1║ │
│ 18│ │Noctis, _Hew._ ║ │ │ 1│ ║ │
│ 19│ │Erebus, _Wall._ ║ 1│ │ 1│ ║ │
│ 20│ │Varuna, _White_ ║ 1│ │ │ ║ │
│ 21│ │Semperi, _Feld._ ║ │ │ │ ║ 1│
│ │ │ ║ │ │ │ ║ │
│ │b.│_Coon_-group. ║ │ │ │ ║ │
│ 22│ │Neptunus, _Guér._ ║ 1│ │ 1│ ║ │
│ 23│ │Coon, _Fab._ ║ │ 1│ 1│ 1║ │
│ │ │ ║ │ │ │ ║ │
│ │c.│_Polydorus_-group. ║ │ │ │ ║ │
│ 24│ │Polydorus, _L._ ║ │ │ │ ║ │
│ 25│ │Leodamas, _Wall._ ║ │ │ │ ║ │
│ 26│ │Diphilus, _Esper_ ║ 1│ │ │ 1║ 1│
│ 27│ │Antiphus, _Fab._ ║ │ 1│ 1│ 1║ 1│
│ 28│ │Polyphontes, _Bd._ ║ │ │ │ ║ │
│ 29│ │Annæ, _Feld._ ║ │ │ │ ║ 1│
│ 30│ │Liris, _Godt._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ B.│d.│_Ulysses_-group. ║ │ │ │ ║ │
│ 31│ │Ulysses, _L._ ║ │ │ │ ║ │
│ 32│ │Penelope, _Wall._ ║ │ │ │ ║ │
│ 33│ │Telegonus, _Feld._ ║ │ │ │ ║ │
│ 34│ │Telemachus, _Mont._║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │e.│_Peranthus_-group. ║ │ │ │ ║ │
│ 35│ │Peranthus, _Fab._ ║ │ │ │ 1║ │
│ 36│ │Pericles, _Wall._ ║ │ │ │ ║ │
│ 37│ │Philippus, _Wall._ ║ │ │ │ ║ │
│ 38│ │Macedon, _Wall._ ║ │ │ │ ║ │
│ 39│ │Brama, _Guér._ ║ 1│ 1│ │ ║ │
│ 40│ │Dædalus, _Feld._ ║ │ │ │ ║ 1│
│ 41│ │Blumei, _Bd._ ║ │ │ │ ║ │
│ 42│ │Arjuna, _Horsf._ ║ │ 1│ 1│ 1║ │
│ │ │ ║ │ │ │ ║ │
│ │f.│_Memnon_-group. ║ │ │ │ ║ │
│ 43│ │Memnon, _L._ ║ │ 1│ 1│ 1║ │
│ 44│ │Androgeus, _Cr._ ║ 1│ │ │ ║ │
│ 45│ │Lampsacus, _Bd._ ║ │ │ │ 1║ │
│ 46│ │Priapus, _Bd._ ║ │ 1│ 1│ 1║ │
│ 47│ │Emalthion, _Hübn._ ║ │ │ │ ║ 1│
│ 48│ │Deiphontes, _Wall._║ │ │ │ ║ │
│ 49│ │Deiphobus, _L._ ║ │ │ │ ║ │
│ 50│ │Ascalaphus, _Bd._ ║ │ │ │ ║ │
│ 51│ │Ænomaus, _Godt._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │g.│_Helenus_-group. ║ │ │ │ ║ │
│ 52│ │Severus, _Cr._ ║ │ │ │ ║ │
│ 53│ │Pertinax, _Wall._ ║ │ │ │ ║ │
│ 54│ │Albinus, _Wall._ ║ │ │ │ ║ │
│ 55│ │Phæstus, _Bd._ ║ │ │ │ ║ │
│ 56│ │Helenus, _L._ ║ │ 1│ │ 1║ │
│ 57│ │Hecuba, _Wall._ ║ │ │ │ ║ │
│ 58│ │Iswara, _White_ ║ 1│ │ 1│ ║ │
│ 59│ │Hystaspes, _Feld._ ║ │ │ │ ║ 1│
│ 60│ │Araspes, _Feld._ ║ │ │ │ ║ 1│
│ 61│ │Nephelus, _Bd._ ║ 1│ 1│ 1│ ║ │
│ │ │ ║ │ │ │ ║ │
│ │h.│_Pammon_-group. ║ │ │ │ ║ │
│ 62│ │Pammon, _L._ ║ 1│ │ │ ║ │
│ 63│ │Theseus, _Cr._ ║ │ 1│ 1│ 1║ │
│ 64│ │Alphenor, _Cr._ ║ │ │ │ ║ 1│
│ 65│ │Nicanor, _Wall._ ║ │ │ │ ║ │
│ 66│ │Hipponous, _Feld._ ║ │ │ │ ║ 1│
│ 67│ │Ambrax, _Bd._ ║ │ │ │ ║ │
│ 68│ │Ambracia, _Wall._ ║ │ │ │ ║ │
│ 69│ │Epirus, _Wall._ ║ │ │ │ ║ │
│ 70│ │Dunali, _Montr._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │i.│_Erectheus_-group. ║ │ │ │ ║ │
│ 71│ │Ormenus, _Guér._ ║ │ │ │ ║ │
│ 72│ │Pandion, _Wall._ ║ │ │ │ ║ │
│ 73│ │Tydeus, _Feld._ ║ │ │ │ ║ │
│ 74│ │Adrastus, _Wall._ ║ │ │ │ ║ │
│ 75│ │Gambrisius, _Cr._ ║ │ │ │ ║ │
│ 76│ │Amphytrion, _Cr._ ║ │ │ │ ║ │
│ 77│ │Euchenor, _Guér._ ║ │ │ │ ║ │
│ 78│ │Godartii, _Montr._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │k.│_Demolion_-group. ║ │ │ │ ║ │
│ 79│ │Demolion, _Cr._ ║ 1│ 1│ 1│ 1║ │
│ 80│ │Gigon, _Wall._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │l.│_Erithonius_-group.║ │ │ │ ║ │
│ 81│ │Erithonius, _Cr._ ║ 1│ │ │ ║ 1│
│ │ │ ║ │ │ │ ║ │
│ │m.│_Paradoxa_-group. ║ │ │ │ ║ │
│ 82│ │Paradoxa, _Zink._ ║ │ 1│ 1│ 1║ │
│ 83│ │Ænigma, _Wall._ ║ 1│ 1│ 1│ ║ │
│ 84│ │Caunus, _Westw._ ║ │ 1│ 1│ ║ │
│ 85│ │Astina, _Westw._ ║ │ │ │ 1║ │
│ 86│ │Hewitsonii, ║ │ │ 1│ ║ │
│ │ │ _Westw._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │n.│_Dissimilis_-group.║ │ │ │ ║ │
│ 87│ │Echidna, _De Haan._║ │ │ │ ║ │
│ 88│ │Paëphates, _Westw._║ │ │ │ ║ 1│
│ │ │ ║ │ │ │ ║ │
│ │o.│_Macareus_-group. ║ │ │ │ ║ │
│ 89│ │Veiovis, _Hew._ ║ │ │ │ ║ │
│ 90│ │Encelades, _Bd._ ║ │ │ │ ║ │
│ 91│ │Deucalion, _Bd._ ║ │ │ │ ║ │
│ 92│ │Idæoides, _Hew._ ║ │ │ │ ║ 1│
│ 93│ │Delessertii, ║ 1│ │ │ ║ │
│ │ │ _Guér._ ║ │ │ │ ║ │
│ 94│ │Dehaanii, _Wall._ ║ 1│ │ 1│ 1║ │
│ 95│ │Leucothoë, _Westw._║ 1│ │ │ ║ │
│ 96│ │Macareus, _Godt._ ║ 1│ │ 1│ 1║ │
│ 97│ │Stratocles, _Feld._║ │ │ │ ║ 1│
│ 98│ │Thule, _Wall._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │p.│_Antiphates_-group.║ │ │ │ ║ │
│ 99│ │Antiphates, _Cr._ ║ 1│ 1│ 1│ 1║ │
│100│ │Euphrates, _Feld._ ║ │ │ │ ║ 1│
│101│ │Androcles, _Bd._ ║ │ │ │ ║ │
│102│ │Dorcus, _De Haan._ ║ │ │ │ ║ │
│103│ │Rhesus, _Bd._ ║ │ │ │ ║ │
│104│ │Aristæus, _Cr._ ║ │ │ │ ║ │
│105│ │Parmatus, _G. R. ║ │ │ │ ║ │
│ │ │ G._ ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │q.│_Eurypylus_-group. ║ │ │ │ ║ │
│106│ │Codrus, _Cr._ ║ │ │ │ ║ │
│107│ │Melanthus, _Feld._ ║ │ │ │ ║ 1│
│108│ │Empedocles, _Fab._ ║ │ │ 1│ 1║ │
│109│ │Payeni, _Bd._ ║ │ │ 1│ 1║ │
│110│ │Sarpedon, _L._ ║ 1│ 1│ 1│ 1║ │
│111│ │Miletus, _Wall._ ║ │ │ │ ║ │
│112│ │Wallacei, _Hew._ ║ │ │ │ ║ │
│113│ │Bathycles, _Zink._ ║ 1│ │ 1│ 1║ │
│114│ │Eurypylus, _L._ ║ │ │ │ ║ │
│115│ │Jason, _Esp._ ║ 1│ 1│ 1│ 1║ │
│116│ │Telephus, _Wall._ ║ │ │ │ ║ │
│117│ │Ægistus, _L._ ║ │ │ │ ║ │
│118│ │Agamemnon, _L._ ║ 1│ 1│ 1│ 1║ 1│
│119│ │Rama, _Feld._ ║ 1│ 1│ │ ║ │
│ │ │(? Arycles, _Rd._) ║ │ │ │ ║ │
│ │ │ ║ │ │ │ ║ │
│ │ │=Leptocircus.= ║ │ │ │ ║ │
│120│ │Meges, _Zink._ ║ 1│ │ │ 1║ │
│121│ │Curtius, _Wall._ ║ │ │ │ ║ │
│122│ │Decius, _Feld._ ║ │ │ │ ║ 1│
│123│ │Curius, _Fab._ ║ │ │ │ 1║ │
│ │ Totals:— ║ │ │ │ ║ │
│ │ │Ornithoptera ║ 2│ 2│ 3│ 2║ 2│
│ │ │Papilio ║ 22│ 19│ 26│ 23║ 17│
│ │ │Leptocircus ║ 1│ │ │ 2║ 1│
├───┼──┼───────────────────╫────────┼────────┼────────┼─────╫────────────┤
│ │ │Species in each ║ 25│ 21│ 29│ 27║ 20│
│ │ │ island ║ │ │ │ ║ │
├───┼──┼───────────────────╫────────┴────────┴────────┴─────╫────────────┤
│ │ │ Total ║ 45 ║ 20│
│ │ │ ║ Sixty-one, Indo-Malayan Region. │
└───┴──┴───────────────────╨─────────────────────────────────────────────┘
┌───┬──┬───────────────────╥─────────────────────────────────────────────────┐
│ │ │ ║ AUSTRO-MALAYAN REGION. │
├───┼──┼───────────────────╫────────╥────────┬──────╥───────┬─────────┬──────┤
│ │ │ =Ornithoptera.= ║Celebes.║Lombock.│Timor.║Gilolo.│Batchian.│Bouru.│
│ │ │ ║ ║ │ ║ │ │ │
├───┼──┼───────────────────╫────────╥────────┬──────╥───────┬─────────┬──────┤
│ │a.│_Priamus_-group. ║ ║ │ ║ │ │ │
│ 1│ │Priamus, _L._ ║ ║ │ ║ │ │ │
│ 2│ │Poseidon, _Db._ ║ ║ │ ║ │ │ │
│ 3│ │Crœsus, _Feld._ ║ ║ │ ║ 1│ 1│ │
│ 4│ │Tithonus, _De ║ ║ │ ║ │ │ │
│ │ │ Haan._ ║ ║ │ ║ │ │ │
│ 5│ │Urvilliana, _Guér._║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │b.│_Pompeus_-group. ║ ║ │ ║ │ │ │
│ 6│ │Remus, _Cr._ ║ 1║ │ ║ 1│ │ │
│ 7│ │Helena, _L._ ║ 1║ │ ║ │ │ 1│
│ 8│ │Leda, _Wall._ ║ 1║ │ ║ │ │ │
│ 9│ │Pompeus, _Cr._ ║ ║ 1│ ║ │ │ │
│ 10│ │Nephereus, _G. R. ║ ║ │ ║ │ │ │
│ │ │ G._ ║ ║ │ ║ │ │ │
│ 11│ │Magellanus, _Feld._║ ║ │ ║ │ │ │
│ 12│ │Criton, _Feld._ ║ ║ │ ║ 1│ 1│ │
│ 13│ │Plato, _Wall._ ║ ║ │ 1║ │ │ │
│ 14│ │Haliphron, _Bd._ ║ 1║ │ ║ │ │ │
│ 15│ │Amphrisius, _Cr._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │c.│_Brookeana_-group. ║ ║ │ ║ │ │ │
│ 16│ │Brookeana, _Wall._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │ │ =Papilio.= ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ A.│a.│_Nox_-group. ║ ║ │ ║ │ │ │
│ 17│ │Nox, _Sw._ ║ ║ │ ║ │ │ │
│ 18│ │Noctis, _Hew._ ║ ║ │ ║ │ │ │
│ 19│ │Erebus, _Wall._ ║ ║ │ ║ │ │ │
│ 20│ │Varuna, _White_ ║ ║ │ ║ │ │ │
│ 21│ │Semperi, _Feld._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │b.│_Coon_-group. ║ ║ │ ║ │ │ │
│ 22│ │Neptunus, _Guér._ ║ ║ │ ║ │ │ │
│ 23│ │Coon, _Fab._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │c.│_Polydorus_-group. ║ ║ │ ║ │ │ │
│ 24│ │Polydorus, _L._ ║ ║ │ ║ │ 1│ 1│
│ 25│ │Leodamas, _Wall._ ║ ║ │ ║ │ │ │
│ 26│ │Diphilus, _Esper_ ║ ║ │ ║ │ │ │
│ 27│ │Antiphus, _Fab._ ║ ║ 1│ ║ │ │ │
│ 28│ │Polyphontes, _Bd._ ║ 1║ │ ║ 1│ 1│ │
│ 29│ │Annæ, _Feld._ ║ ║ │ ║ │ │ │
│ 30│ │Liris, _Godt._ ║ ║ │ 1║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ B.│d.│_Ulysses_-group. ║ ║ │ ║ │ │ │
│ 31│ │Ulysses, _L._ ║ ║ │ ║ │ │ │
│ 32│ │Penelope, _Wall._ ║ ║ │ ║ │ │ │
│ 33│ │Telegonus, _Feld._ ║ ║ │ ║ 1│ 1│ │
│ 34│ │Telemachus, _Mont._║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │e.│_Peranthus_-group. ║ ║ │ ║ │ │ │
│ 35│ │Peranthus, _Fab._ ║ ║ 1│ ║ │ │ │
│ 36│ │Pericles, _Wall._ ║ ║ │ 1║ │ │ │
│ 37│ │Philippus, _Wall._ ║ ║ │ ║ │ 1│ │
│ 38│ │Macedon, _Wall._ ║ 1║ │ ║ │ │ │
│ 39│ │Brama, _Guér._ ║ ║ │ ║ │ │ │
│ 40│ │Dædalus, _Feld._ ║ ║ │ ║ │ │ │
│ 41│ │Blumei, _Bd._ ║ 1║ │ ║ │ │ │
│ 42│ │Arjuna, _Horsf._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │f.│_Memnon_-group. ║ ║ │ ║ │ │ │
│ 43│ │Memnon, _L._ ║ ║ 1│ ║ │ │ │
│ 44│ │Androgeus, _Cr._ ║ ║ │ ║ │ │ │
│ 45│ │Lampsacus, _Bd._ ║ ║ │ ║ │ │ │
│ 46│ │Priapus, _Bd._ ║ ║ │ ║ │ │ │
│ 47│ │Emalthion, _Hübn._ ║ ║ │ ║ │ │ │
│ 48│ │Deiphontes, _Wall._║ ║ │ ║ 1│ 1│ │
│ 49│ │Deiphobus, _L._ ║ ║ │ ║ │ │ 1│
│ 50│ │Ascalaphus, _Bd._ ║ 1║ │ ║ │ │ │
│ 51│ │Ænomaus, _Godt._ ║ ║ │ 1║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │g.│_Helenus_-group. ║ ║ │ ║ │ │ │
│ 52│ │Severus, _Cr._ ║ ║ │ ║ 1│ 1│ 1│
│ 53│ │Pertinax, _Wall._ ║ 1║ │ ║ │ │ │
│ 54│ │Albinus, _Wall._ ║ ║ │ ║ │ │ │
│ 55│ │Phæstus, _Bd._ ║ ║ │ ║ │ │ │
│ 56│ │Helenus, _L._ ║ ║ │ ║ │ │ │
│ 57│ │Hecuba, _Wall._ ║ 1║ │ ║ │ │ │
│ 58│ │Iswara, _White_ ║ ║ │ ║ │ │ │
│ 59│ │Hystaspes, _Feld._ ║ ║ │ ║ │ │ │
│ 60│ │Araspes, _Feld._ ║ ║ │ ║ │ │ │
│ 61│ │Nephelus, _Bd._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │h.│_Pammon_-group. ║ ║ │ ║ │ │ │
│ 62│ │Pammon, _L._ ║ ║ │ ║ │ │ │
│ 63│ │Theseus, _Cr._ ║ ║ 1│ 1║ │ │ │
│ 64│ │Alphenor, _Cr._ ║ 1║ │ ║ │ │ 1│
│ 65│ │Nicanor, _Wall._ ║ ║ │ ║ 1│ 1│ │
│ 66│ │Hipponous, _Feld._ ║ ║ │ ║ │ │ │
│ 67│ │Ambrax, _Bd._ ║ ║ │ ║ │ │ │
│ 68│ │Ambracia, _Wall._ ║ ║ │ ║ │ │ │
│ 69│ │Epirus, _Wall._ ║ ║ │ ║ │ │ │
│ 70│ │Dunali, _Montr._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │i.│_Erectheus_-group. ║ ║ │ ║ │ │ │
│ 71│ │Ormenus, _Guér._ ║ ║ │ ║ │ │ │
│ 72│ │Pandion, _Wall._ ║ ║ │ ║ │ │ │
│ 73│ │Tydeus, _Feld._ ║ ║ │ ║ 1│ 1│ │
│ 74│ │Adrastus, _Wall._ ║ ║ │ ║ │ │ │
│ 75│ │Gambrisius, _Cr._ ║ ║ │ ║ │ │ 1│
│ 76│ │Amphytrion, _Cr._ ║ 1║ │ ║ │ │ │
│ 77│ │Euchenor, _Guér._ ║ ║ │ ║ │ │ │
│ 78│ │Godartii, _Montr._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │k.│_Demolion_-group. ║ ║ │ ║ │ │ │
│ 79│ │Demolion, _Cr._ ║ ║ │ ║ │ │ │
│ 80│ │Gigon, _Wall._ ║ 1║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │l.│_Erithonius_-group.║ ║ │ ║ │ │ │
│ 81│ │Erithonius, _Cr._ ║ ║ │ 1║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │m.│_Paradoxa_-group. ║ ║ │ ║ │ │ │
│ 82│ │Paradoxa, _Zink._ ║ ║ │ ║ │ │ │
│ 83│ │Ænigma, _Wall._ ║ ║ │ ║ │ │ │
│ 84│ │Caunus, _Westw._ ║ ║ │ ║ │ │ │
│ 85│ │Astina, _Westw._ ║ ║ │ ║ │ │ │
│ 86│ │Hewitsonii, ║ ║ │ ║ │ │ │
│ │ │ _Westw._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │n.│_Dissimilis_-group.║ ║ │ ║ │ │ │
│ 87│ │Echidna, _De Haan._║ ║ │ 1║ │ │ │
│ 88│ │Paëphates, _Westw._║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │o.│_Macareus_-group. ║ ║ │ ║ │ │ │
│ 89│ │Veiovis, _Hew._ ║ 1║ │ ║ │ │ │
│ 90│ │Encelades, _Bd._ ║ 1║ │ ║ │ │ │
│ 91│ │Deucalion, _Bd._ ║ 1║ │ ║ │ │ │
│ 92│ │Idæoides, _Hew._ ║ ║ │ ║ │ │ │
│ 93│ │Delessertii, ║ ║ │ ║ │ │ │
│ │ │ _Guér._ ║ ║ │ ║ │ │ │
│ 94│ │Dehaanii, _Wall._ ║ ║ │ ║ │ │ │
│ 95│ │Leucothoë, _Westw._║ ║ │ ║ │ │ │
│ 96│ │Macareus, _Godt._ ║ ║ │ ║ │ │ │
│ 97│ │Stratocles, _Feld._║ ║ │ ║ │ │ │
│ 98│ │Thule, _Wall._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │p.│_Antiphates_-group.║ ║ │ ║ │ │ │
│ 99│ │Antiphates, _Cr._ ║ ║ │ ║ │ │ │
│100│ │Euphrates, _Feld._ ║ ║ │ ║ │ │ │
│101│ │Androcles, _Bd._ ║ 1║ │ ║ │ │ │
│102│ │Dorcus, _De Haan._ ║ 1║ │ ║ │ │ │
│103│ │Rhesus, _Bd._ ║ 1║ │ ║ │ │ │
│104│ │Aristæus, _Cr._ ║ ║ │ ║ │ 1│ │
│105│ │Parmatus, _G. R. ║ ║ │ ║ │ │ │
│ │ │ G._ ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │q.│_Eurypylus_-group. ║ ║ │ ║ │ │ │
│106│ │Codrus, _Cr._ ║ 1║ │ ║ 1│ 1│ 1│
│107│ │Melanthus, _Feld._ ║ ║ │ ║ │ │ │
│108│ │Empedocles, _Fab._ ║ ║ │ ║ │ │ │
│109│ │Payeni, _Bd._ ║ ║ │ ║ │ │ │
│110│ │Sarpedon, _L._ ║ ║ │ ║ 1│ 1│ 1│
│111│ │Miletus, _Wall._ ║ 1║ │ ║ │ │ │
│112│ │Wallacei, _Hew._ ║ ║ │ ║ │ 1│ │
│113│ │Bathycles, _Zink._ ║ ║ │ ║ │ │ │
│114│ │Eurypylus, _L._ ║ ║ │ ║ 1│ 1│ 1│
│115│ │Jason, _Esp._ ║ ║ │ ║ │ │ │
│116│ │Telephus, _Wall._ ║ 1║ │ ║ │ │ │
│117│ │Ægistus, _L._ ║ ║ │ ║ 1│ 1│ │
│118│ │Agamemnon, _L._ ║ 1║ 1│ 1║ 1│ 1│ 1│
│119│ │Rama, _Feld._ ║ ║ │ ║ │ │ │
│ │ │(? Arycles, _Rd._) ║ ║ │ ║ │ │ │
│ │ │ ║ ║ │ ║ │ │ │
│ │ │=Leptocircus.= ║ ║ │ ║ │ │ │
│120│ │Meges, _Zink._ ║ ║ │ ║ │ │ │
│121│ │Curtius, _Wall._ ║ 1║ │ ║ │ │ │
│122│ │Decius, _Feld._ ║ ║ │ ║ │ │ │
│123│ │Curius, _Fab._ ║ ║ │ ║ │ │ │
│ │ Totals:— ║ ║ │ ║ │ │ │
│ │ │Ornithoptera ║ 4║ 1│ 1║ 3│ 2│ 1│
│ │ │Papilio ║ 19║ 5│ 7║ 11│ 15│ 9│
│ │ │Leptocircus ║ 1║ │ ║ │ │ │
├───┼──┼───────────────────╫────────╫────────┼──────╫───────┼─────────┼──────┤
│ │ │Species in each ║ 24║ 6│ 8║ 14│ 17│ 10│
│ │ │ island ║ ║ │ ║ │ │ │
├───┼──┼───────────────────╫────────╫────────┴──────╫───────┴─────────┴──────┤
│ │ │ Total ║ 24║ 12 ║ │
│ │ │ ║ ║ Seventy-two, Austro-Malayan Region. │
└───┴──┴───────────────────╨────────╨────────────────────────────────────────┘
┌───┬──┬───────────────────╥────────────────────────────┐
│ │ │ ║ AUSTRO-MALAYAN REGION. │
├───┼──┼───────────────────╫──────┬──────┬──────┬───────┤
│ │ │ =Ornithoptera.= ║Ceram.│Banda.│Goram.│ Ké │
│ │ │ ║ │ │ │Island.│
├───┼──┼───────────────────╫──────┬──────┬──────┬───────┤
│ │a.│_Priamus_-group. ║ 1│ │ │ │
│ 1│ │Priamus, _L._ ║ 1│ │ │ │
│ 2│ │Poseidon, _Db._ ║ │ │ │ │
│ 3│ │Crœsus, _Feld._ ║ │ │ │ │
│ 4│ │Tithonus, _De ║ │ │ │ │
│ │ │ Haan._ ║ │ │ │ │
│ 5│ │Urvilliana, _Guér._║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │b.│_Pompeus_-group. ║ │ │ │ │
│ 6│ │Remus, _Cr._ ║ 1│ │ │ │
│ 7│ │Helena, _L._ ║ 1│ │ │ │
│ 8│ │Leda, _Wall._ ║ │ │ │ │
│ 9│ │Pompeus, _Cr._ ║ │ │ │ │
│ 10│ │Nephereus, _G. R. ║ │ │ │ │
│ │ │ G._ ║ │ │ │ │
│ 11│ │Magellanus, _Feld._║ │ │ │ │
│ 12│ │Criton, _Feld._ ║ │ │ │ │
│ 13│ │Plato, _Wall._ ║ │ │ │ │
│ 14│ │Haliphron, _Bd._ ║ │ │ │ │
│ 15│ │Amphrisius, _Cr._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │c.│_Brookeana_-group. ║ │ │ │ │
│ 16│ │Brookeana, _Wall._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │ │ =Papilio.= ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ A.│a.│_Nox_-group. ║ │ │ │ │
│ 17│ │Nox, _Sw._ ║ │ │ │ │
│ 18│ │Noctis, _Hew._ ║ │ │ │ │
│ 19│ │Erebus, _Wall._ ║ │ │ │ │
│ 20│ │Varuna, _White_ ║ │ │ │ │
│ 21│ │Semperi, _Feld._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │b.│_Coon_-group. ║ │ │ │ │
│ 22│ │Neptunus, _Guér._ ║ │ │ │ │
│ 23│ │Coon, _Fab._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │c.│_Polydorus_-group. ║ │ │ │ │
│ 24│ │Polydorus, _L._ ║ 1│ │ │ 1│
│ 25│ │Leodamas, _Wall._ ║ │ │ │ │
│ 26│ │Diphilus, _Esper_ ║ │ │ │ │
│ 27│ │Antiphus, _Fab._ ║ │ │ │ │
│ 28│ │Polyphontes, _Bd._ ║ │ │ │ │
│ 29│ │Annæ, _Feld._ ║ │ │ │ │
│ 30│ │Liris, _Godt._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ B.│d.│_Ulysses_-group. ║ │ │ │ │
│ 31│ │Ulysses, _L._ ║ 1│ │ │ │
│ 32│ │Penelope, _Wall._ ║ │ │ │ │
│ 33│ │Telegonus, _Feld._ ║ │ │ │ │
│ 34│ │Telemachus, _Mont._║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │e.│_Peranthus_-group. ║ │ │ │ │
│ 35│ │Peranthus, _Fab._ ║ │ │ │ │
│ 36│ │Pericles, _Wall._ ║ │ │ │ │
│ 37│ │Philippus, _Wall._ ║ 1│ │ │ │
│ 38│ │Macedon, _Wall._ ║ │ │ │ │
│ 39│ │Brama, _Guér._ ║ │ │ │ │
│ 40│ │Dædalus, _Feld._ ║ │ │ │ │
│ 41│ │Blumei, _Bd._ ║ │ │ │ │
│ 42│ │Arjuna, _Horsf._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │f.│_Memnon_-group. ║ │ │ │ │
│ 43│ │Memnon, _L._ ║ │ │ │ │
│ 44│ │Androgeus, _Cr._ ║ │ │ │ │
│ 45│ │Lampsacus, _Bd._ ║ │ │ │ │
│ 46│ │Priapus, _Bd._ ║ │ │ │ │
│ 47│ │Emalthion, _Hübn._ ║ │ │ │ │
│ 48│ │Deiphontes, _Wall._║ │ │ │ │
│ 49│ │Deiphobus, _L._ ║ 1│ │ │ │
│ 50│ │Ascalaphus, _Bd._ ║ │ │ │ │
│ 51│ │Ænomaus, _Godt._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │g.│_Helenus_-group. ║ │ │ │ │
│ 52│ │Severus, _Cr._ ║ 1│ │ │ │
│ 53│ │Pertinax, _Wall._ ║ │ │ │ │
│ 54│ │Albinus, _Wall._ ║ │ │ │ │
│ 55│ │Phæstus, _Bd._ ║ │ │ │ │
│ 56│ │Helenus, _L._ ║ │ │ │ │
│ 57│ │Hecuba, _Wall._ ║ │ │ │ │
│ 58│ │Iswara, _White_ ║ │ │ │ │
│ 59│ │Hystaspes, _Feld._ ║ │ │ │ │
│ 60│ │Araspes, _Feld._ ║ │ │ │ │
│ 61│ │Nephelus, _Bd._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │h.│_Pammon_-group. ║ │ │ │ │
│ 62│ │Pammon, _L._ ║ │ │ │ │
│ 63│ │Theseus, _Cr._ ║ │ │ │ │
│ 64│ │Alphenor, _Cr._ ║ 1│ │ │ │
│ 65│ │Nicanor, _Wall._ ║ │ │ │ │
│ 66│ │Hipponous, _Feld._ ║ │ │ │ │
│ 67│ │Ambrax, _Bd._ ║ │ │ │ │
│ 68│ │Ambracia, _Wall._ ║ │ │ │ │
│ 69│ │Epirus, _Wall._ ║ │ │ │ │
│ 70│ │Dunali, _Montr._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │i.│_Erectheus_-group. ║ │ │ │ │
│ 71│ │Ormenus, _Guér._ ║ │ │ 1│ 1│
│ 72│ │Pandion, _Wall._ ║ │ │ │ │
│ 73│ │Tydeus, _Feld._ ║ │ │ │ │
│ 74│ │Adrastus, _Wall._ ║ │ 1│ │ │
│ 75│ │Gambrisius, _Cr._ ║ 1│ │ │ │
│ 76│ │Amphytrion, _Cr._ ║ │ │ │ │
│ 77│ │Euchenor, _Guér._ ║ │ │ │ 1│
│ 78│ │Godartii, _Montr._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │k.│_Demolion_-group. ║ │ │ │ │
│ 79│ │Demolion, _Cr._ ║ │ │ │ │
│ 80│ │Gigon, _Wall._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │l.│_Erithonius_-group.║ │ │ │ │
│ 81│ │Erithonius, _Cr._ ║ │ │ │ 1│
│ │ │ ║ │ │ │ │
│ │m.│_Paradoxa_-group. ║ │ │ │ │
│ 82│ │Paradoxa, _Zink._ ║ │ │ │ │
│ 83│ │Ænigma, _Wall._ ║ │ │ │ │
│ 84│ │Caunus, _Westw._ ║ │ │ │ │
│ 85│ │Astina, _Westw._ ║ │ │ │ │
│ 86│ │Hewitsonii, ║ │ │ │ │
│ │ │ _Westw._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │n.│_Dissimilis_-group.║ │ │ │ │
│ 87│ │Echidna, _De Haan._║ │ │ │ │
│ 88│ │Paëphates, _Westw._║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │o.│_Macareus_-group. ║ │ │ │ │
│ 89│ │Veiovis, _Hew._ ║ │ │ │ │
│ 90│ │Encelades, _Bd._ ║ │ │ │ │
│ 91│ │Deucalion, _Bd._ ║ │ │ │ │
│ 92│ │Idæoides, _Hew._ ║ │ │ │ │
│ 93│ │Delessertii, ║ │ │ │ │
│ │ │ _Guér._ ║ │ │ │ │
│ 94│ │Dehaanii, _Wall._ ║ │ │ │ │
│ 95│ │Leucothoë, _Westw._║ │ │ │ │
│ 96│ │Macareus, _Godt._ ║ │ │ │ │
│ 97│ │Stratocles, _Feld._║ │ │ │ │
│ 98│ │Thule, _Wall._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │p.│_Antiphates_-group.║ │ │ │ │
│ 99│ │Antiphates, _Cr._ ║ │ │ │ │
│100│ │Euphrates, _Feld._ ║ │ │ │ │
│101│ │Androcles, _Bd._ ║ │ │ │ │
│102│ │Dorcus, _De Haan._ ║ │ │ │ │
│103│ │Rhesus, _Bd._ ║ │ │ │ │
│104│ │Aristæus, _Cr._ ║ 1│ │ │ │
│105│ │Parmatus, _G. R. ║ │ │ │ │
│ │ │ G._ ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │q.│_Eurypylus_-group. ║ │ │ │ │
│106│ │Codrus, _Cr._ ║ 1│ │ │ │
│107│ │Melanthus, _Feld._ ║ │ │ │ │
│108│ │Empedocles, _Fab._ ║ │ │ │ │
│109│ │Payeni, _Bd._ ║ │ │ │ │
│110│ │Sarpedon, _L._ ║ 1│ │ │ │
│111│ │Miletus, _Wall._ ║ │ │ │ │
│112│ │Wallacei, _Hew._ ║ │ │ │ │
│113│ │Bathycles, _Zink._ ║ │ │ │ │
│114│ │Eurypylus, _L._ ║ 1│ │ │ │
│115│ │Jason, _Esp._ ║ │ │ │ │
│116│ │Telephus, _Wall._ ║ │ │ │ │
│117│ │Ægistus, _L._ ║ 1│ │ │ 1│
│118│ │Agamemnon, _L._ ║ 1│ │ │ 1│
│119│ │Rama, _Feld._ ║ │ │ │ │
│ │ │(? Arycles, _Rd._) ║ │ │ │ │
│ │ │ ║ │ │ │ │
│ │ │=Leptocircus.= ║ │ │ │ │
│120│ │Meges, _Zink._ ║ │ │ │ │
│121│ │Curtius, _Wall._ ║ │ │ │ │
│122│ │Decius, _Feld._ ║ │ │ │ │
│123│ │Curius, _Fab._ ║ │ │ │ │
│ │ Totals:— ║ │ │ │ │
│ │ │Ornithoptera ║ 3│ │ 1│ 1│
│ │ │Papilio ║ 13│ 1│ 2│ 4│
│ │ │Leptocircus ║ │ │ │ │
├───┼──┼───────────────────╫──────┼──────┼──────┼───────┤
│ │ │Species in each ║ 16│ 1│ 3│ 5│
│ │ │ island ║ │ │ │ │
├───┼──┼───────────────────╫──────┴──────┴──────┴───────┤
│ │ │ Total ║ 27 │
│ │ │ ║ Seventy-two, │
│ │ │ ║ Austro-Malayan Region. │
└───┴──┴───────────────────╨────────────────────────────┘
┌───┬──┬───────────────────╥──────────────────────────────────────────────────┐
│ │ │ ║ AUSTRO-MALAYAN REGION. │
├───┼──┼───────────────────╫────────┬──────┬────────┬───────┬────────┬────────┤
│ │ │ =Ornithoptera.= ║ Aru │Mysol.│Waigiou.│ New │ New │Woodlark│
│ │ │ ║Islands.│ │ │Guinea.│Ireland.│ Isl. │
├───┼──┼───────────────────╫────────┬──────┬────────┬───────┬────────┬────────┤
│ │a.│_Priamus_-group. ║ │ │ │ │ │ │
│ 1│ │Priamus, _L._ ║ │ │ │ │ │ │
│ 2│ │Poseidon, _Db._ ║ 1│ 1│ 1│ 1│ │ 1│
│ 3│ │Crœsus, _Feld._ ║ │ │ │ │ │ │
│ 4│ │Tithonus, _De ║ │ │ │ 1│ │ │
│ │ │ Haan._ ║ │ │ │ │ │ │
│ 5│ │Urvilliana, _Guér._║ │ │ │ │ 1│ │
│ │ │ ║ │ │ │ │ │ │
│ │b.│_Pompeus_-group. ║ │ │ │ │ │ │
│ 6│ │Remus, _Cr._ ║ │ │ │ │ │ │
│ 7│ │Helena, _L._ ║ │ │ │ 1_v_│ │ │
│ 8│ │Leda, _Wall._ ║ │ │ │ │ │ │
│ 9│ │Pompeus, _Cr._ ║ │ │ │ │ │ │
│ 10│ │Nephereus, _G. R. ║ │ │ │ │ │ │
│ │ │ G._ ║ │ │ │ │ │ │
│ 11│ │Magellanus, _Feld._║ │ │ │ │ │ │
│ 12│ │Criton, _Feld._ ║ │ │ │ │ │ │
│ 13│ │Plato, _Wall._ ║ │ │ │ │ │ │
│ 14│ │Haliphron, _Bd._ ║ │ │ │ │ │ │
│ 15│ │Amphrisius, _Cr._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │c.│_Brookeana_-group. ║ │ │ │ │ │ │
│ 16│ │Brookeana, _Wall._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │ │ =Papilio.= ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ A.│a.│_Nox_-group. ║ │ │ │ │ │ │
│ 17│ │Nox, _Sw._ ║ │ │ │ │ │ │
│ 18│ │Noctis, _Hew._ ║ │ │ │ │ │ │
│ 19│ │Erebus, _Wall._ ║ │ │ │ │ │ │
│ 20│ │Varuna, _White_ ║ │ │ │ │ │ │
│ 21│ │Semperi, _Feld._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │b.│_Coon_-group. ║ │ │ │ │ │ │
│ 22│ │Neptunus, _Guér._ ║ │ │ │ │ │ │
│ 23│ │Coon, _Fab._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │c.│_Polydorus_-group. ║ │ │ │ │ │ │
│ 24│ │Polydorus, _L._ ║ 1│ │ │ │ │ │
│ 25│ │Leodamas, _Wall._ ║ │ 1│ │ 1│ │ │
│ 26│ │Diphilus, _Esper_ ║ │ │ │ │ │ │
│ 27│ │Antiphus, _Fab._ ║ │ │ │ │ │ │
│ 28│ │Polyphontes, _Bd._ ║ │ │ │ │ │ │
│ 29│ │Annæ, _Feld._ ║ │ │ │ │ │ │
│ 30│ │Liris, _Godt._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ B.│d.│_Ulysses_-group. ║ │ │ │ │ │ │
│ 31│ │Ulysses, _L._ ║ │ │ │ │ │ │
│ 32│ │Penelope, _Wall._ ║ 1│ │ 1│ 1│ │ │
│ 33│ │Telegonus, _Feld._ ║ │ │ │ │ │ │
│ 34│ │Telemachus, _Mont._║ │ │ │ │ │ 1│
│ │ │ ║ │ │ │ │ │ │
│ │e.│_Peranthus_-group. ║ │ │ │ │ │ │
│ 35│ │Peranthus, _Fab._ ║ │ │ │ │ │ │
│ 36│ │Pericles, _Wall._ ║ │ │ │ │ │ │
│ 37│ │Philippus, _Wall._ ║ │ │ │ │ │ │
│ 38│ │Macedon, _Wall._ ║ │ │ │ │ │ │
│ 39│ │Brama, _Guér._ ║ │ │ │ │ │ │
│ 40│ │Dædalus, _Feld._ ║ │ │ │ │ │ │
│ 41│ │Blumei, _Bd._ ║ │ │ │ │ │ │
│ 42│ │Arjuna, _Horsf._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │f.│_Memnon_-group. ║ │ │ │ │ │ │
│ 43│ │Memnon, _L._ ║ │ │ │ │ │ │
│ 44│ │Androgeus, _Cr._ ║ │ │ │ │ │ │
│ 45│ │Lampsacus, _Bd._ ║ │ │ │ │ │ │
│ 46│ │Priapus, _Bd._ ║ │ │ │ │ │ │
│ 47│ │Emalthion, _Hübn._ ║ │ │ │ │ │ │
│ 48│ │Deiphontes, _Wall._║ │ │ │ │ │ │
│ 49│ │Deiphobus, _L._ ║ │ │ │ │ │ │
│ 50│ │Ascalaphus, _Bd._ ║ │ │ │ │ │ │
│ 51│ │Ænomaus, _Godt._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │g.│_Helenus_-group. ║ │ │ │ │ │ │
│ 52│ │Severus, _Cr._ ║ 1│ │ │ │ │ │
│ 53│ │Pertinax, _Wall._ ║ │ │ │ │ │ │
│ 54│ │Albinus, _Wall._ ║ │ │ │ 1│ │ │
│ 55│ │Phæstus, _Bd._ ║ │ │ │ 1│ │ │
│ 56│ │Helenus, _L._ ║ │ │ │ │ │ │
│ 57│ │Hecuba, _Wall._ ║ │ │ │ │ │ │
│ 58│ │Iswara, _White_ ║ │ │ │ │ │ │
│ 59│ │Hystaspes, _Feld._ ║ │ │ │ │ │ │
│ 60│ │Araspes, _Feld._ ║ │ │ │ │ │ │
│ 61│ │Nephelus, _Bd._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │h.│_Pammon_-group. ║ │ │ │ │ │ │
│ 62│ │Pammon, _L._ ║ │ │ │ │ │ │
│ 63│ │Theseus, _Cr._ ║ │ │ │ │ │ │
│ 64│ │Alphenor, _Cr._ ║ │ │ │ │ │ │
│ 65│ │Nicanor, _Wall._ ║ │ │ │ │ │ │
│ 66│ │Hipponous, _Feld._ ║ │ │ │ │ │ │
│ 67│ │Ambrax, _Bd._ ║ │ 1│ │ 1│ │ │
│ 68│ │Ambracia, _Wall._ ║ │ │ 1│ │ │ │
│ 69│ │Epirus, _Wall._ ║ 1│ │ │ │ │ │
│ 70│ │Dunali, _Montr._ ║ │ │ │ │ │ 1│
│ │ │ ║ │ │ │ │ │ │
│ │i.│_Erectheus_-group. ║ │ │ │ │ │ │
│ 71│ │Ormenus, _Guér._ ║ 1│ │ 1│ │ │ │
│ 72│ │Pandion, _Wall._ ║ │ 1│ │ 1│ │ │
│ 73│ │Tydeus, _Feld._ ║ │ │ │ │ │ │
│ 74│ │Adrastus, _Wall._ ║ │ │ │ │ │ │
│ 75│ │Gambrisius, _Cr._ ║ │ │ │ │ │ │
│ 76│ │Amphytrion, _Cr._ ║ │ │ │ │ │ │
│ 77│ │Euchenor, _Guér._ ║ 1│ │ 1│ 1│ │ │
│ 78│ │Godartii, _Montr._ ║ │ │ │ │ │ 1│
│ │ │ ║ │ │ │ │ │ │
│ │k.│_Demolion_-group. ║ │ │ │ │ │ │
│ 79│ │Demolion, _Cr._ ║ │ │ │ │ │ │
│ 80│ │Gigon, _Wall._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │l.│_Erithonius_-group.║ │ │ │ │ │ │
│ 81│ │Erithonius, _Cr._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │m.│_Paradoxa_-group. ║ │ │ │ │ │ │
│ 82│ │Paradoxa, _Zink._ ║ │ │ │ │ │ │
│ 83│ │Ænigma, _Wall._ ║ │ │ │ │ │ │
│ 84│ │Caunus, _Westw._ ║ │ │ │ │ │ │
│ 85│ │Astina, _Westw._ ║ │ │ │ │ │ │
│ 86│ │Hewitsonii, ║ │ │ │ │ │ │
│ │ │ _Westw._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │n.│_Dissimilis_-group.║ │ │ │ │ │ │
│ 87│ │Echidna, _De Haan._║ │ │ │ │ │ │
│ 88│ │Paëphates, _Westw._║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │o.│_Macareus_-group. ║ │ │ │ │ │ │
│ 89│ │Veiovis, _Hew._ ║ │ │ │ │ │ │
│ 90│ │Encelades, _Bd._ ║ │ │ │ │ │ │
│ 91│ │Deucalion, _Bd._ ║ │ │ │ │ │ │
│ 92│ │Idæoides, _Hew._ ║ │ │ │ │ │ │
│ 93│ │Delessertii, ║ │ │ │ │ │ │
│ │ │ _Guér._ ║ │ │ │ │ │ │
│ 94│ │Dehaanii, _Wall._ ║ │ │ │ │ │ │
│ 95│ │Leucothoë, _Westw._║ │ │ │ │ │ │
│ 96│ │Macareus, _Godt._ ║ │ │ │ │ │ │
│ 97│ │Stratocles, _Feld._║ │ │ │ │ │ │
│ 98│ │Thule, _Wall._ ║ │ │ 1│ 1│ │ │
│ │ │ ║ │ │ │ │ │ │
│ │p.│_Antiphates_-group.║ │ │ │ │ │ │
│ 99│ │Antiphates, _Cr._ ║ │ │ │ │ │ │
│100│ │Euphrates, _Feld._ ║ │ │ │ │ │ │
│101│ │Androcles, _Bd._ ║ │ │ │ │ │ │
│102│ │Dorcus, _De Haan._ ║ │ │ │ │ │ │
│103│ │Rhesus, _Bd._ ║ │ │ │ │ │ │
│104│ │Aristæus, _Cr._ ║ │ │ │ │ │ │
│105│ │Parmatus, _G. R. ║ 1│ │ 1│ │ │ │
│ │ │ G._ ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │q.│_Eurypylus_-group. ║ │ │ │ │ │ │
│106│ │Codrus, _Cr._ ║ 1│ │ 1│ │ │ │
│107│ │Melanthus, _Feld._ ║ │ │ │ │ │ │
│108│ │Empedocles, _Fab._ ║ │ │ │ │ │ │
│109│ │Payeni, _Bd._ ║ │ │ │ │ │ │
│110│ │Sarpedon, _L._ ║ 1│ │ │ 1│ │ │
│111│ │Miletus, _Wall._ ║ │ │ │ │ │ │
│112│ │Wallacei, _Hew._ ║ 1│ │ │ │ │ │
│113│ │Bathycles, _Zink._ ║ │ │ │ │ │ │
│114│ │Eurypylus, _L._ ║ │ │ │ 1│ │ │
│115│ │Jason, _Esp._ ║ │ │ │ │ │ │
│116│ │Telephus, _Wall._ ║ │ │ │ │ │ │
│117│ │Ægistus, _L._ ║ 1│ │ │ │ │ │
│118│ │Agamemnon, _L._ ║ 1│ 1│ 1│ 1│ │ │
│119│ │Rama, _Feld._ ║ │ │ │ │ │ │
│ │ │(? Arycles, _Rd._) ║ │ │ │ │ │ │
│ │ │ ║ │ │ │ │ │ │
│ │ │=Leptocircus.= ║ │ │ │ │ │ │
│120│ │Meges, _Zink._ ║ │ │ │ │ │ │
│121│ │Curtius, _Wall._ ║ │ │ │ │ │ │
│122│ │Decius, _Feld._ ║ │ │ │ │ │ │
│123│ │Curius, _Fab._ ║ │ │ │ │ │ │
│ │ Totals:— ║ │ │ │ │ │ │
│ │ │Ornithoptera ║ 1│ 1│ 1│ 3│ 1│ 1│
│ │ │Papilio ║ 12│ 4│ 8│ 11│ │ 4│
│ │ │Leptocircus ║ │ │ │ │ │ │
├───┼──┼───────────────────╫────────┼──────┼────────┼───────┼────────┼────────┤
│ │ │Species in each ║ 13│ 5│ 9│ 14│ 1│ 5│
│ │ │ island ║ │ │ │ │ │ │
├───┼──┼───────────────────╫────────┴──────┴────────┴───────┴────────┴────────┤
│ │ │ Total ║ 27 │
│ │ │ ║ Seventy-two, Austro-Malayan Region. │
└───┴──┴───────────────────╨──────────────────────────────────────────────────┘
The exceeding richness of the Malayan region in these fine insects is
seen by comparing the number of species found in the different tropical
regions of the earth. From all Africa only 33 species of _Papilio_ are
known; but as several are still undescribed in collections, we may raise
their number to about 40. In all tropical Asia there are at present
described only 65 species, and I have seen in collections but two or
three which have not yet been named. In South America, south of Panama,
there are 120 species, or about the same number as I make in the Malayan
region; but the area of the two countries is very different; for while
South America (even excluding Patagonia) contains 5,000,000 square
miles, a line encircling the whole of the Malayan islands would only
include an area of 2,700,000 square miles, of which the land-area would
be about 1,000,000 square miles. This superior richness is partly real
and partly apparent. The breaking up of a district into small isolated
portions, as in an archipelago, seems highly favourable to the
segregation and perpetuation of local peculiarities in certain groups;
so that a species which on a continent might have a wide range, and
whose local forms, if any, would be so connected together that it would
be impossible to separate them, may become by isolation reduced to a
number of such clearly defined and constant forms that we are obliged to
count them as species. From this point of view, therefore, the superior
number of Malayan species may be considered as apparent only. Its true
superiority is shown, on the other hand, by the possession of three
genera and twenty groups of Papilionidæ against a single genus and eight
groups in South America, and also by the much greater average size of
the Malayan species. In most other families, however, the reverse is the
case, the South American _Nymphalidæ_, _Satyridæ_, and _Erycinidæ_ far
surpassing those of the East in number, variety, and beauty.
The following list, exhibiting the range and distribution of each group,
will enable us to study more easily their internal and external
relations.
_Range of the Groups of Malayan_ Papilionidæ.
Ornithoptera.
1. _Priamus_-group. Moluccas to Woodlark Island.
2. _Pompeus_-group. Himalayas to New Guinea (Celebes, maximum).
3. _Brookeana_-group. Sumatra and Borneo.
Papilio.
4. _Nox_-group. North India, Java, and Philippines.
5. _Coon_-group. North India to Java.
6. _Polydorus_-group. India to New Guinea and Pacific.
7. _Ulysses_-group. Celebes to New Caledonia.
8. _Peranthus_-group. India to Timor and Moluccas (India, max.).
9. _Memnon_-group. India to Timor and Moluccas (Java, max.).
10. _Helenus_-group. Africa and India to New Guinea.
11. _Pammon_-group. India to Pacific and Australia.
12. _Erechtheus_-group. Celebes to Australia.
13. _Demolion_-group. India to Celebes.
14. _Erithonius_-group. Africa, India, Australia.
15. _Paradoxa_-group. India to Java (Borneo, max.).
16. _Dissimilis_-group. India to Timor (India, max.).
17. _Macareus_-group. India to New Guinea.
18. _Antiphates_-group. Widely distributed.
19. _Eurypylus_-group. India to Australia.
Leptocircus.
20. _Leptocircus_-group. India to Celebes.
This Table shows the great affinity of the Malayan with the Indian
Papilionidæ, only three out of the nineteen groups ranging beyond, into
Africa, Europe, or America. The limitation of groups to the Indo-Malayan
or Austro-Malayan divisions of the archipelago, which is so well marked
in the higher animals (see ‘Journal of Linnean Society,’ vol. iv. 172,
and ‘Journal of the Royal Geographical Society,’ 1863, p. 230), is much
less conspicuous in insects, but is shown in some degree by the
Papilionidæ. The following groups are either almost or entirely
restricted to one portion of the Archipelago:—
_Indo-Malayan Region._
_Nox_-group.
_Coon_-group.
_Macareus_-group (nearly).
_Paradoxa_-group.
_Dissimilis_-group (nearly).
_Brookeanus_-group.
LEPTOCIRCUS (genus).
_Austro-Malayan Region._
_Priamus_-group.
_Ulysses_-group.
_Erechtheus_-group.
The remaining groups, which range over the whole archipelago, are, in
many cases, insects of very powerful flight, or they frequent open
places and the sea-beach, and are thus more likely to get blown from
island to island. The fact that three such characteristic groups as
those of _Priamus_, _Ulysses_, and _Erechtheus_ are strictly limited to
the Australian region of the archipelago, while five other groups are
with equal strictness confined to the Indian region, is a strong
corroboration of that division which has been founded almost entirely on
the distribution of Mammalia and Birds.
If the various Malayan islands have undergone recent changes of level,
and if any of them have been more closely united within the period of
existing species than they are now, we may expect to find indications of
such changes in community of species between islands now widely
separated; while those islands which have long remained isolated would
have had time to acquire peculiar forms by a slow and natural process of
modification.
An examination of the relations of the species of the adjacent islands
will thus enable us to correct opinions formed from a mere consideration
of their relative positions. For example, looking at a map of the
archipelago, it is almost impossible to avoid the idea that Java and
Sumatra have been recently united; their present proximity is so great,
and they have such an obvious resemblance in their volcanic structure.
Yet there can be little doubt that this opinion is erroneous, and that
Sumatra has had a more recent and more intimate connexion with Borneo
than it has had with Java. This is strikingly shown by the mammals of
these islands—very few of the species of Java and Sumatra being
identical, while a considerable number are common to Sumatra and Borneo.
The birds show a somewhat similar relationship; and we shall find that
the group of insects we are now treating of tells exactly the same tale.
Thus:—
Sumatra 21 sp.│20 sp. common to both islands;
Borneo 29 sp.│ „
──────────────┼──────────────────────────────
Sumatra 21 sp.│11 sp. common to both islands;
Java 27 sp.│ „
──────────────┼──────────────────────────────
Borneo 29 sp.│20 sp. common to both islands;
Java 27 sp.│ „
showing that both Sumatra and Java have a much closer relationship to
Borneo than they have each other—a most singular and interesting result
when we consider the wide separation of Borneo from them both, and its
very different structure. The evidence furnished by a single group of
insects would have had but little weight on a point of such magnitude if
standing alone; but coming as it does to confirm deductions drawn from
whole classes of the higher animals, it must be admitted to have
considerable value.
We may determine in a similar manner the relations of the different
Papuan Islands to New Guinea. Of thirteen species of Papilionidæ
obtained in the Aru Islands, five were also found in New Guinea, and
eight not. Of nine species obtained at Waigiou, five were New Guinea,
and four not. The five species found at Mysol were all New Guinea
species. Mysol, therefore, has closer relations to New Guinea than the
other islands; and this is corroborated by the distribution of the
birds, of which I will only now give one instance. The Paradise Bird
found in Mysol is the common New Guinea species, while the Aru Islands
and Waigiou have each a species peculiar to themselves.
The large island of Borneo, which contains more species of Papilionidæ
than any other in the archipelago, has nevertheless only two peculiar to
itself; and it is quite possible, and even probable, that one of these
may be found in Sumatra or Java. The last-named island has also two
species peculiar to it; Sumatra has not one, and the peninsula of
Malacca only one. The identity of species is even greater than in birds
or in most other groups of insects, and points very strongly to a recent
connexion of the whole with each other and the continent. But when we
pass to the next island (Celebes), separated from them by a strait not
wider than that which divides them from each other, we have a striking
contrast; for with a total number of species less than either Borneo or
Java, no less than eighteen are absolutely restricted to it. Further
east, the large islands of Ceram and New Guinea have only three species
peculiar to each, and Timor has five. We shall have to look, not to
single islands, but to whole groups, in order to obtain an amount of
individuality comparable with that of Celebes. For example, the
extensive group comprising the large islands of Java, Borneo, and
Sumatra, with the peninsula of Malacca, possessing altogether 45
species, has about 21, or less than half, peculiar to it; the numerous
group of the Philippines possess 21 species, of which 16 are peculiar;
the seven chief islands of the Moluccas have 27, of which 12 are
peculiar; and the whole of the Papuan Islands, with an equal number of
species, have 17 peculiar. Comparable with the most isolated of these
groups is Celebes, with its 24 species, of which the large proportion of
18 are peculiar. We see, therefore, that the opinion I have already
expressed, in the papers before quoted, of the high degree of isolation
and the remarkable distinctive features of this interesting island, is
fully borne out by the examination of this conspicuous family of
insects. A single straggling island, with a few small satellites, it is
zoologically of equal importance with extensive groups of islands many
times as large as itself; and standing in the very centre of the
archipelago, surrounded on every side with islets connecting it with the
larger groups, and which seem to afford the greatest facilities for the
migration and intercommunication of their respective productions, it yet
stands out conspicuous with a character of its own in every department
of nature, and presents peculiarities which are, I believe, without a
parallel in any similar locality on the globe.
Briefly to summarize these peculiarities, Celebes possesses three genera
of mammals (out of the very small number which inhabit it) which are of
singular and isolated forms, viz., _Cynopithecus_, a tailless Ape allied
to the Baboons; _Anoa_, a straight-horned Antelope of obscure
affinities, but quite unlike anything else in the whole archipelago or
in India; and _Babirusa_, an altogether abnormal wild Pig. With a rather
limited bird population, Celebes has an immense preponderance of species
confined to it, and has also five remarkable genera (_Meropogon_,
_Streptocitta_, _Enodes_, _Scissirostrum_, and _Megacephalon_) entirely
restricted to its narrow limits, as well as two others (_Prioniturus_
and _Basilornis_) which only range to a single island beyond it.
Mr. Smith’s elaborate tables of the distribution of Malayan Hymenoptera
(see ‘Proc. Linn. Soc.’ Zool. vol. vii.) show that, out of the large
number of 301 species collected in Celebes, 190 (or nearly two-thirds)
were absolutely restricted to it, although Borneo, on one side, and the
various islands of the Moluccas on the other, were equally well explored
by me; and no less than twelve of the genera are not found in any other
island of the archipelago. I have just shown in the present paper that,
in the Papilionidæ, it has far more species of its own than any other
island, and a greater proportion of peculiar species than many of the
large groups of islands in the archipelago—and that it gives to a large
number of the species and varieties which inhabit it, 1st, an increase
of size, and, 2nd, a peculiar modification in the form of the wings,
which stamp upon the most dissimilar insects a mark distinctive of their
common birth-place.
What, I would ask, are we to do with phenomena such as these? Are we to
rest content with that very simple, but at the same time very
unsatisfying explanation, that all these insects and other animals were
created exactly _as_ they are, and originally placed exactly _where_
they are, by the inscrutable will of their Creator, and that we have
nothing to do but to register the facts and wonder? Was this single
island selected for a fantastic display of creative power, merely to
excite a child-like and unreasoning admiration? Is all this appearance
of gradual modification by the action of natural causes—a modification
the successive steps of which we can almost trace—all delusive? Is this
harmony between the most diverse groups, all presenting analogous
phenomena, and indicating a dependence upon physical changes of which we
have independent evidence, all false testimony? If I could think so, the
study of nature would have lost for me its greatest charm. I should feel
as would the geologist, if you could convince him that his
interpretation of the earth’s past history was all a delusion—that
strata were never formed in the primeval ocean, and that the fossils he
so carefully collects and studies are no true record of a former living
world, but were all created just as they now are, and in the rocks where
he now finds them.
I must here express my own belief that none of these phenomena, however
apparently isolated pr insignificant, can ever stand alone—that not the
wing of a butterfly can change in form, or vary in colour, except in
harmony with, and as a part of, the grand march of nature. I believe,
therefore, that all the curious phenomena I have just recapitulated are
immediately dependent on the last series of changes, organic and
inorganic, in these regions; and as the phenomena presented by the
island of Celebes differ from those of all the surrounding islands, it
can, I conceive, only be because the past history of Celebes has been to
some extent unique and different from theirs. We must have much more
evidence to determine exactly in what that difference has consisted. At
present, I only see my way clear to one deduction, viz., that Celebes
represents one of the oldest parts of the archipelago, that it has been
formerly more completely isolated both from India and from Australia
than it is now, and that, amid all the mutations it has undergone, a
relic or substratum of the fauna and flora of some more ancient land has
been here preserved to us.
It is only since my return home, and since I have been able to compare
the productions of Celebes side by side with those of the surrounding
islands, that I have been fully impressed with their peculiarity, and
the great interest that attaches to them. The plants and the reptiles
are still almost unknown; and it is to be hoped that some enterprising
naturalist may soon devote himself to their study. The geology of the
country would also be well worth exploring, and its recent fossils would
be of especial interest as elucidating the changes which have led to its
present anomalous condition. This island stands, as it were, upon the
boundary-line between two worlds. On one side is that ancient Australian
fauna which preserves to the present day the facies of an early
geological epoch; on the other is the rich and varied fauna of Asia,
which seems to contain, in every class and order, the most perfect and
highly organized animals. Celebes has relations to both, yet strictly
belongs to neither; it possesses characteristics which are altogether
its own; and I am convinced that no single island upon the globe would
so well repay a careful and detailed research into its past and present
history.
In the following catalogue of the Malayan species of Papilionidæ I have
included those from Woodlark Island, collected by M. Montrouzier, as
that island comes fairly within the limits of the archipelago; while I
exclude New Caledonia as belonging more to the Australian and Pacific
fauna. I have given full particulars of the variation of the several
species, and have described all new species, forms, varieties, and
undescribed sexes. The distribution of each species is noted chiefly
from my own observations[11]. As the full synonymy and references to
almost every work on Lepidoptera are given in the British Museum List of
Papilionidæ, I have not thought it necessary to do more than to refer to
a good figure and description in well-known works; and I have quoted
Boisduval’s ‘Species Général des Lépidoptères’ throughout. In all cases,
however, where I have myself corrected the synonymy, or determined sexes
which had been before improperly located, I have given much fuller
references.
Footnote 11:
Species collected by myself have (Wall.) after the localities where I
have found them.
I have found it necessary to describe and name twenty new species, and
to separate six or seven more which have been hitherto considered as
varieties or sexes of other species. I have also described and separated
twenty-five local forms or races, and twenty polymorphous forms or
sexes, as well as several simple varieties. On the other hand, I have
reduced fourteen species, which figure in some of our latest lists, to
the rank of sexes or local or polymorphic forms of other species. For
convenience of reference, I add a list of these, with a reference to the
page where will be found the reasons for not adopting them.
Ornithoptera Pronomus, _G. R. Gray_, = O. Poseidon, _Db._ (var.), p. 36.
Ornithoptera Archideus, _G. R. Gray_, = O. Poseidon, _Db._ (var.), p. 36.
Ornithoptera Euphorion, _G. R. Gray_, = O. Poseidon, _Db._ (♀ var.), p. 36.
Ornithoptera Amphimedon, _Cr._, = O. Helena, _L._ ♀, p. 38.
Papilio Hegemon, _G. R. Gray_, = P. Polyphontes, _Bd._, p. 43.
Papilio Melanides, _De Haan_, = P. Theseus, _Fab._ (♀ form), p. 53.
Papilio Romulus, _Cr._, = P. Pammon, _L._ (♀ form), p. 52.
Papilio Rumanzovia, _Eschsch._, = P. Emalthion, _Hübn._ (♀ form), p. 48.
Papilio Polytes, _L._, = P. Pammon, _L._, ♀, p. 51.
Papilio Orophanes, _Bd._, = P. Ambrax, _Bd._, ♀, p. 54.
Papilio Elyros, _G. R. Gray_, = P. Alphenor, _Cr._ (♀ form), p. 53.
Papilio Amanga, _Bd._, = P. Ormenus, _Guér._ (♀ form), p. 55.
Papilio Onesimus, _Hewits._, = P. Ormenus, _Guér._ (♀ form), p. 55.
Papilio Drusius, _Cr._, = P. Gambrisius, _Cr._, ♀, p. 58.
As the arrangement of the species of _Papilio_ which I have adopted in
this paper is somewhat new, and I hope will be found to be more natural
than those which have been previously used, I here add lists of the
Indian and Australian species arranged in the same manner. Those already
included in my Malayan list will be indicated thus, (Mal.), and printed
in _italics_.
_List of the_ PAPILIONIDÆ _of the Indian Region_.
1. Teinopalpus imperialis, _Hope_.
2. Ornithoptera Darsius, _G. R. G._ (Ceylon).
3. —— Rhadamanthus, _Bd._
4. —— _Pompeus_, Cr. (Mal.).
5. —— _Amphrisius_, Cr. (Mal.).
_Papilio_ (Sect. A).
_Nox group._
6. Papilio _Varuna_, White (Mal.).
7. —— Aidoneus, _Db._
8. —— Philoxenus, _G. R. G._
9. —— Polyceutes, _Db._
10. —— Dasarada, _Moore_.
11. —— Ravana, _Moore_.
12. —— Minereus, _G. R. G._
13. —— Icarius, _Westw._
14. —— Bootes, _Westw._
15. —— Janaka, _Moore_.
_Coon group._
16. Papilio Doubledayi, _Wall._
_Polydorus group._
17. Papilio Jophon, _G. R. G._ (Ceylon).
18. —— _Diphilus_, Esp. (Mal.).
19. —— Alcinous, _Klug._
20. —— Mencius, _Feld._
21. —— Hector, _L._
_Papilio_ (Sect. B).
_Protenor group._
22. Papilio Protenor, _Cr._
23. Papilio Elphenor, _Db._
24. —— Rhetenor, _Westw._
25. —— Sakontala, _Hewits._
_Peranthus group._
26. Papilio Crino, _Fab._ (Ceylon).
27. —— Bianor, _Cr._
28. —— Polyctor, _Bd._
29. —— Ganesa, _Db._
30. —— Arcturus, _Westw._
31. —— Paris, _L._
32. —— Palinurus, _Fab._?
33. —— Krishna, _Moore_.
_Memnon group._
34. Papilio _Androgeus_, Cr. (Mal.).
35. —— Polymnestor, _Cr._ (Ceylon).
36. —— Demetrius, _Cr._
_Helenus group._
37. Papilio _Helenus_, L. (Mal.).
38. —— Chaon, _Westw._
39. —— Castor, _Westw._
40. —— _Nephelus_, Bd. (Mal.).
_Pammon group._
41. Papilio _Pammon_, L. (Mal.).
_Demolion group._
42. Papilio _Demolion_, Cr. (Mal.).
_Papilio_ (Sect. C).
_Erithonius group._
43. Papilio _Erithonius_, Cr. (Mal.).
_Paradoxa group._
44. Papilio Telearchus, _Hewits_.
45. —— Slateri, _Hewits_.
_Dissimilis group._
46. Papilio dissimilis, _L._
47. —— Panope, _L._
48. —— Lacedæmon, _Fab._
49. —— Pollux, _Westw._
_Papilio_ (Sect. D).
_Macareus group._
50. Papilio _Macareus_, God. (Mal.).
51. —— _Leucothoë_, Westw. (Mal.).
52. —— Megarus, _Westw._
53. —— Agestor, _G. R. G._
54. —— Epytides, _Hewits_.
55. —— Xenocles, _Db._
_Antiphates group._
56. Papilio _Antiphates_, Cr. (Mal.).
57. —— Agetes, _Westw._
58. —— Anticrates, _Db._
59. —— Orestes, _Fab._
60. —— Alebion, _G. R. G._
61. —— Glycerion, _G. R. G._
_Eurypylus group._
62. Papilio Gyas, _Westw._
63. —— Evan, _Db._
64. —— Cloanthus, _Westw._
65. —— _Sarpedon_, L. (Mal.).
66. —— Chiron, _Wall._
67. —— _Jason_, Esp. (Mal.).
68. —— _Agamemnon_, L. (Mal.).
69. —— _Rama_, Feld. (Mal.).
4. Chinese species.
61. Indian species.
4. Ceylon species.
_List of the_ PAPILIONIDÆ _of the Australian Region_.
_Ornithoptera (Priamus group)._
1. Ornithoptera _Poseidon_, Db. (Mal.).
2. —— Richmondia, _G. R. G._
_Papilio_ (Sect. A).
_Polydorus group._
3. Papilio _Leodamas_, Wall. (Mal.).
4. —— _Liris_, Godt. (Mal.).
5. —— Godartianus, _Bd._ (Pacific Islands).
_Papilio_ (Sect. B).
_Helenus group._
6. Papilio Capaneus, _Westw._
7. —— Ilioneus, _Don._
_Ulysses group._
8. Papilio Ulyssinus, _Westw._
9. —— Montrouzieri, _Bd._ (New Caledonia).
_Pammon group._
10. Papilio Canopus, _Westw._
_Erectheus group._
11. Papilio Erectheus, _Don._
12. —— Amyntor, _Bd._ (New Caledonia).
_Papilio_ (Sect. C).
_Erithonius group._
13. Papilio _Erithonius_, Cr. (Mal.).
_Anactor group._
14. Papilio Anactor, _McL._
_Papilio_ (Sect. D).
_Antiphates group._
15. Papilio Leosthenes, _Db._
16. —— _Permatus_, G. R. G. (Mal.).
_Eurypylus group._
17. Papilio _Sarpedon_, L. (Mal.).
18. —— Gelon, _Bd._ (New Caledonia).
19. —— Lycaon, _Westw._
20. —— Macleayanus, _Leach._
21. —— Scottianus, _Feld._ (Ash Islands).
22. Eurycus Cressida, _Fab._
6. Pacific Islands.
16. Australia.
_Catalogue of Malayan_ PAPILIONIDÆ.
ORNITHOPTERA (Boisd.).
[Illustration:
Fig. 1.
Anal valves of _O. Amphrisius_.
]
The characters in the _larva_ and _pupa_ which have been supposed to
distinguish this genus from PAPILIO are erroneous, or at least do not
exist in all the species. My own observations on _O. Poseidon_ show that
the _larva_ has no “external sheath” to the thoracic tentacles, and that
the suspending thread passes round the pupa, and is not “fastened on
each side to a silky tubercle.” There remain therefore only the
characters of the perfect insect, the most important of which are the
anal valves in the male. These are very large, ovate or rounded,
coriaceous, and not hairy, and are furnished with projecting points or
spines (sometimes very conspicuous) which serve to attach the male more
firmly to the female _in copulâ_. In several species I have observed,
these points or hooks were buried in the protruded anal gland of the
female, and thus effectually prevented the great weight of the insects
causing them to separate upon suddenly taking flight. The great strength
and size of these insects, the thick texture of their wings, their long
curved and stout antennæ, their peculiar form, colour, and distribution,
are the only other characters that separate them from _Papilio_. Though
these may not perhaps be technically sufficient, I think it advisable
and convenient to retain a genus so well known and long established.
_Ornithoptera_ is pre-eminently a Malayan genus, seventeen species
inhabiting the archipelago, one (_Rhadamanthus_, Bd.) India and China,
one (_Darsius_, G. R. Gray) peculiar to Ceylon, one (_Richmondia_, G. R.
Gray) North Australia. _O. Victoriæ_, G. R. Gray, from some island east
of New Guinea, should probably be included in the Malayan list; and
_Æacus_, Felder, from an unknown locality. The following are the
well-established Malayan species.
a. _Priamus_ group.
1. ORNITHOPTERA PRIAMUS, Linnæus.
♂. _Papilio Priamus_, L.; Cram. Pap. Ex. t. 23. f. A, B; Godart,
Enc. Méth. ix. p. 25. _O. Priamus_, Bd. Sp. Gén. Lép. p. 173.
♀. _P. Panthous_, L.; Cram. Pap. Ex. t. 123. f. A, t. 124. f. A.
This may be at once distinguished from all the allied species with which
it has been often confounded—in the _male_, by the more rounded and
deeply scalloped hind wings, with larger black spots and a broader
border, the upper wings with no green on the median nervure or its
branches, and the sooty patch extending only to the second median
nervule; in the _female_, by the very constant and peculiar light
olive-brown colour, the absence of any spots in the discoidal cell of
the upper wings, and the broad shallow scallops of the hinder margin.
_Hab._ Amboyna and Ceram, probably also Bouru (_Wall._).
2. ORNITHOPTERA POSEIDON, Doubleday.
_O. Poseidon_, Db. Ann. of Nat. Hist. xvi. p. 173; Westwood, Cat. of
Orient. Ent. pl. 11, 14.
The numerous specimens of _Ornithoptera_ which I obtained in various
parts of New Guinea and the adjacent islands show so much instability of
form, colouring, and even of neuration, no two individuals being exactly
alike, that I am obliged to include them all in one variable species, to
which I believe must also be referred _O. Pronomus_, G. R. Gray, from
Cape York, _O. Euphorion_, G. R. Gray, from North Australia, _O.
Archideus_, G. R. Gray (ex Boisd.), erroneously said to be from Celebes,
and _O. Boisduvalii_, Montrouzier, from Woodlark Island.
Var. _a_, Aru Islands (_Wall._). _O. Arruana_, Feld. Lep. Frag. p. 24.
Individuals from this locality differ in the arrangement of the
nervures; in some the third subcostal nervure of the upper wings
branches from the same point with the upper disco-cellular, in others
considerably beyond it; the points from which the subcostal nervures
branch also vary. The amount of green colour on the median nervure and
its branches varies. In some specimens there is a spot at the anal angle
of lower wings beneath, agreeing with _O. Pronomus_, G. R. Gray; but
this is generally wanting.
Var. _b_, Dorey, Salwatty, south-west coast of New Guinea (_Wall._).
These agree very closely with _O. Poseidon_, as figured by Westwood;
they differ individually in the same manner as the last, and also in the
length of the lower disco-cellular nervure on the under wings. They have
generally no golden spots beneath the wings. They vary also in the
outline of the under wings, the outer and anal angles being more acute
in some specimens than in others. Some have the under wings of a uniform
green entirely without spots, while others have a range of black spots
more or less fully developed.
Var. _c_, Waigiou (_Wall._). _Archideus_, G. R. Gray, ♀.
This agrees with the last; but the male is of a more delicate green than
any of the others, and has less of that colour on the median veins. On
the under side there are no golden spots. The whole surface has a golden
tinge, and the central portion of the lower wings is tinged with
amber-brown.
The females of all the above vary extremely, much more even than the
males, and from the same locality two specimens are rarely alike. The
discoidal cell is in some specimens more than half occupied by a whitish
patch, while in others there are only a few small spots. One of my
specimens from Salwatty almost exactly agrees with that figured by
Westwood (Cat. of Or. Ent. pl. 14) as from Cape York. One of the Waigiou
specimens is the same as _Archideus_, G. R. G., figured by Boisduval
(Voy. de l’Astrolabe, t. 4. f. 1, 2); and another, from New Guinea,
differs very little from _Euphorion_, G. R. G. (Brit. Mus. Cat. Lep. pt.
1. pl. 2. f. 3), from North Australia.
From these facts I am led to conclude that we have here a variable form
spread over an extensive area, and kept variable by the continual
intercrossing of individuals, which would otherwise segregate into
distinct and sharply defined races. The same area is inhabited by many
species of birds common to all parts of it; and just as the birds of
Ceram and Amboyna are almost all distinct species from those of New
Guinea, so do we find those islands inhabited by the _Ornithoptera
Priamus_, a well-marked and constant species, readily distinguishable in
either sex from the inconstant forms of New Guinea proper. The same
parallel holds in North Australia. Many New Guinea species of birds
extend, with very slight variation, to the country about Cape York; but
when we reach the Moreton Bay district all these have disappeared, and
we find only true Australian species. So the variable forms of _O.
Poseidon_ reach North Australia and Cape York, while in the Moreton Bay
district we find the comparatively well-marked species _O. Richmondia_.
Similar causes, whether geographical or climatal, have thus produced an
analogous distribution in these widely separated groups of animals.
3. ORNITHOPTERA CRŒSUS, Felder.
_O. Crœsus_, Feld. Wien. Ent. Monats., Dec. 1859. _O. Crœsus_, G. R.
Gray, Proc. Zool. Soc. 1859, p. 424.
_Hab._ Batchian (Moluccas) (_Wall._).
Local form, _a._—_Male_: has the orange colour of the upper surface of a
much deeper fiery-red hue; on the under surface, the black spots of the
lower wings are nearer the margin, and the yellow spots below them are
entirely absent; there is also a green line between the subcostal
nervure and the margin; on the under surface of the fore wings the green
patch in the discoidal cell extends to its base, and is reflexed in a
narrow line along its upper edge.
_Female_: differs still more from that sex in _O. Crœsus_; the white
markings on all the wings are so large as almost to fill up the spaces
between the veins, the lower part of the discoidal cell in both upper
and under wings being also occupied with a whitish patch; the range of
spots occupying the posterior margin are of a dusky yellow colour.
_Hab._ Ternate (♂), Gilolo (♀) (_Wall._).
This well-marked local form is no doubt peculiar to Gilolo and the small
adjacent islands, as the original species is to Batchian.
I was three months in the island of Batchian before I obtained a
specimen of this fine insect, which I had seen once or twice only flying
high in the air. I at length came upon it flying about a beautiful
cinchonaceous shrub with white bracts and yellow flowers (_Mussænda_,
sp.); and having cleared a path round about, I visited the place every
morning on my way to the forest, and once or twice a week had the
satisfaction of capturing a fine male specimen of _O. Crœsus_. The
females were more plentiful and more easily caught. I afterwards sent
out one of my men with a net every day to look after this insect only.
He would stay out all day long, wandering up a broad rocky torrent,
where the males flew up and down occasionally or settled on the rocks
which just appeared above the water. He generally brought me one, and
sometimes even two or three specimens; and thus, with those that I
myself captured at the flowers, I secured a fine series of this richly
coloured species.
4. ORNITHOPTERA TITHONUS, De Haan.
_O. Tithonus_, De Haan, Verh. Nat. Gesch. Ned. t. 1. f. 1.
_Hab._ S.W. Coast of New Guinea (_Leyden Museum_).
This remarkable species must be very rare, as I never saw it in any part
of the New Guinea district that I visited; nor was it seen during the
exploration, a few years ago, by a Dutch steamer which visited the part
of the coast where the only specimen known was said to have been
obtained.
5. ORNITHOPTERA URVILLIANA, Guérin.
_Papilio Urvilliana_, Guér. Voy. de la Coquille, Lép. t. 13. f. 1, 2,
♂.
_O. Urvilliana_, Boisd. Sp. Gén. Lép. p. 175.
_Hab._ New Ireland (_Paris Museum_).
b. _Pompeus_ group.
6. ORNITHOPTERA REMUS, Cramer.
_Papilio Remus_, Cr. Pap. Ex. t. 135. f. A, t. 136. f. A (♀), t. 386.
f. A, B (♂); Fab. Syst. Ent. iii. 1. p. 11.
_O. Remus_, Bd. Sp. Gén. Lép. p. 176. _Papilio Panthous_ ♂, Clerck,
Icon. t. 18 (♀).
_Hab._ Amboyna, Ceram, Gilolo, Morty Island, Sulla Island, Celebes
(_Wall._).
The specimens above quoted agree well with Cramer’s figures. The female
from the Sulla Islands differs only in having more yellow towards the
anal angle of the lower wings. The specimens figured by Cramer in pls.
10, 11, under the name of “_Hypolitus_” seem to be a remarkable variety,
in which the female has much of the character of the male. Messrs.
Doubleday and G. R. Gray have adopted _Panthous_ as the specific name of
this insect; but this name was first used by Linnæus for the female of
_Priamus_ only, in the 10th ed. of the ‘Systema Naturæ’ (1758). Clerck
(in 1759) adopted the name, but supposed he had found the male in the
female of _Remus_. Linnæus, in Mus. Lud. Ulric. (1764), and in the 12th
ed. of the ‘Systema Naturæ’ (1766), adopts this error, so far as
referring to Clerck’s two figures; but in both these works his
description refers only to the female of _P. Priamus_, indicating that
the supposed other sex (_P. Remus_) was not known to him personally. The
name of _Panthous_ must therefore altogether drop, it having been
applied to this species only through a double error—first, that of
Linnæus, in supposing his _Panthous_ to be distinct from _Priamus_, and
then that of Clerck, in thinking that a female _Remus_ was the male of
the Linnean _Panthous_.
7. ORNITHOPTERA HELENA, Linnæus.
♂. _P. Helena_, Cram. Pap. Ex. t. 140. f. A, B. _O. Helena_, Boisd.
Sp. Gén. Lép. p. 177.
♀. _P. Amphimedon_, Cram. Pap. Ex. t. 194. f. A. _O. Amphimedon_,
Boisd. Sp. Gén. p. 176.
_Hab._ Amboyna and Ceram (_Wall._).
The females from these localities are always sooty, with the spots and
markings on the hinder wings of a dull buff-colour even in the freshest
specimens.
_a._ Local form _Bouruensis_.—_Male_: exactly resembles the Amboyna
specimens, except that the yellow patch is more variable in form and
extent.
_Female_: nearly black, and with the markings on the lower wings almost
as pure and deep yellow as in the males: size a little smaller than in
the type.
_Hab._ Bouru (_Wall._).
_b._ Local form _Papuensis_.—_Female_: sooty black, the two first
branches of the subcostal nervure margined with whitish near their
origin; markings of the lower wings of the same tint of orange-yellow as
is _O. Helena_ ♂, but not so glossy.
_Male_ not known.
_Hab._ New Guinea, Salwatty (_Wall._).
_c._ Local form _Celebensis_.—_Male_: wings a little more pointed than
in _O. Helena_; yellow patch of lower wings extending nearer to the
posterior margin, and bounded towards the abdominal margin by the first
branch of the median nervure. Beneath, having the nervures between the
discoidal cell and the outer border ashy-margined.
_Female_ not known.
_Hab._ Macassar (Celebes) (_Wall._).
_Remarks._—Of these three local modifications of _O. Helena_, the first
is very distinct in the female, but not separable in the male sex. Of
the second and third, only one sex is known; and they may very probably
prove to be well-marked species when more materials are obtained.
8. ORNITHOPTERA LEDA, n. s.
_Male_: upper wings elongate, triangular, glossy black, quite uniform
and immaculate; the outer margin delicately white-marked at the
termination of the nervures. Lower wings yellow, as in the allied
species, with a black border about the same width as in _O. Pompeus_ on
the outer and abdominal margins, narrower on the inner margin; the
posterior scalloping of the yellow patch not so deep as in _O. Pompeus_,
and having a spot at the anal angle connected more or less with the
margin.
The under surface differs from that of _O. Pompeus_ by the ashy margins
of the veins of the upper wings being entirely absent, and in having
much less white on the outer edge. There are no submarginal spots except
the anal one, much red at the base of the wings, and no black spots on
the abdomen.
_Female_: this sex varies very much, some having the upper wings
immaculate, while others have the veins about the end of the discoidal
cell broadly margined with whitish. The marginal series of spots on the
lower wings vary as they do in _O. Pompeus_ and _O. Amphrisius_. The
best distinction from _O. Pompeus_ (♀) seems to be the more elongated
wings, the less crenellated margin, and the more produced outer angle of
the lower wings. The yellow patch is also of a deeper colour both on the
upper and under surfaces.
_Hab._ Celebes (Macassar and Menado) (_Wall._)
9. ORNITHOPTERA POMPEUS, Cramer.
_P. Pompeus_, Cr. Pap. Ex. t. 25. f. A (♂). _P. Minos_, Cr. Pap. Ex.
t. 195. f. A (♀). _P. Heliacon_, Fab. Ent. Syst. 3. i. p. 19, 60.
_O. Heliacon_, Boisd. Sp. Gén. Lép. p. 178.
_Hab._ Sumatra, Borneo, Java, Lombock (_Wall._), India (var.).
_Remark._—The form that occurs in India, in its more elongate wings and
darker colouring, approaches very closely to _O. Rhadamanthus_.
10. ORNITHOPTERA NEPHEREUS, G. R. Gray.
_P. Astenous_, Eschscholtz, Voy. Kotzebue, t. 4. f. A, B, C. (nec
Fab.).
_O. Nephereus_, G. R. G., List of Lep. B. M. Papilionidæ, p. 6.
_Hab._ Philippine Islands.
_Remark._—This is quite distinct from _O. Rhadamanthus_, Bd., with which
it has generally been identified.
11. ORNITHOPTERA MAGELLANUS, Felder.
_O. Magellanus_, Feld. Lep. Nov. Phil. p. 11.
_Hab._ North of Luzon (Philippines).
_Remark._—This fine species has a beautiful opalescent glow on the lower
wings when viewed obliquely.
12. ORNITHOPTERA CRITON, Felder.
_O. Criton_, Feld. Lep. Fragm. p. 49.
_Hab._ Batchian, Ternate, Gilolo, Morty Island (_Wall._).
13. ORNITHOPTERA PLATO, n. s.
_Male_: resembles _O. Criton_ in the form and extent of the yellow
patch, but the upper wings differ in having the outer half of a lighter
tint; on the under surface this outer half of the wing is of a light
ash-colour. Abdomen almost wholly black beneath. No red patches at the
base of the wings, or any red collar.
_Female_ unknown.
_Hab._ Timor (_Wall._).
This is a very distinct species, though at first sight resembling
several others. I obtained a single male specimen only.
14. ORNITHOPTERA HALIPHRON, Boisduval.
_O. Haliphron_, Bd. Sp. Gén. Lép. p. 181 (♂); Felder, Lep. Fragm. p.
37, Taf. ii. f. 2, 3 (♂, ♀).
_Hab._ Macassar (Celebes) (_Wall._).
15. ORNITHOPTERA AMPHRISIUS, Cramer.
_P. Amphrisius_, Cr. Pap. Ex. t. 219. f. A; Godardt, Enc. Méth. ix. p.
27, pt.
_O. Amphrisius_, Bd. Sp. Gén. Lép. p. 178.
_Hab._ Malacca, Java, Borneo (_Wall._).
This may be readily distinguished from the allied species by the upper
wings in the male being yellow-marked, and by the absence of red spots
at the base of the wings beneath in both sexes.
c. _Brookeana_ group.
16. ORNITHOPTERA BROOKEANA, Wallace.
_O. Brookeana_, Wall. Proc. Ent. Soc. 1855, p. 104; Hewitson, Ex.
Butt. Papilionidæ, i. f. 1. _Papilio Trogon_, V. Voll. Tijdschrift
voor Ent. 1860, p. 69, pl. 6.
_Hab._ Borneo (Sarawak) (_Wall._), Sumatra (_Leyden Museum_).
_Remarks._—I have been in much doubt about the position of this
remarkable species, and was for some time inclined to place it among the
Papilios. It agrees, however, far better with _Ornithoptera_ in the form
and stoutness of the wings, the long stout and curved antennæ, the red
collar and patches at the base of the wings beneath, the abdominal fold,
and the flight and general appearance. It is powerful on the wing, and
occasionally settles on the ground in damp sunny places. It inhabits the
interior of North-west Borneo and the mountains of West Sumatra. The
female is unknown. It is peculiar in the great length of the discoidal
cell of the wings and its altogether unique style of coloration, and
must be considered as the type of a distinct group of the genus
_Ornithoptera_.
PAPILIO.
This is without doubt the finest and most remarkable genus of Diurnal
Lepidoptera. About 360 species are now known, all, except ten, being
tropical or subtropical. I have given at p. 23 the characters of the
sections and groups into which I divide the Malayan species.
SECTION A.
a. _Nox_ group.
17. PAPILIO NOX, Swainson.
_P. Nox._, Sw. Zool. Ill. pl. 102; Horsf. Lep. Ins. E. I. C. pl. 1. f.
1; Boisd. Sp. Gén. Lép. p. 277.
_P. Neesius_, Zink. Nov. Act. Acad. Nat. Cur. xv. t. 14. f. 1.
_Hab._ Java (♂, ♀) (_Wall._), Penang (♂) (_Brit. Mus._).
18. PAPILIO NOCTIS, Hewitson. Tab. V. fig. 1 (♂)[12].
Footnote 12:
In all the Plates, the wings on one side of each figure are detached
from the body, and represent the _under surface_ of the same insect.
In one case only (Tab. VII. f. 1.) the upper surfaces of two varieties
of the same species are given.
_P. Noctis_, Hewits. Proc. Zool. Soc. 1859, p. 423, pl. 66. f. 5 (♀).
_Male_: differs from the same sex of _P. Nox_ by the broader apex of the
fore wings, and by the hind wings being more elongate, more glossy, and
especially by the entire nondentated hinder margin.
_Hab._ Borneo (Sarawak) (_Wall._), (♂, ♀ Mus. nost.)
19. PAPILIO EREBUS, Wallace.
_P. Nox_, var., De Haan, Verh. Nat. Gesch. t. 5. f. 3 (♀).
_Hab._ Malacca (_Wall._), Banjermassing, Borneo (_De Haan_).
_Remarks._—I am somewhat doubtful of the species, the female only being
known; but it differs so strikingly from the same sex of _P. Nox_ and
_P. Noctis_ (the former of which seems very constant), that I think it
better to separate it in order to draw attention to other specimens that
may exist in collections. It differs from _P. Nox_ (♀) by its narrower
and more elongate hind wings, which are black, glossed with steel-blue;
the fore wings are black, with the veins beyond the cell clearly
white-margined. The lower margin is also much less strongly dentated.
20. PAPILIO VARUNA, White.
♀. _P. Varuna_, Wh., Entomologist, 1842, p. 280; Westw. Ann. Nat.
Hist. ix. p. 37. _P. Chara_, Westw. Arc. Ent. pl. 66. f. 2.
♂. _P. Astorion_, Westw. Ann. Nat. Hist. ix. p. 37; Arc. Ent. pl. 66.
f. 1.
_Hab._ Pulo Penang, Sylhet.
21. PAPILIO SEMPERI, Felder.
_P. Semperi_, Feld. Lep. Nov. Philipp, pp. 1, 11.
_Hab._ Luzon, Philippines (♂, ♀).
N.B. The _Philoxenus_ group peculiar to India follows on after these.
b. _Coon_ group.
22. PAPILIO NEPTUNUS, Guérin.
_P. Neptunus_, Guér. Deless. Voy. dans l’Inde, p. 69, t. 19 (_P.
Saturnus_).
_Hab._ Malacca, Borneo (♂, ♀) (_Wall._).
23. PAPILIO COON, Fabricius.
_P. Coon_, Fab. Ent. Syst. iii. 1. pp. 10, 27; Don. Ins. China, pl.
24. f. 1; Lucas, Lep. Ex. t. 6. f. 2; Boisd. Sp. Gén. Lép. p. 201.
_Hab._ Java, Sumatra (_Wall._), Borneo (_De Haan_).
_Remarks._—The specimens from Sumatra are constantly larger than those
from Java. The Indian form, in which the markings are red instead of
yellow, with other differences, I consider a distinct species, for which
I propose the name of _P. Doubledayi_, after the late Mr. Edward
Doubleday of the British Museum[13].
Footnote 13:
PAPILIO DOUBLEDAYI, Wallace. (_P. Coon_, var., B. M. Cat.)
_Above_: upper wings as in _P. Coon_, but the base darker. Lower wings
broader than in _P. Coon_; the white spot in the cell toothed below,
and divided by one or two faint blackish lines, cut off at the middle
of the cell by the black triangular basal patch. The marginal spot
next within the tail wanting; the two anal spots, end of abdomen, and
its rings (which are yellow in _P. Coon_) red; collar behind the eyes
and palpi (which are black in _P. Coon_) also red.
_Beneath_: base of lower wings broadly black; white spots all much
broader and rounder than in _P. Coon_; sides of the thorax, end of the
abdomen, and the marginal spots in the caudal and anal region red.
The female differs in a corresponding manner from _P. Coon_ ♀. Size
about the same.
_Hab._ Moulmein, Assam.
c. _Polydorus_ group.
24. PAPILIO POLYDORUS, Linnæus.
_P. Polydorus_, L.; Clerck, Icon. t. 33. f. 3. _P. Leobates_, Reinw.
Verh. Nat. Gesch. Zool. t. 6. f. 3 (♀).
_Hab._ Ceram, Matabello Island, Bouru, Batchian (♂, ♀) (_Wall._).
Local form or variety _a_.—The white markings on the fore wings forming
a patch below the cell; red spots on the hind wings nearer to the
posterior margin and that next the anal angle larger.
_Hab._ Ké Island, Aru Island (♂, ♀) (_Wall._).
25. PAPILIO LEODAMAS, n. s. Tab. V. fig. 2 (♂).
_P. Polydorus_, in Brit. Mus. List of Papilionidæ, p. 10.
_Male._ Above, glossy black, upper wings immaculate (the veins
pale-margined in the female). Lower wings with a rounded white spot
divided into six parts by fine nervures, of which the outermost and that
in the cell are sometimes reduced to points; marginal row of red spots
obscured with black, and but faintly indicated.
Beneath, the white patch has a small red spot attached to the part next
the anal angle; and the marginal row of six red spots are clearly
marked, that at the anal angle being twice the size of the rest. Wings
short, much rounded, scarcely or not at all produced in the caudal
region.
Expanse of wings 3¾ in. to 4 in.
_Hab._ New Guinea, Mysol (♂, ♀) (_Wall._), Rockingham Bay (Australia),
(_Brit. Mus._, ♀).
26. PAPILIO DIPHILUS, Esper.
_P. Diphilus_, Esp. Ausl. Schmett. t. 40. f. 1. P. _Polydorus_, Boisd.
Sp. Gén. Lép. p. 267; and most authors.
_Hab._ Java, Malacca (_Wall._), Philippine Islands, India.
_Remarks._—The specimens from Manilla are larger, and the females
paler-coloured, than those from other localities, all of which have
slight characteristic peculiarities; but they also vary in the
individuals from each locality, so that no perfect segregation of local
forms has yet taken place.
27. PAPILIO ANTIPHUS, Fabricius.
_P. Antiphus_, Fab. Syst. Ent. iii. 1. pp. 10–28; Boisd. Sp. Gén. Lép.
p. 266.
_Hab._ Sumatra, Borneo, Lombock, Java (_Wall._), Philippine Islands.
_Remarks._—The Philippine form (_P. Kotzebuea_, Eschsch.) is rather
larger and of a more uniform glossy black than those from other
localities. _P. Theseus_, Cram., has been erroneously supposed to be the
female of this species, whereas it is the female of one of the _Pammon_
group, belonging to a different section of the genus. De Haan figures
_P. Theseus_ as _P. Antiphus_ ♀, in Verh. Nat. Gesch. t. 8. f. 2. As has
been already pointed out, _P. Theseus_ mimics this species.
28. PAPILIO POLYPHONTES, Boisduval.
_P. Polyphontes_, Bd. Sp. Gén. Lép. p. 268. _P. Hegemon_, G. R. G.,
List of Papilionidæ in B. Mus.
_Hab._ Celebes, Batchian, Morty Isl. (♂, ♀) (_Wall._).
_Remarks._—The markings vary from pure white to a smoky tint; but
otherwise all the specimens from the above localities agree. De Haan
gives (Verh. Nat. Gesch. t. 8. f. 4) a female of one of the _Pammon_
group as _P. Polyphontes_ ♀.
29. PAPILIO ANNÆ, Felder.
_P. Annæ_, Feld. Lep. Nov. Philipp, p. 1.
_Hab._ Mindoro (Philippines).
30. PAPILIO LIRIS, Godart.
_P. Liris_, God. Enc. Méth. iv. p. 72; Boisd. Sp. Gén. Lép. p. 268; De
Haan, Verh. Nat. Gesch. p. 38, t. 4. f. 3 (♀).
_Hab._ Timor (_Wall._), N.W. Australia (_Brit. Mus._).
_Remarks._—The Australian specimens are smaller. The female of _P.
Œnomaus_ mimics this species, as has been already mentioned (p. 22).
Both species were taken by myself on the same spot, and, though such
large and conspicuous insects, they could never be distinguished without
a close examination after capture. The female of this species differs
very little from the male, being rather larger, with broader wings and
less vivid coloration.
SECTION B.
d. _Ulysses_ group.
31. PAPILIO ULYSSES, Linnæus.
_P. Ulysses_, L., Cramer, Pap. Ex. t. 121. f. A, B (♀), t. 122 A (♀).
_P. Diomedes_, Boisd. Sp. Gén. Lép. p. 202.
_Hab._ Amboyna, Ceram (♂, ♀) (_Wall._).
_Remark._—The largest specimens of this glorious insect are found in the
island of Amboyna, where it is rather common, hovering about the forest
pathways. It sometimes visits the gardens in the town of Amboyna.
32. PAPILIO PENELOPE, n. s.
_Male_: rather smaller than _P. Ulysses_. Upper wings with six black
cottony patches, and all separate from each other; whereas in _P.
Ulysses_ there are seven, and the four lower ones are always united at
their margins. The blue colour fills the discoidal cell, and generally
extends beyond it at the extremity; the upper disco-cellular nervure not
black-bordered as in _P. Ulysses_. Lower wings with the blue colour
extending further along the abdominal margin, and not quite so far
towards the outer angle.
_Female_: has the blue colour of the same form and extent as in _P.
Ulysses_ ♀, but of the same bright tint as in the male; the marginal
lunules more deeply curved.
Expanse of wings 5 inches.
_Hab._ New Guinea, Waigiou, Aru Is. (♂, ♀) (_Wall._).
_Remark._—As all the other forms closely allied to _P. Ulysses_ have
received names (_Telemachus_, Montr., _Chaudoiri_, Feld., _Telegonus_,
Feld., and _Ulyssinus_, Westw.), I have also given one to this form
peculiar to New Guinea and the Papuan Islands, the distinctive
characters of which, though very slight, seem sufficiently constant.
33. PAPILIO TELEGONUS, Felder.
_P. Telegonus_, Feld. Lep. Fragm. p. 50.
_Hab._ Batchian, Gilolo (♂, ♀) (_Wall._).
_Remark._—A very distinct species, separated from _P. Ulysses_ by the
extent of the cottony patch on the upper wings, and by the different
form and colour of the blue markings.
34. PAPILIO TELEMACHUS, Montrouzier.
_P. Telemachus_, Mont. Ann. de la Soc. d’Agriculture de Lyon, 1856, p.
395.
_Hab._ Woodlark Isl. (S. E. of New Guinea).
_Remark._—This is a small species (exp. 4 in.), with less blue on the
lower wings.
e. _Peranthus_ group.
35. PAPILIO PERANTHUS, Fabricius.
_P. Peranthus_, Fab. Syst. Ent. iii. 1. p. 15; Don. Ins. China, pl.
26; Lucas, Lep. Ex. t. 12. f. 2; Boisd. Sp. Gén. Lép. p. 203.
_Hab._ Java, Lombock (_Wall._).
36. PAPILIO PERICLES, n. sp. Tab. VI. fig. 1 (♂).
Wings more elongate, and upper wings more pointed, than in _P.
Peranthus_.
_Above_: black, the basal half of a silvery blue, greenish towards the
base of the costa, and purplish on the outer margin, where on the lower
wings it shades off into separate scales. On the submedian and two lower
branches of the median nervure are elongate black cottony patches as in
_P. Ulysses_, the lower ones joined at the base, the upper one separate;
above these the outer margin is of a brown-black, with a few atoms of
yellow and blue scales towards the apex; the blue colour extends beyond
the discoidal cell of the upper wings in a line parallel with the outer
margin, on the lower wings it rounds away to the anal angle, and below
it are five submarginal lunules of blue atoms, the outer one almost
obsolete, and that next the tail largest and most deeply coloured.
Thorax and body green.
_Beneath_: as in _P. Peranthus_, but the posterior range of lunules
margined with brilliant blue and orange brown.
Expanse of wings 3½ inches.
_Hab._ Timor (♂) (_Wall._).
37. PAPILIO PHILIPPUS, Wallace. Tab. VI. fig. 3.
_P. Peranthus_, var. A, Boisd. Sp. Gén. Lép. p. 204.
_Above_: basal half of the wings of a rich green-blue, the rest black,
with a triangular patch at the apex of the uppers, formed of green atoms
situated between the nervures; on the lower wings six large submarginal
lunules, the lowest of which sends out some green atoms along the tail.
The black cottony spot is of a different form from that of _P.
Peranthus_, the separate patches being only joined in the middle, and
two of them extending along the nervures in a point nearly to the
discoidal cell.
_Beneath_: brilliantly marked with lunules of buff, black, and blue.
Expanse of wings 4½–5 inches.
_Hab._ Moluccas (_Wall._).
_Remarks._—My specimen from Ceram is of a greener tinge, and the colour
extends a little beyond the end of the discoidal cell; that from
Batchian is smaller, of a bluer tinge, and the colour of less extent.
The species seems to be very rare.
38. PAPILIO MACEDON, Wallace. Tab. VI. fig. 2 (♂).
_P. Peranthus_, var. B., Boisd. Sp. Gén. Lép. p. 204.
Boisduval’s description sufficiently shows the remarkable differences of
form, size, and colouring which this species presents, compared with
that of which he considers it a variety. The female agrees with the
male, except that the colours are a little less brilliant, and the
cottony patches of the fore wings are absent.
Expanse of wings, ♂, 5 inches; ♀, 5–6 inches.
_Hab._ Macassar, Menado (Celebes) (_Wall._).
39. PAPILIO BRAMA, Guérin.
_P. Brama_, Guér. Rev. Zool. 1840, p. 43, t. 1. f. 3, 4. _P.
Palinurus_, De Haan, Verh. Nat. Gesch. pp. 5, 29.
_Hab._ Malacca, Sumatra (_Wall._).
40. PAPILIO DÆDALUS, Felder.
_P. Dædalus_, Feld. Lep. Nov. Philipp. p. 2.
_Hab._ Luzon (Philippine Islands).
41. PAPILIO BLUMEI, Boisduval. Tab. VI. fig. 4 (♂).
_P. Blumei_, Boisd. Sp. Gén. Lép. p. 206.
_Hab._ Menado (Celebes) (_Wall._). “Amboyna,” _Bd._, error of
locality.
_Remark._—This very fine species comes nearest to the last, but is of
much larger size, and is conspicuous by its brilliantly coloured tails.
42. PAPILIO ARJUNA, Horsfield.
_P. Arjuna_, Horsf. Cat. Lep. E. I. Comp. pl. 1. f. 14; Boisd. Sp.
Gén. Lép. p. 209. _P. Arjuna_, var. _a._, Brit. Mus. Cat. of
Papilionidæ, p. 17.
_Hab._ Java, Borneo, Sumatra (_Wall._).
The Bornean form differs from that of Java by its larger size, and on
the under surface by the three middle lunules being formed of a violet
line only, with scarcely a trace of red beneath it, and by the
orange-red lunules both at the anal and outer angles being divided (not
margined) by a violet line. The scales sprinkled at the base of the
lower wings are white and blue, and are neither so dense nor do they
extend so far as the yellowish scales of the Java specimens. In all
these particulars the Sumatra specimens are somewhat intermediate, but
approach most to those of Borneo. This is one of the examples which show
the isolation of Java, notwithstanding its proximity to Sumatra.
f. _Memnon_ group.
(N.B. The _Protenor_ group of India is intermediate between this and the
last group.)
43. PAPILIO MEMNON, Linnæus. Tab. I. figs. 1 (♂), 2, 3, 4 (♀s).
♂, _P. Memnon_, L., Cram. Pap. Ex. t. 91. f. C (♂); Boisd. Sp. Gén.
Lép. p. 192.
♀, 1st dimorphic form, _P. Anceus_, Cr. Pap. Ex. t. 222. f. A, B.
? _P. Laomedon_, Cr. Pap. Ex. t. 50. f. A, B; De Haan, Verh. Nat.
Gesch. p. 24, t. 3. f. 2.
♀, 2nd dimorphic form, _P. Achates_, Cr. Pap. Ex. t. 243. A.
_Hab._ Java, Sumatra (_Wall._).
Local form _a_.—_Male_: border of posterior wings beneath narrow and of
an ashy-blue colour.
_Female_: near _P. Anceus_, Cr., and _P. Laomedon_, Cr., but of an
olive-ashy colour.
_Hab._ Borneo (_Wall._).
Local form _b_.—_Male_: band on under side of posterior wings ashy; the
spots large, with reddish-orange lunules between the two series, and
below the four outer ones.
_Hab._ Lombock (_Wall._).
_Remarks._—The difference between the male and the 2nd form of female is
so great, both in form and colouring, that they could not have been
imagined to be the same, had they not been bred from the same larvæ.
They have also been taken “_in copulâ_” by myself. Each form varies
considerably, both individually and locally; yet there are none
intermediate between the two. I consider them, therefore, as presenting
a fine instance of dimorphism; and I also believe that the second form
mimics _P. Coon_, for reasons which I have explained at p. 21.
44. PAPILIO ANDROGEUS, Cramer.
♂, _P. Androgeus_, Cr. Pap. Ex. t. 91. f. A, B.
♀, 1st dimorphic form, _P. Agenor_, L., Cr. Pap. Ex. t. 32. f. A, B.
♀, 2nd dimorphic form, _P. Achates_, Cr. Pap. Ex. t. 182. f. A, B; _P.
Alcanor_, Cr. Pap. Ex. t. 166, f. A.
_Hab._ Malacca (_Wall._), India.
_Remarks._—Ever since it was discovered that the insects figured by the
old authors as _P. Anceus_, _P. Agenor_, _P. Achates_, &c. were varying
females of _P. Memnon_ and _P. Androgeus_, the whole of these were very
naturally concluded to belong to one varying species. An examination of
many extensive collections, however, has convinced me that the
continental forms, on the one hand, and the insular ones, on the other,
can be readily distinguished, and really form two very well-marked
species. The red lunules at the anal region beneath characterize all
specimens from India (_Androgeus_, Cr.), while these are entirely absent
in all the insular specimens (_Memnon_, Cr.); and the same
characteristic difference can be traced in a greater or less degree
throughout all the infinitely varying female specimens. My specimen from
Malacca has a faint trace only on the upper surface of the
characteristic red mark at the base of the anterior wings; in other
respects it resembles the continental individuals. This form mimics the
Indian form of _P. Coon_ (_P. Doubledayi_, Wall.).
45. PAPILIO LAMPSACUS, Boisduval.
_P. Lampsacus_, Bd. Sp. Gén. Lép. p. 190; De Haan, Verh. Nat. Gesch.
p. 23, t. 2. f. 2.
_Hab._ Java (♂) (_Wall._).
46. PAPILIO PRIAPUS, Boisduval.
_P. Priapus_, Bd. Sp. Gén. Lép. p. 190; De Haan, Verh. Nat. Gesch. p.
23, t. 2. f. 1.
_Hab._ Java (_Boisd._), Sumatra (_Raffles_), Borneo (_De Haan_).
47. PAPILIO EMALTHION, Hübner.
♂, _Iliades Emalthion_, Hübn. Samml. Exot. ii. t. 117; _P. Emalthion_,
Boisd. Sp. Gén. Lép. p. 196; _P. Floridor_, Godt. Enc. Méthod. ix.
p. 809; _P. Kruscusterina_ in Eschsch. Voy. Kotzebue, t. 3. f. 5.
♀, 1st form, _P. Emalthion_, Cat. of Lep. Brit. Mus. pl. 5. f. 4.
♀, 2nd form, _P. Rumanzovia_, Eschsch. Voy. Kotz. t. 2. f. 4; _P.
Descombesi_, Boisd. Sp. Gén. p. 197; _P. Floridor_, ♀. Godt. Enc.
Méth. ii. p. 809.
_Hab._ Philippine Islands.
_Remarks._—I have no doubt whatever that we have here another case of
dimorphism, and I therefore unhesitatingly place these supposed species
under one name. The male of _P. Emalthion_ very closely resembles the
next species (_P. Deiphontes_), and the 2nd form of female (_P.
Rumanzovia_, Eschsch.) as closely resembles the female of the same
species; so that there can be no doubt that Godardt was right in
describing them as the sexes of his _P. Floridor_. The female figured in
the British Museum Catalogue is intermediate between these, but has more
of the characters of the male; and it is to be remarked that both these
forms of female have arrived in Europe accompanied by the same male. I
am therefore obliged to reduce by one the hitherto received species of
Philippine Papilios.
48. PAPILIO DEIPHONTES, n. s.
_P. Deiphobus_, var. A., Bd. Sp. Gén. Lép. p. 201.
♂. _Above_: exactly as in _P. Deiphobus_, but having a small tooth only
in place of the tail, and the posterior band of a clear ashy blue.
_Beneath_: with the markings as in _P. Emalthion_, except that the red
patch at the base of the upper wings is smaller.
♀. Also tailless, but resembling in markings the same sex of _P.
Deiphobus_, the pale patch on the upper wings not extending into the
discoidal cell.
Expanse of wings, ♂, 5½ inches; ♀, 5¾ inches.
_Hab._ Batchian, Gilolo, Ternate, Morty Isl. (_Wall._).
49. PAPILIO DEIPHOBUS, Linnæus.
_P. Deiphobus_, L., Cramer, Pap. Ex. t. 181. f. A, B; Donovan, Ins.
Ind. pl. 17. f. 2; Lucas, Lep. Ex. t. 11; Boisd. Sp. Gén. Lép. p.
200.
♀, _P. Alcandor_, Cr. Pap. Ex. t. 40. f. A, B.
_Hab._ Ceram, Amboyna, Bouru (_Wall._).
_Remark._—A simple variety of both this and the last species frequently
occurs, in which all the markings on the under side are ochre-yellow
instead of red.
50. PAPILIO ASCALAPHUS, Boisduval.
_P. Ascalaphus_, Bd. Sp. Gén. Lép. p. 200 (♂); De Haan, Verh. Nat.
Gesch. p. 26, t. 1. f. 2 (♀).
_Hab._ Menado, Macassar (Celebes), Sulla Isl. (_Wall._).
51. PAPILIO ŒNOMAUS, Godardt.
_P. Œnomaus_, Godt. Encyc. Méth. ix. p. 72; Boisd. Sp. Gén. Lép. p.
190; De Haan, Verh. Nat. Gesch. p. 24, t. 4. f. 1 (♂), 2 (♀).
_Hab._ Timor (♂, ♀) (_Wall._).
_Remark._—As has been already noticed (p. 22), the female of this
species closely resembles _P. Liris_ ♀, in company with which it was
captured.
g. _Helenus_ group.
52. PAPILIO SEVERUS, Cramer.
_P. Severus_, Cr. Pap. Ex. t. 227. f. A, B (♂), t. 278. f. A, B (♀);
Boisd. Sp. Gén. Lép. p. 212.
_Hab._ Bouru, Ceram, Amboyna, Gilolo, Batchian, Aru Isl. (_Wall._).
_Remarks._—This species exhibits a large amount of simple variation, in
the presence or absence of a pale patch on the uppers, in the brown
submarginal marks on the lower wings, in the form and extent of the
yellow band, and in the size of the specimens. The most extreme forms,
as well as the intermediate ones, are often found in one locality and in
company with each other, indicating that over the above range continual
intermixture probably takes place, and thus prevents any one form from
becoming specialized in a restricted area. The two following
modifications of it, however, have acquired perfect stability, each in a
large island situated on the extreme limits of the species. I therefore
consider them to be distinct, though the actual differences are but
small.
53. PAPILIO PERTINAX, n. s. Tab. V. fig. 4 (♂).
_Upper side_: anterior wings rather more elongate and pointed than in
_P. Severus_, dusky brown, with faint longitudinal rows of yellow scales
in the cell, and with rather denser scales between the nervures beyond
it; these are condensed into a narrow yellowish band parallel to the
outer margin, and rather nearer to the cell than to it. Hind wings
black, with three yellowish white subquadrate spots (the upper one
smallest) situate between the outer angle and the discoidal nervule;
beyond these and continued to the anal angle are a few very faint and
minute groups of scales.
_Under side_ as above, but the transverse band on the upper wings is
whiter, and on the lower wings are seven submarginal brownish-yellow
lunules, the middle ones least marked, and those at the outer and anal
angles having above them a very small group of minute blue scales.
The female is paler-coloured, with the markings rather more diffused,
and has on the under side an imperfect ocellus at the anal angle, a row
of faint brown lunules extending to the three white spots, and two
irregular lunules of blue atoms below those next the abdominal margin.
Expanse of wings, ♂, 4¼ inches; ♀, 5 inches.
_Hab._ Macassar (Celebes) (_Wall._).
_Remark._—This species was rather abundant near Macassar, in woody
places, and was very constant in its markings and general aspect.
54. PAPILIO ALBINUS, n. s. Tab. V. fig. 5 (♂).
Wings broader than in _P. Severus_, costa less arched, tail smaller, and
the caudal margin less produced.
_Upper side_: brown-black; anterior wings with very faint horizontal
lines of yellowish scales in the cell; apical portion of the wing more
thickly powdered between the nervures, the powdering fading away towards
the outer angle. Posterior wings with a large yellowish-white patch,
commencing close to the anterior margin, widening in the middle so as to
cross the end of the cell, and ending in a triangle with prolonged apex
at the abdominal margin; the outer edge of this spot is regularly
angulated and scalloped; two very faint brown lunules occur next the
anal angle; and the outer margin is rather broadly white-edged between
the dentations.
_Under side_: the anterior wings have distinct greyish lines of scales
between the nervures in the apical region; posterior wings not dotted
with scales as in _P. Severus_, but with two or three single rows of
scales in the cell only; the yellowish band consisting of a lunule next
the upper margin, followed by three rhomboidal spots notched below, of
which the middle one is the largest, then a roundish spot and a small
horizontal mark; a row of seven submarginal lunules, of which the three
middle ones are smallest and nearly obsolete, and that at the anal angle
much the largest and, with the whitish marginal spot below it, forming
an incomplete ocellus.
Expanse of wings 3½–3¾ inches.
_Hab._ New Guinea (♂) (_Wall._).
55. PAPILIO PHESTUS, Guérin.
_P. Phestus_, Guér. Voyage de la Coquille, t. 14. f. 2; Bd. Voy. de
l’Astrolabe, i. p. 41; Sp. Gén. Lép. p. 212.
_Hab._ New Guinea (_Paris Museum_).
56. PAPILIO HELENUS, Linnæus.
_P. Helenus_, L.; Cramer, Pap. Ex. t. 153. f. A, B; Lucas, Lep. Ex. t.
15. f. 2; Boisd. Sp. Gén. Lép. p. 211.
_Hab._ China (“_type_,” _Cramer’s figure_).
Local form _a_. Has more falcate wings and longer tail; the red marks at
the anal angle beneath are divided by a violet-white mark.
_Hab._ North India.
Local form _b_. Same form of wings as the last, but smaller; the third
and fourth lunules from the anal angle beneath very small or quite
absent.
_Hab._ Java, Sumatra (_Wall._).
57. PAPILIO HECUBA, n. s. Tab. V. fig. 3 (♂).
Upper wings falcate, and their outer margin much hollowed out, as in
many of the Celebes butterflies.
♂. _Upper side_: the outer half of the anterior wings of a fine cottony
texture, as in _P. Helenus_, but more marked; the red lunule at the anal
angle wanting; the rest as in _P. Helenus_.
_Under side_: the lunules and ocelli are ochre-yellow instead of deep
red, the two outer ones very small, the third almost obsolete, and the
next two absent; the anal ocellus is bordered with blue above, and
adjoining it is a blue lunule in the place of the red one in _P.
Helenus_.
♀. _Upper side_: of a browner colour; two orange-brown ocelli at the
anal angle.
_Under side_: the lunules and ocelli all larger; the two intermediate
ones entirely absent, as in the male.
Expanse of wings 5½–5¾ inches.
_Hab._ Macassar, Menado (Celebes) (_Wall._).
58. PAPILIO ISWARA, White.
_P. Iswara_, White, Entom. 1842, p. 280; Doub. and Hew. Gen. of Diurn.
Lep. pl. 2. f. 1 (♀).
_Hab._ Penang, Malacca, Singapore, Borneo (♂, ♀) (_Wall._).
59. PAPILIO HYSTASPES, Felder.
_P. Hystaspes_, Feld. Lep. Nov. Philipp. p. 12.
_Hab._ Luzon (Philippines).
This is the Philippine form of _P. Helenus_.
60. PAPILIO ARASPES, Felder.
_P. Araspes_, Feld. Ent. Fragm. p. 17.
_Hab._ Philippine Islands.
This comes near to _P. Iswara_.
61. PAPILIO NEPHELUS, Boisduval.
_P. Nephelus_, Bd. Sp. Gén. Lép. p. 210; De Haan, Verh. Nat. Gesch. p.
29, t. 4. f. 4, ♂.
_Hab._ Malacca, Sumatra, Borneo (♂, ♀) (_Wall._), Assam (_Brit.
Mus._).
h. _Pammon_ group.
62. PAPILIO PAMMON, Linnæus. Tab. II. figs. 1 (♂), 3, 5, 6 (♀ ♀).
♂, _P. Pammon_, L.; Cram. Pap. Ex. t. 141. f. B; Boisd. Sp. Gén. Lép.
p. 272.
♀, _P. Polytes_, L.; Cram. Pap. Ex. t. 265. f. A, B, C.
_Hab._ Malacca, Singapore (_Wall._), China, India, Ceylon.
The continental specimens of _P. Pammon_ have all considerably developed
tails in both sexes; the insular specimens on the other hand, (which I
treat as a separate species), have only a prominent tooth or very short
tail in the males. The females also differ considerably, presenting an
analogous but distinct series of forms. In the true _P. Pammon_ the
males are very constant; but the females exist under three distinct
forms, each of them presenting more or less numerous varieties, viz.:—
_1st form_ of female. Tab. II. fig. 3.
This exactly resembles the male, except in the possession of a distinct
ocellus at the anal angle on the upper surface. Rarely a variety occurs
having in addition a submarginal row of red lunules, indicating a slight
approximation towards some varieties of the second form.
_2nd form_ of female (_P. Polytes_). Tab. II. fig. 5.
This is by far the most common form of female. A variety of this rarely
occurs, which wants the red patch at the anal angle, and has the white
patch formed of a row of spots all situated a little below the discoidal
cell. This is the nearest approach to the first form.
_3rd form_ of female (_P. Romulus_, Cram. Pap. Ex. t. 43. f. A; _P.
Mutius_, Fab., Boisd. Sp. Gén. p. 270; _P. Hector_ ♀, De Haan). Tab. II.
fig. 6.
This not uncommon Indian butterfly I consider to be a third form of the
female of _P. Pammon_. I was first led to suspect this by finding that
no males of it are known (the male and female from Ceylon, noted in the
British Museum List, I have ascertained to be both females), nor have I
been able to find any after an examination of the chief collections in
England. It is also to be observed that it has been received from no
locality which is not also inhabited by _P. Pammon_; there is no other
known Indian butterfly that can possibly be the other sex of it; and
lastly, it agrees very closely with the second form of female (_P.
Polytes_) in all its details of form, texture, and neuration; and though
at first sight having a very different aspect, specimens are to be found
which by a very slight modification could be changed so as to resemble
that form. I am therefore quite satisfied in my own mind that I am right
in sinking this species into a form of _P. Pammon_. I have already
stated my opinion that it mimics _P. Hector_, with which, however, it
has no affinity. The resemblance was such as to induce De Haan to place
it as the female of that species.
63. PAPILIO THESEUS, Cramer. Tab. II. figs. 2, 4, 7 (♀ ♀).
_P. Theseus_, Cr. Pap. Ex. t. 180. f. B (♀); Boisd. Sp. Gén. Lép. p.
276.
_P. Antiphus_ ♀, De Haan, Verh. Nat. Gesch. p. 49, t. 8. f. 2; Brit.
Mus. List. Pap. p. 12.
_P. Polyphontes_ ♀, De Haan, Verh. Nat. Gesch. t. 8. f. 4.
_P. Melanides_, De Haan, Verh. Nat. Gesch. t. 8. f. 3 (♀).
Male like _P. Pammon_ ♂, but smaller, and the tail always reduced to a
projecting tooth.
_Hab._ Java, Sumatra, Borneo, Lombock, Timor (_Wall._).
Local form _a_. Much larger; more falcate wings; a broad short tail.
_Hab._ Macassar (_Wall._)
_1st form_ of female. Tab. II. fig. 2.
Like the male, but with a very slightly marked blue and red ocellus at
the anal angle. This is very rare in the islands. I found one specimen
only in Timor, which I took “_in copulâ_” with a male almost exactly
resembling it.
_2nd form_ of female (_P. Polyphontes_ ♀, De Haan). Tab. II. fig. 4.
Like the 2nd form of _P. Pammon_ ♀; but has the pale portion of the
anterior wing of a much lighter colour, and not extending so far towards
the base of the wing; the white spot on the hind wings is more rounded,
and has always a rather large portion within the cell. This form is to
some extent local, not existing, I believe, in Sumatra, where it is
replaced by the next.
_Hab._ Borneo, Java, Timor (_Wall._).
_3rd form_ of female (_P. Theseus_, Cr.; _P. Antiphus_ ♀, De Haan). Tab.
II. fig. 7.
This is well characterized by the entire absence of the white spot from
the hind wings. The red spots and lunules remain; but in some specimens
only those in the anal region are visible, and these have a very close
resemblance to _P. Antiphus_. This is also a local form, not occurring,
I believe, in company with the last.
_Hab._ Sumatra, Lombock (_Wall._).
_4th form_ of female (_P. Melanides_, De Haan, Verh. Nat. Gesch. t. 8.
f. 3).
I consider this to be an isolated modification of _P. Theseus_, Cr.,
peculiar to Borneo. It possesses all the characteristics of a female of
this species.
_Hab._ Banjarmassing (Borneo) (_Leyden Museum_).
N.B. The 2nd, 3rd, and 4th forms of ♀ are all tailed, as in the females
of _P. Pammon_.
64. PAPILIO ALPHENOR, Cramer.
_P. Alphenor_, Cr. Pap. Ex. t. 90. f. B (♀); Boisd. Sp. Gén. Lép. p.
274 (♂, ♀); _P. Ledebouria_, Eschsch. Voy. Kotz. t. 3. f. 7.
This is very closely allied to _P. Theseus_. The male is larger, has the
caudal tooth scarcely perceptible, and on the under side has white
instead of red marginal lunules. The female is tailed, much larger than
_P. Theseus_ ♀ form 2nd, from which it further differs by the white
patch on the hind wings having the red markings blended with it, and
more prominent.
_Hab._ Celebes, Bouru, Amboyna, Ceram (_Wall._), Philippine Islands.
_1st form_ of female (_P. Ledebouria_, Eschsch.).
Like the male, but with a brown tinge and an obscure anal lunule. This
has been noticed only in the Philippine Islands.
_2nd form_ of female (_P. Alphenor_, Cr.).
Distribution the same as the male.
_3rd form_ of female (_P. Elyros_, G. R. Gray, B. M. List Pap. p. 26).
The white patch on the lower wings reduced to a small spot, or quite
absent. There are many varieties of this, showing very instructively how
such isolated forms of female as occur in the two preceding species may
have been produced by simple variation followed by a “natural selection”
of the forms best adapted to special conditions.
_Hab._ Philippine Islands (_B. M._)
65. PAPILIO NICANOR, Felder, ‘Voyage of the Novara,’ pl. ... f. _c_,
_d_.
_Male._ Upper side:—like _P. Alphenor_ ♂; but the band of white spots is
broader and more regular, and there is a row of four white submarginal
lunules.
Under side as in _P. Alphenor_; but the marginal spots of the upper
wings, and the submarginal lunules of the lower wings, are larger and
more distinct.
_Female_ quite tailless, like the male. Upper side:—like _P. Alphenor_
♀; but the rufous anal spots are much smaller, not forming an ocellus at
the anal angle, and they do not join the white central patch.
Under side, differs from _P. Alphenor_ in nearly the same manner as on
the upper side.
_Hab._ Batchian, Gilolo, Morty Island (_Wall._).
_Remarks._—The absence of tails in the female, and the white submarginal
lunules in the male, distinguish this at a glance from all its allies.
It has a comparatively restricted range, and is very constant in both
sexes. The plate sent me by Dr. Felder is not numbered.
66. PAPILIO HIPPONOUS, Felder[14].
Footnote 14:
Having obtained a specimen of this insect while these sheets are
passing through the press, I find that it should have been placed next
to _P. Severus_.
_P. Hipponous_, Feld. Lep. Nov. Philipp. p. 12; _P. Dironus_, B. M.
List (no description).
_Hab._ Luzon, Mindanao (Philippines).
67. PAPILIO AMBRAX, Boisduval.
_P. Ambrax_, Bd. Sp. Gén. Lép. p. 218; Voy. au Pôle Sud, Lép. t. 1. f.
3, 4 (♂); De Haan, Verh. Nat. Gesch. t. 7. f. 2 (♀). _P. Orophanes_,
Boisd. Sp. Gén. p. 275 (♀).
_Hab._ Mysol, Salwatty, Dorey (_Wall._).
_Remark._—I believe that two, if not three, well-marked forms or species
have been mixed up under the name of _P. Ambrax_, as I have endeavoured
to show by the references. My specimens of the two sexes of each show a
uniformity of character in each locality.
68. PAPILIO AMBRACIA, Wallace.
_P. Ambrax_, Bd.; De Haan, Verh. Nat. Gesch. t. 7. f. 1 (♂).
_Male._ Differs from _P. Ambrax_, Bd., by the ashy-white patch at the
apex of the anterior wings.
_Female._ Has a large, roundish, white patch on the anterior wings,
extending from the discoidal cell to the hinder angle. The red lunules
on the hind wings are smaller. Same size as _P. Ambrax_.
_Hab._ Waigiou (♂, ♀) (_Wall._).
69. PAPILIO EPIRUS, n. s.
_Male._ Above:—anterior wings as in _P. Ambrax_; posterior wings more
elongate, the white band much narrower, notched behind at the nervures,
with the portions between regularly rounded; the part which crosses the
cell is cut by black nervures, and there is an oblique red mark at the
anal angle.
Beneath:—with a submarginal of seven lunules on the hinder wings, the
one above the anal angle very large; whereas the last two species have
one small lunule only beneath, at the anal angle.
_Female._ Is probably that figured in ‘Voy. au Pôle Sud,’ Lép. t. 1, f.
5, which resembles most the female of _P. Ambracia_, but differs in the
form of the white and red patches. It is said to be from “the coasts of
New Guinea”; but as the expedition touched at the Aru Islands, it is
very probable that there is an error of locality, as I have ascertained
to be very often the case in the indications furnished by these and
other ‘Voyages.’
_Hab._ Aru Islands (_Wall._).
70. PAPILIO DUNALI, Montrouzier.
_P. Dunali_, Mont. Ann. Soc. d’Agricult. de Lyon, 1856, p. 394.
_Hab._ Woodlark Island (S.E. of New Guinea).
_Remark._—This seems closely allied to the last species.
i. _Erectheus_ group.
71. PAPILIO ORMENUS, Guérin. Tab. III. figs. 2 (♂), 1, 3, 4 (♀ ♀).
_P. Ormenus_, Guér. Voy. de la Coquille, pl. 14. f. 3; Boisd. Sp. Gén.
Lép. p. 211.
_P. Erectheus_, var., Voy. au Pôle Sud, Lép. t. 1. f. 1, 2.
_P. Amanga_, Boisd. Sp. Gén. p. 216, ♀ (_P. Onesimus_, Hew. Ex. Butt.
Pap. iii. f. 8).
_Hab._ Waigiou, Aru Isl., Ké Isl., Matabello and Goram Isl. (_Wall._).
This belongs to a remarkable group of Papilios inhabiting the
Austro-Malayan region, and which are especially interesting as
exhibiting a good instance of polymorphism, the females being of two or
three distinct forms.
The male in this species is characterized by the small amount of marking
on the under surface.
_1st form_ of female. Tab. III. fig. 1.
Almost exactly intermediate between the male and the normal female,
which resembles _P. Erectheus_ ♀.
Upper side brown-black; a band of four whitish-yellow spots across the
anterior wings beyond the cell, the upper one of the same size and
position as in the male, the 2nd and 3rd elongated towards the cell, the
4th rather shorter than the 3rd, and immediately beneath it. Posterior
wings with a central patch of a pale sulphur-yellow just crossing the
end of the cell, and separated below into five truncate lobes; below
this, and next the anal margin, are two irregular blue lunules, with a
red lunule at the anal angle and a smaller one lower down beneath the
second blue lunule.
Under side as above; on the hind wings the upper half of the yellow
patch is dusky, and there is a complete submarginal series of seven red
lunules.
_Hab._ Waigiou (a single specimen) (_Wall._).
_2nd form_ of female. Tab. III. fig. 3.
Resembles very closely _P. Erectheus_ ♀; but the white patch on the hind
wings does not cover so much of the cell, and the two middle lobes are
much elongated posteriorly, and separated by wedge-shaped spaces; the
blue lunules are but slightly marked, and do not exceed two in number.
Under side:—differs from _P. Erectheus_ in the white patch never
reaching the anterior margin of the hind wings. In a specimen from
Waigiou, the four middle lunules are nearly white. This may be
considered the typical form of female, as it occurs everywhere in
company with the male.
_3rd form_ of female (_Amanga_, Bd.). Tab. III. fig. 4.
I have three specimens of this form from three of the localities in
which the male occurs. They differ slightly from each other, but agree
generally with the figure and description above quoted. An allied form
of female (of the next species) was observed closely followed by two
males of the ordinary form; they were watched for some time, the males
hovering over the females in the manner usual before pairing; and the
three were then captured at one stroke of the net. This occurred three
years after the capture of the specimen figured by Mr. Hewitson, and at
once convinced me that these puzzling specimens were an additional form
of female to a well-known male. The fact that the only females known of
an allied species (_P. Tydeus_) are intermediate between these forms
confirms this determination.
_Hab._ Aru Island, Mysol, Goram Isl. (_Wall._)
72. PAPILIO PANDION, n. s.
_Male._ Closely resembles _P. Ormenus_, but presents the following
differences:—
Upper side:—the band of spots across the fore wings is faintly marked,
or more frequently quite absent; the grey lines bordering the nervures
at the apex are more distinct; on the hind wings, the first three
indentations of the whitish patch are followed by faint powdered lunules
of the same colour.
Under side:—the apex of the fore wings is strongly marked with grey
lines between the nervures, but has generally no spots; on the hind
wings there is a curved submarginal band of lunules across the wing,
viz., at the anal angle a large irregular red lunulate spot with a blue
and a grey mark above it—2nd, a larger grey lunule with an angular blue
mark below it, and a red lunule nearer the margin—3rd, a similar grey
lunule and blue mark—4th, a larger grey lunule, and a smaller blue mark
with a faint red lunule below—5th, a grey lunule and a faint blue dash
below—6th, a blue lunule with a faint grey mark above—7th, a blue lunule
with a very faint mark above it. These vary somewhat in different
specimens, but the whole series can always be traced.
_1st form_ of female.
Scarcely distinguishable from the typical female of the last species:
the blue lunules on the under surface form a complete series, almost as
in _P. Erectheus_ ♀.
_Hab._ New Guinea, Salwatty, Mysol Island (with the male) (_Wall._).
_2nd form_ of female.
Upper surface:—fore wings as in _P. Onesimus_, Hew.; hind wings
yellowish-white, a broad black border along the anterior, and a narrow
one along the posterior margin, two yellowish lunules near the outer
angle, anal angle pale yellow, then an oblong black spot with a bluish
mark in its upper part, followed by a second (half-obliterated) black
spot.
Under surface with the same markings; but there are a series of six blue
angulated marks upon a black ground, the two intermediate ones being
smaller and less distinct. Abdomen yellow; under side black.
_Hab._ Dorey (New Guinea) (_Wall._)
_Remarks._—This specimen was taken in company with two males, as before
mentioned. An insect, described by M. Montrouzier as the female of his
_P. Godartii_ (from Woodlark Island), agrees very closely with this, and
is no doubt the female of the same species, or a closely allied one
which he puts in his list as _P. Ormenus_. The fact, therefore, that
this peculiar pale form of female _Papilio_ has been found in five
islands, from no one of which is a male insect known which can be mated
with it, except those of the _Ormenus_-form (which always occur in the
same places), may, in conjunction with the observation already given of
the companionship of the two forms, be taken to prove that this is
really a case of polymorphism. I believe also it will be found that
these extreme departures from the typical form of a species are
connected with mimetic resemblances and the safety of the individuals.
We have already seen that the extreme forms of _P. Memnon_ ♀ and _P.
Pammon_ ♀ respectively resemble other species which from their habits
and abundance seem to have some peculiar immunity from danger. In this
case also there is a resemblance to quite a different family of
butterflies, the Morphidæ. In form, coloration, and general appearance
these pale-coloured Papilios resemble species of the genus _Drusilla_;
and the same genus is also imitated by other butterflies—one of these,
_Melanitis Agondas_ ♀, having been actually confounded with _Drusilla
bioculata_ as the same species, so great is the resemblance. This fact
of species of several genera imitating the Drusillas would indicate that
they have some special immunities which make it advantageous to other
insects to be mistaken for them; and their habits confirm this opinion.
They have all a very similar style of dress, and fly very slowly, low
down in damp woods, often settling on the ground or on rotten wood; and
they are exceedingly abundant in individuals. Now these are the general
characteristics of all groups which are the subjects of imitation; and
we may therefore presume, when we see forms departing widely from the
general appearance of their close relations, and resembling closely
other groups with which they have no affinity, that what we must call
_accidental_ variations have been accumulated and rendered definite by
natural selection for the protection and benefit of those forms.
73. PAPILIO TYDEUS, Felder. Tab. IV. figs. 3 (♂), 2 (♀).
_P. Tydeus_, Feld. Lep. Fragm. p. 52 (♂).
_Female._—Upper side dusky brown; fore wings with the central portion
below the cell nearly white; hind wings with the basal two-thirds white,
with an irregular and obtusely dentated margin, and edged with
ochre-yellow; the rest black, with a submarginal row of seven broad
yellowish lunules, and above those nearest the anal angle three
irregular blue patches.
Under side nearly as above; the white space on the upper wings is more
extensive and better defined; the marginal lunules are dilated so as to
form a crenellated band, and the blue marks are increased to six or
seven in number. Head and thorax dusky; abdomen yellowish.
_Hab._ Batchian, Morty Island (_Wall._).
_Remark._—The female, which seems to be of only one form in this
species, is especially interesting as being allied to the pale-yellow
form of _P. Ormenus_ and _P. Pandion_.
74. PAPILIO ADRASTUS, n. s. Tab. IV. fig. 1 (♀).
_Male._—Upper side, like _P. Ormenus_ ♂; but has the band of the hind
wings narrower, not crossing the cell, and more pointed towards the anal
angle.
Under side with a single red anal spot, and three blue lunules beyond
it.
_Female._—Upper side brown-black; anterior wings with the apical half
browner, a whitish patch around the end of the cell, and an ovate spot
within it; posterior wings with a small central whitish patch more or
less tinged with ochreous; a submarginal row of very large deep-red
lunules, that at the anal angle forming an irregular ocellus bordered
above with pale blue, and a few blue atoms on the side of it.
Indentations of all the wings broadly margined with ochreous.
Under side:—the white patch of the anterior wings larger and well
defined, and continued by smaller and fainter patches to the outer
angle; posterior wings with the small central patch and marginal lunules
as above, with the addition of a faint row of angulated blue marks
between them.
Wings elongated posteriorly, and somewhat angulated at the termination
of the first median nervure.
Expanse of wings, ♂, 5¼ inches; ♀, 6 inches.
_Hab._ Banda Island (_Wall._).
_Remarks._—This species is near _P. Ormenus_ in the male, but approaches
_P. Gambrisius_ in the female, which differs from all others in this
group by its dark colouring and the short narrow band on the hind wings.
A male and two females were obtained in the small island of Banda.
75. PAPILIO GAMBRISIUS, Cramer.
_P. Gambrisius_, Cr. Pap. Ex. t. 157. f. A, B (♂); Boisd. Sp. Gén.
Lép. p. 213.
_P. Drusius_, Cr. Pap. Ex. t. 229. f. A, t. 230. f. A (♀); Boisd. Sp.
Gén. Lép. p. 218.
_Hab._ Amboyna, Ceram, Bouru (_Wall._).
_Remarks._—The males of this fine species are not uncommon in Ceram, and
in hot weather come down to the beach and settle on the wet sand. The
females, however, are very rare; I obtained one in the mountainous
forests of Ceram, and this is, I believe, the only fine and perfect
specimen now in Europe.
Expanse of male 5½–6½ inches, of female 7 inches.
76. PAPILIO AMPHITRION, Cramer.
_P. Amphitrion_, Cr. Pap. Ex. t. 7 f. A, B; Boisd. Sp. Gén. Lép. p.
217.
_Hab._ Celebes?
_Remarks._—The habitat of this rare species is doubtful. Cramer says,
“America;” Godart, “Amboyna;” but I believe its true locality will be
found to be Celebes. It forms a transition to the next species.
77. PAPILIO EUCHENOR, Guérin.
_P. Euchenor_, Guér. Voy. de la Coquille, t. 13. f. 3 (♂); _P. Axion_,
Boisd. Sp. Gén. Lép. p. 46 (♂).
_Female._—Similar to the male; but the markings are all of a dull
ochre-yellow, and the second and third spots, reckoning from the inner
margin of the upper wings, are almost entirely wanting. This sex is much
rarer than the male.
_Hab._ New Guinea, Aru Island, Ké Island (_Wall._).
78. PAPILIO GODARTII, Montrouzier.
_P. Godartii_, Montr. Ann. Soc. d’Agric. de Lyon, 1856, p. 394.
_Hab._ Woodlark Island.
_Remark._—Closely allied to the last; perhaps a variation only.
k. _Demolion_ group.
79. PAPILIO DEMOLION, Cramer.
_P. Demolion_, Cr. Pap. Ex. t. 89. f. A, B; _P. Cresphontes_, Fabr.;
Boisd. Sp. Gén. Lép. p. 220.
_Hab._ Java, Borneo, Sumatra, Singapore (_Wall._), Moulmein (_Brit.
Mus._).
80. PAPILIO GIGON, n. s. Tab. VII. fig. 6 (♀).
“_P. Gigon_,” List of Papilionidæ in Brit. Mus. p. 27 (no
description).
Much larger than _P. Demolion_; costal margin of the fore wings very
much arched from the base; tail proportionally shorter.
Upper side:—markings as in _P. Demolion_, with the following
differences. In the cell of the fore wings are four longitudinal curved
greyish-yellow lines; the yellow band begins higher on the abdominal
margin, and curves outward toward the tip, where the spots are obliquely
elongate, and the three last distinctly notched; on the hind wings the
lunulate spots are much deeper and are rather further from the margin,
and the two spots at the outer angle (often obsolete in _P. Demolion_)
are large and well marked.
Under side:—the markings resemble those of _P. Demolion_, but are
stronger; the band of silvery spots is much more sinuate, and possesses
an additional lunule above the outer angle; a patch of ochre-yellow
covers the lower margin of the cell, extending a little along the
nervures which radiate from it.
Abdomen blackish, with numerous stripes and spots of pale yellow.
Expanse of wings 4¾ to 5⅓ inches.
_Hab._ Celebes, Sulla Island (_Wall._).
_Remark._—This was regarded by Boisduval as a large variety of _P.
Demolion_ (see Sp. Gén. Lép. p. 221); but it offers remarkable
differences both in form and markings.
l. _Erithonius_ group.
81. PAPILIO ERITHONIUS, Cramer.
_P. Erithonius_, Cr. Pap. Ex. t. 232. f. A, B.
_P. Epius_, Fabr.; Don. Ins. China, pl. 29. f. 2; Boisd. Sp. Gén. Lép.
p. 238.
_Hab._ India, China (_type_).
Local form _a_ (_Malayanus_).—The two spots on the lower margin of the
cell of the hind wings wanting; anal spots redder, and the ocellus at
the outer angle darker: two spots in cell of fore wings, as in the type;
but in Flores specimens these approach so as almost to unite.
_Hab._ Singapore, Flores (_Wall._), Manilla.
Local form _b_ (_Sthenelus_, Macleay).—A single large spot in the cell
of the fore wings; one small detached spot on the margin of the cell of
the hind wings.
_Hab._ Goram Island (_Wall._), Australia.
SECTION C.
m. _Paradoxa_ group.
82. PAPILIO PARADOXA, Zinken.
_Zelima Paradoxa_, Zink. Beitr. Ins. Java, t. 15. f. 9, 10.
_P. Paradoxa_, Westw. Cab. Or. Ent. pl. 9. f. 1, 1*.
_Hab._ Java (_Wall._).
Local form _a_.—_P. Paradoxa_, var., Hew. Proc. Zool. Soc. 1859, p. 422,
pl. 67. f. 1 (♂), 2 (♀).
_Hab._ Borneo (_Wall._).
Local form _b_.—Smaller; intermediate in the markings between the Java
and Borneo forms; interior row of elongate marks on upper wings light
blue, not descending to the outer angle.
_Hab._ Sumatra (_Wall._).
Both sexes of this species closely resemble the corresponding sexes of
_Euplœa Midamus_, Cr., which is very common in all the above-mentioned
localities.
83. PAPILIO ÆNIGMA, n. s. Tab. VII. fig. 3 (♂).
Size, form, and markings nearly the same as in _P. Paradoxa_.
Above:—purplish black, without any gloss or silky reflexions; a
submarginal row of white spots on all the wings, more or less blue-edged
on the upper wings, sometimes partially obsolete on the lower ones; one
or two spots at the end of the cell, and a row of six or seven elongate
marks beyond it, bright blue.
Beneath, the submarginal row of white spots only.
_Female._—_P. Paradoxa_, var. A, Hewitson, Proc. Zool. Soc. 1859, p.
423, pl. 67. f. 3.
I put this as the female of the above with some hesitation, as it was
not captured in the same island. It agrees, however, in the entire
absence of gloss, and in the peculiar elongation of the outer angle of
the lower wings.
_Hab._ Malacca, Sumatra (♂); Borneo (♀) (_Wall._).
_Female variety?_—_P. Paradoxa_, var. B, Hewitson (Proc. Zool. Soc. pl.
66. f. 4), may be an extreme variation of this, but will more probably,
when the male is discovered, prove to be a distinct species.
84. PAPILIO CAUNUS, Westwood.
_P. Caunus_, Westw. Cab. Or. Ent. pl. 9. f. 2, 2*.
_Hab._ Sumatra, Borneo (♂, ♀) (_Wall._), Java (_Leyden Mus._).
_Remarks._—My specimens have less white on the lower wings than is
represented in Mr. Westwood’s figure. The female is of a brownish
colour, with the same white markings as the male, but without any blue
tinge. This species is very like _Euplœa Rhadamanthus_, one of the most
common butterflies in all the above-mentioned localities. It is
undistinguishable from that insect on the wing, though it flies very
slowly, like the species it mimics.
85. PAPILIO ASTINA, Westwood.
_P. Astina_, Westw. Cab. Or. Ent. pl. 9. f. 3.
_Hab._ Java (_Brit. Mus. ex Coll. Horsf._).
86. PAPILIO HEWITSONII, Westwood.
_P. Hewitsonii_, Westw. Proc. Ent. Soc. 1864, p. 10.
_P. Slateri_ ♀, Hew. Ex. Butt. Pap. pl. 4. f. 9; _P. Cammu_, B. M.
List of Papilionidæ (no description).
_Hab._ Borneo (♂) (_Wall._).
_Remarks._—The last two species should probably form a distinct group,
on account of the peculiar elongation of the cell of the lower wings.
They both resemble dark species of _Euplœa_. _P. Slateri_ is a quite
distinct species from North India, to which Mr. Hewitson referred the
present species as the female. All the specimens known of both species
are, however, males.
n. _Dissimilis_ group.
87. PAPILIO ECHIDNA, De Haan.
_P. Echidna_, De Haan, Verh. Nat. Gesch. p. 42, t. 8. f. 6; _Clytia
dissimilis_, Sw. Zool. Ill. 2nd ser. pl. 120; _P. dissimilis_, var.,
Brit. Mus. List of Papilionidæ.
_Hab._ Timor (♂, ♀) (_Wall._).
_Remarks._—This species has been confounded with _P. dissimilis_, from
which it is very distinct, by the absence of the yellow marginal band
beneath. It is also widely separated geographically from that species,
which inhabits the continent of India only. The sexes are alike, as they
are in _P. dissimilis_. _P. Panope_, L., which has been supposed to be
its female, is a very distinct species, of which also both sexes exist
in most collections.
88. PAPILIO PALEPHATES, Westwood.
_P. Palephates_, Westw. Arc. Ent. pl. 79. f. 1; _P. dissimilis_, var.
_b_, Brit. Mus. List of Papilionidæ.
_Hab._ Philippine Islands.
SECTION D.
o. _Macareus_ group.
89. PAPILIO VEIOVIS, Hewitson.
_P. Veiovis_, Hew. Ex. Butt. Pap. pl. 7. f. 20 (♂).
_Hab._ Menado (Celebes) (“_Coll. Hewitson._”).
_Remark._—This fine new species has been recently received from Menado,
and seems best placed in this group, near _P. Encelades_.
90. PAPILIO ENCELADES, Boisduval.
_P. Encelades_, Bd. Sp. Gén. Lép. p. 376; Hewitson, Ex. Butt. Pap. pl.
4. f. 10 (♂).
_Hab._ Macassar (Celebes) (_Wall._).
91. PAPILIO DEUCALION, Boisduval.
_P. Deucalion_, Bd. Sp. Gén. Lép. p. 375; Hewitson, Ex. Butt. Pap. pl.
4. f. 11 (♀).
_Hab._ Macassar, Menado (Celebes) (_Wall._).
_Remarks._—At Macassar I took only males of _P. Encelades_, and females
of _P. Deucalion_ at the same spot (a half-dry river-bed), and therefore
conjectured that they might be sexes of one species, although so unlike.
Some years afterwards, however, I took at Menado a fine male of _P.
Deucalion_, which only differs in its rather smaller size and brighter
colouring.
92. PAPILIO IDÆOIDES, Hewitson.
_P. Idæoides_, Hew. Ex. Butt. Pap. pl. 1. f. 2.
_Hab._ Philippine Islands (♂) (_Brit. Mus._).
_Remark._—This singular species must closely resemble on the wing
_Hestia Leuconoë_, from the same islands.
93. PAPILIO DELESSERTII, Guérin.
_P. Delessertii_, Guér.; Deless. Souvenirs, t. 17.
_Hab._ Pulo Penang (_Hope Museum, Oxford_).
_Remark._—This resembles the species of _Hestia_ and _Idæopsis_, from
the same locality, and is intermediate in size. It has been confounded
with the next.
94. PAPILIO DEHAANII, Wallace.
_P. Laodocus_, De Haan, Verh. Nat. Gesch. t. 8. f. 5 (nec Fab.); _P.
Melanides_, Erichs. Archiv für Natur. 1843 (nec De Haan, 1839).
_Hab._ Malacca, Borneo (_Wall._), Java (_Leyden Mus._).
_Remarks._—The Bornean specimens are rather larger, and have the yellow
anal spot somewhat differently shaped. The two names which have been
applied to this species having been preoccupied, I have named it after
the first describer.
95. PAPILIO LEUCOTHOË, Westwood.
_P. Leucothoë_, Westw. Arc. Ent. pl. 79. f. 3; _P. Xenocles_, var.,
Brit. Mus. List of Pap.
_Hab._ Singapore, Malacca (_Wall._), N. India.
96. PAPILIO MACAREUS, Godart.
_P. Macareus_, Godt. Enc. Méth. ix. pl. 76; Horsf. Desc. Cat. Lep. E.
1. C. pl. 5. f. 1; Boisd. Sp. Gén. Lép. p. 374.
_P. striatus_, Zink. Beitr. Ins. Java, t. 14. f. 5.
_Hab._ Malacca (_Wall._), Java (_Horsfield_), Borneo (_Leyden Mus._).
This species closely resembles _Danais Aglaë_, Cr., found in the same
islands.
97. PAPILIO STRATOCLES, Felder.
_P. Stratocles_, Feld. Lép. Nov. Philipp, p. 2.
_Hab._ Mindanao (Philippines).
98. PAPILIO THULE, n. s. Tab. VII. fig. 1 (♂).
Form of _P. Macareus_, but smaller.
Above:—brown-black, spotted and marked with greenish white; a row of
spots near the outer margin of all the wings, and on the upper wings a
second row between the first and the end of the cell, three or four
others close to the cell, and 5–7 irregularly placed in the cell; the
spot next the outer angle is double, and the two lower spots of the
second row are continued indistinctly to the cell. The lower wings have
a mark at the end of the cell, and five elongated spots radiating from
it between the nervures.
Beneath:—brown, with the spots all whiter and more distinct. Neck with
four white points; abdomen dusky, with pale lines on the sides and
beneath.
Expanse of wings 3¾ inches.
_Hab._ New Guinea (♂) (_Wall._).
Variety or local form _a_.—Like the above, but with the discal spots of
the lower wings united into a transverse band divided by fine nervures.
_Hab._ Waigiou Island (♂) (_Wall._).
This species imitates _Danais sobrina_, Bd., a New Guinea species. The
figure represents the upper surface of both forms of this insect.
p. _Antiphates_ group.
99. PAPILIO ANTIPHATES, Cramer.
_P. Antiphates_, Cr. Pap. Ex. t. 72, f. A, B; Boisd. Sp. Gén. Lép. p.
248.
_P. Pompilius_, Fab.; Lucas, Lep. Ex. t. 22. f. 1; Godt. Enc. Méthod,
ix. p. 49.
_P. Alcibiades_, Fab.; Godt. Enc. Méthod, ix. p. 49.
_Hab._ India, China (“_type_”).
Local form _a_.—_Podalirius Pompilius_, Sw. Zool. Ill. 2nd ser. pl. 105.
_Hab._ Malacca, Sumatra, Java, Borneo (_Wall._).
These differ from the type in the black apical portion not quite
reaching the outer angle, and in the first and second bands on the upper
wings not extending below the cell. The fourth band varies in extent, as
does the amount of grey colouring in the caudal region.
100. PAPILIO EUPHRATES, Felder.
_P. Euphrates_, Feld. Lep. Nov. Philipp. p. 12; _P. Coretes_, Brit.
Mus. List of Papilionidæ (no description).
_Hab._ Philippine Islands.
101. PAPILIO ANDROCLES, Boisduval. Tab. VII. fig. 5 (♂).
_P. Androcles_, Bd. Sp. Gén. Lép. p. 279.
_Hab._ Macassar (Celebes) (_Wall._).
_Remarks._—I only met with this magnificent species on one occasion, on
the banks of a mountain-stream and on the sands close to a waterfall.
When resting on the ground, the very long white tails are raised up at a
considerable angle, and are very conspicuous.
102. PAPILIO DORCUS, De Haan.
_P. Dorcus_, De Haan, Verh. Nat. Gesch. Zool. t. 7. f. 4.
_Hab._ Gorontalo (N. Celebes) (“_Leyden Museum_”).
103. PAPILIO RHESUS, Boisduval.
_P. Rhesus_, Bd. Sp. Gén. Lép. p. 253.
_Hab._ Macassar (Celebes) (_Wall._). “Bengal,” the locality given by
Boisduval, is erroneous.
104. PAPILIO ARISTÆUS, Cramer.
_P. Aristæus_, Cr. Pap. Ex. t. 318, f. E, F; Boisd. Sp. Gén. Lép. p.
252.
_Hab._ Ceram, Batchian (_Wall._).
105. PAPILIO PARMATUS, G. R. Gray.
_P. Parmatus_, G. R. Gray, Cat. Lep. Ins. Brit. Mus. pl. 3. f. 2.
_Hab._ Aru Islands, Waigiou (_Wall._), Australia (_Brit. Mus._).
_Remarks._—The Aru specimen agrees almost exactly with the type specimen
in the British Museum. The Waigiou insect is rather darker on the under
surface, and has the black markings more sharply defined.
q. _Eurypylus_ group.
106. PAPILIO CODRUS, Cramer.
_P. Codrus_, Cr. Pap. Ex. t. 179. f. A, B; Boisd. Sp. Gén. Lép. p.
228.
_Hab._ Amboyna and Ceram (_type_) (♂, ♀) (_Wall._).
Local form _a_ (_Gilolensis_).—Differs from the true _P. Codrus_ in
having always an additional semiovate spot below the submedian nervure,
and in having a small round spot on the anterior margin of the lower
wings beneath: it is also rather smaller.
_Hab._ Batchian and Gilolo (_Wall._)
Subspecies _b_ (_Celebensis_).—Fore wings in the male more attenuate,
with the costal margin more curved than in true _P. Codrus_; upper
surface more green and glossy; an additional large quadrate spot on the
inner margin of the fore wings. Under surface lighter brown, the whitish
marks near the anal angle wanting; a dark subtriangular band across the
cell of the fore wings. Rather smaller than _P. Codrus_.
_Hab._ Celebes, Sulla Islands (_Wall._).
Subspecies _c_ (_Papuensis_).—Hind wings less elongate than in the true
_P. Codrus_; macular band much broader, and reaching the inner margin of
the upper wings, the lower portion divided by nervures only; the band
continued on the lower wings by means of an obscure white fascia.
Beneath, the greenish white band continues on to the lower wings, but
gradually fades away after reaching the cell. Expanse of wings 4¼
inches.
_Hab._ Waigiou, Aru Island (_Wall._).
_Remarks._—This approaches the next species. Subspecies _b_ and _c_ I
consider to be really as distinct as many universally received species,
differing in form and in several points of coloration. As, however, it
is probable that there are forms in other islands which may present
intermediate characters, I prefer retaining the whole under the old
specific name.
107. PAPILIO MELANTHUS, Felder.
_P. Melanthus_, Feld. Lep. Nov. Philipp. p. 12.
_Hab._ Mindanao (Philippines).
108. PAPILIO EMPEDOCLES, Fabricius.
_P. Empedocles_, Fab. Ent. Syst. iii. 1. p. 70; Don. Ins. Ind. pl. 17.
f. 1; Boisd. Sp. Gén. Lép. p. 229.
_Hab._ Borneo (_Wall._).
109. PAPILIO PAYENI, Boisduval.
_P. Payeni_, Bd. Sp. Gén. Lép. p. 235; Van der Hoeven, Tijd. von Nat.
Gesch. v. t. 8, f. 1, 2, 6.
_Hab._ Borneo (_Wall._), Java (_Van der Hoeven_).
_Remarks._—This remarkable species has been placed by Boisduval in a
group by itself. It, however, agrees very closely in habits and
structure with this group, and can hardly, I think, be separated, though
very abnormal in colouring. _P. Evan_, Db., is a closely allied species
from India; and _P. Gyas_, Westw., from the same country, is also nearly
related, though it has been hitherto placed in another section of the
genus.
110. PAPILIO SARPEDON, Linnæus.
_P. Sarpedon_, L.; Cram. Pap. Ex. t. 122. f. D, E.; Boisd. Sp. Gén.
Lép. p. 235.
_Chlorisses Sarpedon_, Sw. Zool. Ill. 2nd ser. pl. 89.
_Hab._ Borneo, Sumatra (typical), New Guinea, Aru Is. (darker), Java
(broader band) (_Wall._).
Local form _a_ (_Moluccensis_, Cram. Pap. Ex. t. 122. f. D, E).—Black,
with the bands and spots rich blue.
_Hab._ Ceram, Batchian, Gilolo, Bouru (_Wall._). (The Ceylon form
closely resembles this.)
111. PAPILIO MILETUS, n. s. Tab. VII. fig. 2 (♂).
Wings larger and more falcate than in _P. Sarpedon_, costal margin
abruptly curved near the base of the wing.
Above, black; macular band rich blue, very narrow, the spots on the
upper wings all more or less rounded and separated by thick black bands;
the marginal lunules large and angularly bent.
Beneath, the upper wings have a row of four pearly-white lunules from
the outer angle; and there is one of the same colour at the outer angle
of the lower wings, which have also an additional red spot on the margin
of the cell, below the first branch of the subcostal nervure. Expanse of
wings 4¾ inches.
_Hab._ Macassar and Menado (Celebes) (_Wall._).
_Remarks._—I have separated this species from all the other forms of _P.
Sarpedon_, because, while they differ in markings and colour only, this
differs greatly in form as well as very strikingly in size, colour, and
markings. I cannot conceive, therefore, why such a combination of
distinctive peculiarities should not entitle it to specific rank.
112. PAPILIO WALLACEI, Hewitson.
_P. Wallacei_, Hew. Ex. Butt., “Papilio,” iii. f. 7.
_Hab._ Aru Islands, Batchian (_Wall._).
_Remark._—This isolated species is very rare: I obtained a single male
specimen in each of the above localities in the virgin forest.
113. PAPILIO BATHYCLES, Zinken.
_P. Bathycles_, Zink. Beitr. Ins. Java, p. 157, tab. 14. f. 6, 7;
Boisd. Sp. Gén. Lép. p. 232.
_Hab._ Java, Borneo, Malacca (_Wall._).
_Remark._—The Indian form generally confounded with this I consider to
be a very distinct species, for which I propose the name of _P. Chiron_,
and add a description below[15].
Footnote 15:
PAPILIO CHIRON, n. s.
_P. Bathycles_ (partly), Brit. Mus. List of Papilionidæ.
Very near _P. Bathycles_, Zinken. Fore wings rather broader at the
tip; hind wings considerably less elongate posteriorly.
Above:—fore wings have the three larger green spots separated by broad
black spaces, the first produced towards the base of the wing, the
second notched above; the fourth spot in the cell much more linear.
Hind wings have the green markings more elongate and narrower, and an
additional narrow mark at the abdominal margin.
Beneath, the spots all separated by broad black lines; the abdominal
stripe, which is quite wanting in _P. Bathycles_, larger than above;
an ochre-yellow spot on the hind wings, near the base of the inner
margin (absent in _P. Bathycles_); the submarginal pale spots larger,
and the row of reddish-ochre spots less developed. Expanse of wings 3⅓
inches.
_Hab._ Assam, Sylhet.
114. PAPILIO EURYPYLUS, Linnæus.
_P. Eurypylus_, L.; Cram. Pap. Ex. t. 122. f. C, D; Boisd. Sp. Gén.
Lép. p. 233.
_Hab._ Amboyna (type), Ceram, Bouru, Batchian, New Guinea (_Wall._).
_Remark._—The _male_ has the abdomen above and abdominal margin white;
the _female_ blackish.
115. PAPILIO JASON, Esper.
_P. Jason_, Esp. Ausl. Schmett. t. 58. f. 5; _P. Jason_, L.? _P.
Eurypylus_, var., Boisd. Sp. Gén. Lép. p. 233.
_Hab._ Malacca, Sumatra, Borneo (♂, ♀) (_Wall._).
_Remarks._—This species is readily distinguished from _P. Eurypylus_ by
the abdomen above, and the abdominal margin, being black in both sexes,
by the smaller size, more pointed upper wings, and by the lower wings
having a narrower band and larger spots. of a deeper green colour. On
the under surface the marginal lunules, the cell-spots, and sub-basal
stripe are all larger.
Variety or dimorphic form _a_.—_Evemon_, Boisd. Sp. Gén. Lép. p. 234.
_Hab._ Malacca, Java, Sumatra, Borneo (♂, ♀). (_Wall._)
This may be a distinct species, but is more probably a case of
dimorphism. The two forms are absolutely identical, except that the red
spot at the base of the lower wings beneath, in _P. Jason_, is
constantly absent in _P. Evemon_.
116. PAPILIO TELEPHUS, n. s. Tab. VII. fig. 4 (♂).
Larger than _P. Eurypylus_; anterior wings more elongated, with their
costal margin abruptly curved near the base.
Above, the four spots in the cell of the upper wings linear, of equal
width, not increasing in thickness from the base outwards, as in _P.
Eurypylus_; the macular band narrower, nearly white on the lower wings;
abdomen and abdominal margin pure white.
Beneath, the red anal spot is not produced upwards along the abdominal
margin, the pearly spots have a distinct dusky border, owing to their
exceeding in size those on the upper surface. Expanse of wings 4¼
inches.
_Hab._ Celebes (_Wall._).
_Remarks._—This is a powerful species of very rapid flight, and
difficult to capture. It comes about muddy places in the villages of
South Celebes, and is also found abundantly at pools in the half-dry
mountain-streams. I consider it quite distinct from all the allied
forms.
117. PAPILIO ÆGISTUS, Linnæus.
_P. Ægistus_, L.; Cram. Pap. Ex. t. 241. f. C, D; Boisd. Sp. Gén. Lép.
p. 231.
_Hab._ Ceram, Gilolo, Batchian, Aru Islands (_Wall._)
118. PAPILIO AGAMEMNON, Linnæus.
_P. Agamemnon_, L.; Cram. Pap. Ex. t. 106. f. C, D; Boisd. Sp. Gén.
Lép. p. 230.
This species presents numerous slight modifications of form and marking,
which seem hardly prominent enough to characterize as species, though
tolerably constant in each locality. Type tailed.
_Hab._ India, Manilla.
Local form _a_. Tail shorter; wings rather pointed.
_Hab._ Timor, Flores (_Wall._).
Local form _b_. Tail as in the last; two outer rows of spots on the
lower wings absent.
_Hab._ Ké Island (_Wall._).
Local form _c_. Size small; tail very short.
_Hab._ Malacca, Sumatra, Borneo, Java (_Wall._).
Local form _d_. Wings much elongated, abruptly curved near the base;
tail very short; size large.
_Hab._ Celebes (_Wall._).
Local form _e_. Broader and less sinuated wings, body large, tail very
short.
_Hab._ Ceram, Bouru, Batchian (_Wall._).
Local form _f_. Form of _c_; tail reduced to a tooth; markings and spots
well defined, rounded.
_Hab._ New Guinea, Aru Islands, Waigiou (_Wall._).
119. PAPILIO RAMA, Felder.
_P. Rama_, Feld. Lep. Nov. Mal. p. 1. _P. Arycles_, Boisd. Sp. Gén.
Lép. p. 231?
_Hab._ Malacca, Sumatra (_Wall._).
_Remarks._—I have little doubt but this is the _P. Arycles_ of
Boisduval. His description, however, does not mention the distinctive
character of the four large spots only in the discoidal cell; I have
therefore used Dr. Felder’s name.
LEPTOCIRCUS, Swainson.
This small but interesting genus differs somewhat from _Papilio_ in the
neuration of the wings, but is best distinguished by the longitudinal
fold and great elongation of the hind wings. The species frequent water,
often settling on the edges of rills, or hovering over pools and
rivulets in the sunshine. The few species known are all very closely
allied, and might with equal propriety have been considered as local
forms of one species. Three have been already described, and I have
therefore thought it better to add one more, than to attempt to reduce
those which have been generally accepted as species to a lower rank.
120. LEPTOCIRCUS MEGES, Zinken.
_P. Meges_, Zink. Beitr. Ins. Java, p. 161, tab. 15. f. 8.
_Leptocircus Curius_, Sw. Zool. Ill. pl. 106; Boisd. Sp. Gén. pl. 7.
f. 1, pl. 17. f. 3, p. 381.
_Hab._ Java, Malacca (_Wall._).
121. LEPTOCIRCUS CURTIUS, n. s.
Larger than _L. Meges_; outer black margin broader, and apical nervures
thicker; bluish band much narrower, of equal width on both wings,
straight, abruptly narrow where it crosses the discoidal cell of the
fore wings, and rounded at the inner margin so as to form a small notch
at the junction of the fore and hind wings.
Under side with the band bluish silvery; the three small bands on the
anal margin differing from those on _L. Curius_ and _L. Meges_, the
first being transverse, and not produced obliquely to join the vertical
band, the second small and nearly obsolete, the third at the anal angle
transverse, very little curved, and sharply defined.
Body beneath and base of all the wings greenish ashy. Expanse of wings
1⁹⁄₁₀–2 inches. Length, head to tip of tail 2⁶⁄₁₀ inches.
_Hab._ Celebes (_Wall._).
122. LEPTOCIRCUS DECIUS, Felder.
_L. Decius_, Feld. Lep. Nov. Philipp. p. 13. _L. Corion_, G. R. Gray,
List of Pap. in Brit. Mus.
_Hab._ Philippine Islands.
123. LEPTOCIRCUS CURIUS, Fabricius.
_L. Curius_, Fab. Ent. Syst. iii. 1. p. 28; Doubleday, Zoologist,
1843, p. 111; Gen. of Diurnal Lep. pl. 4*. f. 1; Don. Ins. Ind. pl.
47. f. 1.
_Hab._ Java (_Wall._), North India.
NOTE.
In referring to the species described by Dr. Felder, I have quoted from
papers which he has sent me, with distinct titles and separate paging,
but which were all first published in the ‘Wiener Entomologischen
Monatschrift,’ viz. “Lepidopterologische Fragmente” (quoted as “Lep.
Fragm.”), published at intervals from June 1859 to August 1860,
“Lepidoptera Nova Malayica” (quoted as “Lep. Nov. Mal.”), published in
1860, and “Lepidoptera Nova a Dr. Carolo Semper in insulis Philippinis
collecta” (quoted as “Lep. Nov. Philipp.”), published in 1861. It is to
be regretted that the titles and paging of these separate papers were
not made to correspond with the original publication, so as to have made
a more exact reference possible.
I have also quoted Zinken’s ‘Beitrag zur Insecten-Fauna von Java’
separated from the ‘Nova Acta Acad. Nat. Curios.’; but in this case the
pages and the numbering of the plates have been preserved as in the
original work.
EXPLANATION OF THE PLATES.
PLATE I.
Represents the various forms of _Papilio Memnon_ (see pages 6 and 46).
N.B. The left side of each figure shows the upper surface, and the
right side the under surface of the same insect.
Fig. 1. A male, from Borneo (a slight local variety).
Fig. 2. A female, from Java (a variety like _P. Agenor_, Cr.).
Fig. 3. A female, from Sumatra (a variety near _P. Anceus_, Cr.). The
last two are varieties of the 1st dimorphic form of female in this
species.
Fig. 4. A female, from Java (_P. Achates_, Cr.). The 2nd dimorphic
form of female of _Papilio Memnon_.
PLATE II.
Represents the various forms of _Papilio Pammon_ (figs. 1, 3, 5, and
6) and _P. Theseus_ (figs. 2, 4, and 7). (See pages 6, 7, 51, 52,
and 53.) N.B. The left side of each figure shows the upper surface,
and the right side the under surface of the same insect.
Fig. 1. _Papilio Pammon_; a male, from Malacca.
Fig. 3. The first form of female, closely resembling the male, from
India.
Fig. 5. The second form of female (_P. Polytes_, L.), from Singapore.
This is the most common and widely distributed form of female,
occurring everywhere with the male.
Fig. 6. The third form of female (_P. Romulus_, Cr.), from India.
Fig. 2. _Papilio Theseus_, the first form of female, almost exactly
resembling the male, from Timor. This form is very rare.
Fig. 4. The second form of female, from Timor.
Fig. 7. The third form of female (_P. Theseus_, Cr.), from Sumatra.
The second and third forms of female seem about equally plentiful,
but are generally confined to separate islands. A fourth form of
female (_P. Melanides_, De Haan) would have been figured, but could
not be brought on to the plate. (See pages 7 and 53.)
PLATE III.
Represents the various forms of _Papilio Ormenus_ (see pages 8, 55,
and 56). N.B. The left side of each figure shows the upper surface,
and the right side the under surface of the same insect.
Fig. 2. A male, from the island of Goram.
Fig. 1. The first form of female, from Waigiou.
Fig. 3. The second form of female, from Waigiou.
Fig. 4. The third form of female (_P. Amanga_, Bd.), from the island
of Goram.
PLATE IV.
Represents two species allied to _Papilio Ormenus_, but whose females
are not _dimorphic_ (see pages 57 and 58). N.B. The left side of
each figure shows the upper surface, and the right side the under
surface of the same insect.
Fig. 1. A female of _Papilio Adrastus_, peculiar to the island of
Banda (see page 57).
Fig. 3. _Papilio Tydeus_; a male, from Batchian.
Fig. 2. The female of _Papilio Tydeus_, exhibiting a single permanent
form confined to a small group of islands (Batchian and Gilolo),
intermediate between the two forms of _Papilio Ormenus_ ♀ which are
represented on Plate III. figs. 3 and 4.
PLATE V.
Represents several new species of Papilio, illustrating “local
variation.” N.B. The right side of each figure shows the upper
surface, and the left side the under surface of the same insect.
Fig. 1. The male of _Papilio Noctis_, from Borneo (see page 41). The
female was figured by Mr. Hewitson in the ‘Proceedings of the
Zoological Society of London,’ 1859, plate 66. fig. 5.
Fig. 2. _Papilio Leodamas_, male, from Mysol (see page 42).
Fig. 3. _Papilio Hecuba_, male, from Celebes (see pages 16 and 50).
Fig. 4. _Papilio Pertinax_, male, from Celebes (see page 49).
Fig. 5. _Papilio Albinus_, male, from New Guinea (see page 49).
PLATE VI.
Represents four species not before figured, belonging to the most
brilliantly coloured group of Eastern Papilios, and illustrating
local modifications of form. N.B. The right side shows the upper
surface, and the left side the under surface of the same insect.
Fig. 1. _Papilio Pericles_, male, from Timor (see page 45).
Fig. 2. _Papilio Macedon_, male, from Celebes (see page 45). This
species exhibits in a marked manner the strongly arched wings
characteristic of those from Celebes, as contrasted with those
represented at figs. 1 and 3, from other islands (see pages 16, 17
and 18).
Fig. 3. _Papilio Philippus_, female, from Ceram (see page 45).
Fig. 4. _Papilio Blumei_, male, from the north of Celebes (see page
46). This also exhibits the arched wing, as compared with its ally
from the Moluccas (fig. 3).
PLATE VII.
Represents six remarkable species of Papilio not before figured. N.B.
Except in fig. 1, the right side shows the upper surface, and the
left side the under surface of the same insect.
Fig. 1. _Papilio Thule_, male. The upper surfaces of two varieties or
local forms of this species are figured. The right side represents
the form found in New Guinea, the left side that obtained in
Waigiou. It resembles _Danais sobrina_, Bd., which inhabits the same
countries, and varies in a somewhat similar manner (see pages 20 and
63).
Fig. 3. _Papilio Ænigma_, male, from Sumatra (see page 60). This
species was named as above, from its puzzling resemblance to
_Papilio Paradoxa_, which is found in the same districts. Both
species appear to mimic _Euplœa Midamus_ (see page 20).
Fig. 2. _Papilio Miletus_, male, from Celebes (see page 65). This
species and the next exhibit in a striking manner the abruptly
curved wing peculiar to Celebes. Figs. 5 and 6 represent species
almost equally remarkable in this respect.
Fig. 4. _Papilio Telephus_, male, from Celebes (see page 67).
Fig. 5. _Papilio Androcles_, male, from Celebes (see page 63).
Fig. 6. _Papilio Gigon_, female, from Celebes (see page 59).
PLATE VIII.
Illustrates, by comparative outlines of the anterior wings, the local
modification of form in the Papilios of Celebes as compared with
those of the surrounding islands. In each pair of outlines, the
upper one represents a species peculiar to Celebes, while the one
beneath it shows the most closely allied species or variety from any
of the surrounding islands. (For details, see page 16.) The
following are the names of the species:—
Fig. 1. _Papilio Gigon_, from Celebes; _P. Demolion_, from Java.
Fig. 2. _Papilio Macedon_, from Celebes; _P. Peranthus_, from Java.
Fig. 3. _Papilio Androcles_, from Celebes; _P. Antiphates_, from
Borneo.
Fig. 4. _Papilio Telephus_, from Celebes; _P. Jason_, from Sumatra.
Fig. 5. _Papilio Miletus_, from Celebes; _P. Sarpedon_, from Java.
Fig. 6. _Papilio Agamemnon_, var., from Celebes; _P. Agamemnon_, var.,
from Sumatra.
[Illustration:
TRANS LINN SOC. VOL. XXV TAB 1
]
[Illustration:
TRANS LINN SOC VOL XXV TAB 2.
]
[Illustration:
TRANS LINN SOC VOL XXV TAB. 3.
J. O. Westwood, del. Day & Son, lith^{rs} to the Queen.
]
[Illustration:
TRANS LINN SOC VOL XXV TAB 4
]
[Illustration:
TRANS LINN. SOC., VOL. XXV, TAB 5
J. O. Westwood, del. Vincent Brooks, Imp.
]
PLATE VI.
Represents four species not before figured, belonging to the most
brilliantly coloured group of Eastern Papilios, and illustrating
local modifications of form. N.B. The right side shows the upper
surface, and the left side the under surface of the same insect.
Fig. 1. _Papilio Pericles_, male, from Timor (see page 45).
Fig. 2. _Papilio Macedon_, male, from Celebes (see page 45). This
species exhibits in a marked manner the strongly arched wings
characteristic of those from Celebes, as contrasted with those
represented at figs. 1 and 3, from other islands (see pages 16, 17,
and 18).
Fig. 3. _Papilio Philippus_, female, from Ceram (see page 45).
Fig. 4. _Papilio Blumei_, male, from the north of Celebes (see page
46). This also exhibits the arched wing, as compared with its ally
from the Moluccas (fig. 3).
Wallace, A. R., “On the phenomena of variation and geographical
distribution as illustrated by the Papilionidæ of the Malayan region,”
_The Transactions of the Linnean Society of London_, V. 25, 1866, p.
1–71.
[Illustration:
TRANS LINN SOC. VOL XXV, TAB 6
J. O. Westwood, del. Vincent Brooks, Imp
]
[Illustration:
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J. O. Westwood, del. Vincent Brooks, Imp
]
[Illustration:
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J. O. Westwood, del. Vincent Brooks, Imp
]
------------------------------------------------------------------------
TRANSCRIBER’S NOTES
Page Changed from Changed to
75 Trans Linn Soc Vol XXV Tab Trans Linn Soc Vol XXV Tab 2.
● Typos fixed; non-standard spelling and dialect retained.
● Used numbers for footnotes.
● Enclosed italics font in _underscores_.
● The caret (^) serves as a superscript indicator, applicable to
individual characters (like 2^d) and even entire phrases (like
1^{st}).
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On the phenomena of variation and geographical distribution as illustrated by the Papilionidæ of the Malayan region
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[_From the_ TRANSACTIONS _of the_ LINNEAN SOCIETY, vol. xxv.]
I. _On the Phenomena of Variation and Geographical Distribution as
illustrated by the_ Papilionidæ _of the Malayan Region. By_ ALFRED R.
WALLACE, _Esq._
When the naturalist studies the habits, the structure, or the affinities
of animals, it matters little to which group he especially devotes
himself; all alike offer him endless materials for observation and
research. But, for the purpose of...
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