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[ From the Transactions or THe Linnean Socrery, vor. XXVI. |

On some remarkable Minetice Analogies among African Butterflies.
By Rovranp Trimen, Mem. Ent. Soc. Lond.

(Plates XLIT. & XLIII.)
Read March 5th, 1868.

xe * * * «<Cest une chose bien remarquable que de voir la nature créer A cété les uns des autres
PEuplea Niavius, le Diadema dubia, et le Papilio Westermanni, trois Lépidoptéres qui se ressemblent
presque complétement par le port, le dessin, et la couleur, quoique appartenant A des genres fort éloignés
et de tribus différentes.”’—Botspuva, Species Général des Lépidopteres, pp. 372, 373.

FROM the year 1836, when Dr. Boisduval published these remarks, a period of twenty-
five years elapsed without any light being thrown upon the meaning of those remarkable
resemblances among the Rhopalocera which are familiar to every lepidopterist, which
have been noted by entomologists in publications both prior and subsequent to the date
of the ‘Species Général,’ and to one of the most striking of which the above quotation
refers. ‘The extraordinary fact, that in all parts of the world species of Butterflies
occurred which, aberrant from the normal facies of their immediate allies, most closely
resembled other species of wholly different structure, awakened no comment beyond the
admission that it was curious, unless it were some vague suggestion as to “ recurrent
types”? in nature, which left the subject as completely mysterious as before. Entomo-
logists, no less than naturalists generally, appeared content with a child-like wonder at
this and kindred facts, and let them pass as things inscrutable. That this neglect of
inquiry was due in great measure to the absence of reliable observations upon the living
insects in their native haunts, cannot be doubted; but it may be questioned if, with all
the data now accumulated by various explorers, those whose energies were necessarily,
and in many cases exclusively, concentrated upon the arduous work of the systematist
would have been enabled to elucidate the subject. It remained for one of those adven-
turous lovers of nature whose zeal for discovery leads them to years of tropical wander-
ings, to indicate, from his own assiduous observations, and in the light of that compre-
hensive theory of organic nature which we owe to Mr. Darwin, the rational explanation
of these phenomena. That this explanation is absolutely conclusive, or not susceptible
of future modification, it would be premature to assert; but it is indisputably the only
advance yet made towards the solution of the difficulty, and so reasonable a demonstration
as to commend itself to every thoughtful observer. I need scarcely say that I refer to
the well-known treatise by Mr. Bates on the Heliconide of the Amazons Valley (published
in the twenty-third volume of the ‘ Transactions of the Linnean Society ’), in which the
principle of natural selection is most ably applied in elucidation of the origin and
development of those “‘ mimicries ”’ of which many are now so astonishingly exact. The
VOL, XXVI. 3Z

Qa»

498 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

view propounded by Mr. Bates has met with the weighty support of Mr. Wallace, who,
in his interesting paper on Malayan Papilionide*, has called attention to an analogous
series of phenomena presented in India and the great Eastern Archipelago, and can
only account for them on the same theory.

My object in the present paper is to give some account of the most striking cases of
mimetic analogy which have been found to exist among the Butterflies of Africa, more
especially regarding those occurring at the southern extremity of the continent, a region
in which I have had the advantage of several years’ personal research.

In the first place, it is eminently worthy of remark that the Butterflies which are the
subjects of imitation by others, belong to the same families both in the Old World and
the New. The Danaide and Acreide, throughout the warmer parts of the earth, are
unquestionably the special groups that furnish the models after which more or less
perfect copies have been elaborated. This being an acknowledged fact, we are naturally
led to inquire why it should be so—what advantage is to be gained by closely resembling
the members of those families. Is there anything connected with those groups tending
to show that they possess advantages above others of their Order? Do they appear,
from their numbers and habits of life, to be dominant races ?

These inquiries can unhesitatingly be answered in the affirmative. ‘The slow flight,
the conspicuous colours, the complete disregard of concealment, no less than the great
abundance of individuals, are characteristics indicating unmistakably that these Butter-
flies are favoured races, enjoying advantages and immunities above their fellows. I
believe that Mr. Bates (loc. cit. p. 510) has correctly suggested the principal causes of the
evident security of these insects, viz. their emitting an unpleasant odour, and being proba-
bly distasteful to insectivorous animals. The peculiar smell noticed by Mr. Bates in the
Heliconide Danaide is also possessed by the Huplee of the Hastern Archipelago, as Mr.
Wallace has recorded; and I am enabled to add similar evidence as regards the African
species of Danais and Acreat. Judging from the case of dcrwa Horta, a widely-
distributed African species, which I have reared in considerable numbers from the young
larvee, this disagreeable smell is not peculiar to the imago condition, but attends the insect
throughout its life, even the pupe emitting it. On pressing the thorax of a Danais,
a Euplea, or an Acrea, however lightly, a clear yellow liquid, similar to that secreted
by the Ladybird beetles, almost always exudes from that part of the body; and in some
species of the two former genera a pair of bright-yellow fascicled appendages are protruded
from the extremity of the abdomen}. The peculiar scent appears chiefly to reside in
this liquid, being remarkedly stronger with each effusion. Most of the species of Danais
and Acrea feign death very readily ; and they possess another means of defence which,
as far as 1 am aware, has not hitherto been recorded, viz. the remarkable elasticity of
their entire structure. No pressure of the thorax, short of absolute crushing of the
tissues, suffices to kill or even paralyze these Butterflies ; and the collector who treats

* Trans. Linn. Soe. vol. xxv.

+ The Mauritian Huplaw Euphone also gives out a strong odour when handled.
+ Mr. Bates mentions that species of the genera Lycorea and Ituna (Heliconide Danaidw) possess similar exsertile

organs.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 4.99

them as he would species of other families, soon finds his collecting-box alive with its
struggling occupants. So flexible are the wings, that the insect generally sueceeds in
withdrawing them from crossed fences of pins which form a complete barrier to any
motion on the part of ordinary Butterflies ; and however bent and distorted the wings
may become in such exertions, I have never known a fracture of nervures or membrane
to result, the organs resuming their natural position even after having been bent double
for some hours. While entomologizing in Natal, my Kafir collector used often to bring
me numerous examples of the commonest species in his box, and when engaged in the
necessary work of rejection, I constantly found the limp-winged Danaide and Acreide,
as soon as they were released from the transfixing-pin, fly off with perfect ease and
apparent nonchalance.

It is not difficult to perceive how important, as a reserve means of defence, this
unusual elasticity of structure may prove. That birds, and other eaters of insects, may
occasionally capture a Butterfly of these malodorous tribes before discovering its distasteful
character is not an unreasonable supposition, especially in seasons when an exceptional
searcity of some favourite food may prevail. In such a ease it may be safely stated that
the chances are very greatly in favour of a Danais or an Acrea escaping, if not wholly
unhurt, yet without serious injury, after rough treatment that would have proved fatal
to a harder but less elastic animal*.

More extended observation is necessary to prove by positive evidence that persecution
of the Danaide and Acreide has almost wholly ceased, although, negatively, the witness
borne by their abundance is very strongly in favour of this view. Some check upon
their inordinate increase must, of course, exist; and this is probably to be found in the
Ichneumon flies attached to the various species, the females of which are not likely to
be deterred by any odour from depositing their ova in the larvee. Mr. Bates has re-
corded that the Heliconide Danaide+, when at rest on leaves, did not “appear to be
molested by lizards or predaceous flies of the family Asilide, which were very often seen
pouncing on Butterflies of other families.” A well-known naturalist has quoted (in the
Westminster and Foreign Quarterly Review for July 1867, Article 1) an observation
by My. Belt, that a pair of Puff-birds, watched by him in Brazil, during half-an-hour
caught and brought to their nest various butterflies to feed their young, but entirely
avoided the slow-flying Heliconide, which were about the spot in great numbers}. I

* This elasticity of structure is not confined to the Butterflies in question, being a character of many Moths, and
markedly of some belonging to the families Agaristide and Zygenide. It is remarkable that the three South-African
Moths in which I have found this peculiarity most developed, viz. Pais decora, Husemia euphemia, and Gilaucopis
formosa—all have a strong and offensive odour, emit drops of white or yellow fluid, and are slow-flying, brightly
coloured, and abundant species.

+ The term used by Mr. Bates is “ Danaoid Helicontd@”; but as it is clear that that author agrees with Dr. Felder
in recognizing all the so-called /Heliconide except Heliconius and Hueides as forming a part of the family Danaide, |
have ventured to transpose the words in order the more distinctly to express this relationship.

¢ It is not uncommon to hear doubt expressed as to whether birds even feed on butterflies at all; and thus the
case Just mentioned is of importance. I have noticed a Swallow chasing a Pieris Brassice in England; and I never
knew it to be disputed that the Goatsuckers eat the nocturnal Lepidoptera. But when one reflects on the host of
insectivorous birds in all (and especially in tropical) countries, it would be marvellous indeed if they neglected the
large supply of food afforded by the generally defenceless race of Butterflies.

322

500 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

have published (* Rhopalocera Africze Australis,’ pt. ii. p. 835) an interesting note by
Mr. J. H. Bowker on the manner in which Junonia Enone, one of the Nymphalide, is
systematically hunted down by a small Kaffrarian lizard. This Junonia is a particularly
wary, active insect, and must give its pursuers infinitely more trouble to capture than
would the slow, inert, grass-loving dAcree@ of the same district; yet the lizards leave the
latter unmolested. On more than one occasion, I have seen the larger Dragonflies catch
Butterflies, both in England and Natal. Pierid@ were the victims in both countries ; but
in Natal the ubiquitous decree were certainly passed by, and the more active insects
selected. In the colony in question, several species of Acacia secrete a viscid fluid; and
the spaces on the branches where this freshly exudes from the bark are a great resort of
‘insects of all Orders, which assemble to imbibe the liquid. The larger Mantide take
advantage of these gatherings to secure a plentiful and easy living; and one of them is
usually to be seen among or close to the group of butterflies, beetles, wasps, ants, and
flies attracted to the feast. As long as there is anything left to drink, there is no lack
of visitors, and the Mantis fattens on numerous victims. The wings rejected by the
devourer, who seems to prefer butterflies to other insects, in a short time rather con-
spicuously sprinkle the ground or herbage under one of these feeding-places; and in a
few instances, where I chanced upon accessible spots, I searched among the disjecta
memobra for remains of Danais or Acrea, but in vain. It is necessary to observe that I
cannot recollect having distinguished Butterflies of either of those genera at such drinking-
stations; but the exudations were often situated at a height that precluded the certain
distinction of any but very large insects, and there is apparently no reason why the
fluid so universally appreciated should fail to attract those butterflies.

I think that the facts already placed on record, to the more important of which I have
referred, may fairly be urged in support of the position that the Danaide and Acreide
are exempted or protected in a very great degree from the attacks of those enemies to
which the diurnal Lepidoptera generally, in their adult state, are constantly exposed.
There are many instances of Butterflies, of various families, whose great abundance indi-
cates that they have, by some means, attained immunity from persecution, or risen
superior to adverse circumstances; but, as groups, the Danaide and Acreide, wherever
they occur, are unquestionably preeminent in individuals, if not in species, and must
take rank as the most perfectly adapted of all their Order to the existing conditions of
life.

It is not surprising to find that those families of Butterflies which hold the first place
in the battle of life display a structure which has led Mr. Bates to assign to them a
position, in a “natural system” of classification, at the head of the order Lepidoptera”.
The atrophy of the fore legs is a special character of the Rhopalocera as a group, which
most widely distinguishes them from the Heterocera and from other Orders of insects ;
and this has been tacitly recognized by the universal consent with which entomologists
have assigned the lowest place among Butterflies to the Hesperiide, a family which
shows its affinity to the Moths by the fully developed fore legs, as well as by other cha-

* See Trans. Linn. Soe. vol. xxiii., and Journal of Entomology, December 1861, and No. X. 1864.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 501

racters of importance. The splendid family Pupilionide, by mere force of the size,
number, and beauty of its members, has so long held the place of honour in English and
in most continental classifications, that many lepidopterists are loth to supplant it by a
comparatively insignificant-looking group; but, regarding the question structurally,
there can be no doubt that an arrangement which interposed between the only familics
with fully developed fore legs (viz. the Papilionide and Pieride at one end of the
series, and the Hesperiide at the other) all the groups that more or less widely differed
in the character of those very organs was artificial and unnatural. On the other hand,
Mr. Wallace (/oc. cit. p. 2) has questioned the propriety of claiming a high position for
any group on the ground of extreme imperfection of any of its organs. Taken apart
from other considerations, this at first sight appears a valid objection; but it must be
borne in mind, as suggested by Mr. Bates, that the Lepidopterous type among Articu-
lata, like that of the Bird among Vertebrata, is preeminently aerial, and, consequently,
that a diminution of the ambulatory organs, instead of being a sign of inferiority,
may very possibly indicate a higher (because more thoroughly aerial) form. Mr. Wal-
lace further contends for the first rank being accorded to the Papilionide, on account
of the perfect insects possessing the peculiar and constant character of an apparently
4-branched median nervure, and a “spur” * on the anterior tibixe, and the larve having
an extrusible Y-shaped tentacle. The appendage to the anterior tibie is admitted by
Mr. Wallace to be a character of some Hesperiide ; and not having been found in other
Butterflies, it may fairly be regarded, in conjunction with the full development of the
first pair of legs, as a sign of affinity, however distant, between the two families. The
apparent fourth branch of the median nervure is not an additional nervule, but actually
* of Doubleday) unusually placed in relation
to the third branch of the nervure, and thus can hardly be regarded as in any way signi-
ficant of superior developmentt. Nor can the Y-shaped tentacle of the larva be insisted

the lower radial (‘‘ second discoidal nervule’

* This is more strictly a small foliated expansion or appendage.

+ More remarkable points in the neuration of the Papilionide are the following, viz. :—Ist, the short, transverse,
interno-median nervule, uniting the median and submedian nervures of the fore wings, and closing a small basal
cell; 2ndly, the well-marked internal nervure of the same wings, which has an independent course, and terminates
on the inner margin; and, 3rdly, the distinct prediscoidal cell of the hind wings, formed by the junction of the
branched precostal nervure with the costal. Papilio (including Ornithoptera) appears constantly to present these
characters, with some variation in the size of the prediscoidal cell of the hind wings; the Australian Eurycus also
possesses them all, and has the prediscoidal cell unusully large; in Sericinus, Teinopalpus, and Leptocircus appear
the internal nervure (but nod the interno-median nervule) of the fore wings and a much-narrowed prediscoidal cell ;
Parnassius and Thais present only the internal nervure of the fore wings; and in the aberrant Dovitis I can trace
none of the three characters. In other families we find the transverse interno-median neryule in several of the Mor-
plade, though not so completely developed ; the internal nervure is present in the Danaide, but, instead of having an
independent course, is intimately connected with the submedian nervure, and ends by anastomosing with it; while
the prediscoidal cell of the hind wings has hitherto been employed as the distinguishing feature of the curious Bras-
solide, in which it is formed in precisely the same manner as in the Papilionide.

The presence of additional cells, enclosed by anastomosing nervures, is a feature not rare among the Heterocera
(vide the plates of neuration in Guenée’s * Noctuélites’ and ‘ Phalénites’ ); and in some genera of Geometrae (Opo-
rabia, Ennomos, &c.) a cell is found at the base of the hind wings, occupying exactly the same position as in the
Butterflies mentioned.

502 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

on as indicating perfection of the Rhopalocerous type, when it is considered that the
only known instances of an organ similar in structure and function in the rest of the
Lepidoptera are found, not in Butterflies of any family, but in Cerwra, a genus of Bom-
bycide Moths. The organ of the Puss-Moth caterpillars is double, instead of forked, and
occupies the opposite extremity of the body; but it is plainly homologous to that of the
Papilio larvee, and is protruded on occasion in precisely the same manner. Looking to
the pupz of the several families, we find, receding from those of the Heterocera in
silken, earthen, or other cocoons, or buried in the ground, a gradual advance, cor-
responding with that towards a more aerial type of imago, to the freely suspended chry-
salis of the tetrapod Rhopalocera. The pupze of the Hesperiide are secured, like those
of many moths, either in a slight cocoon or by several silken threads; those of the Papi-
lionide, Pieride, and Lycenide*, by the tail and by a single silken girdle; while,
through the Hrycinide, which present instances both of girt and freely suspended pupet,
there is a gradation to the chrysalides of the true Nymphalide Butterflies, which hang,
head downwards, by a caudal attachment only. The remarkable pupa of Parnassius,
more heteroceroid (in its blunted form, bluish efflorescence, and numerous threads of
support in a cocoon of leaves) than most of those known among the Hesperiide, seems
to afford an additional indication of some connexion between the Papilionide and Moths.
Another singular fact tends to strengthen the idea of a remote, but distinct relationship,
viz. that the pupz of Hphyra, a Geometrine genus, are not only suspended by the tail
and a silken girth, in precisely the same position as those of the Papilionide, hut closely
resemble them in form, M. Guenée observing (‘ Phalénites,’ ix. p. 402) that they can
best be compared with the chrysalides of Thaist. Similar, also, in their caudal attach-
ment and silken girdle, but not so like the Papilio chrysalides in form, are the pupe of
the very curious Heterocerous group of Péerophoride, which stands alone in the structure
of its wings, and has hitherto, by common consent, been placed last in the entire order
Lepidoptera.

It thus appears that the Papilionide exhibit points of structure in common with some
Heterocera in each stage of their growth; and these characters seem, in the aggregate, of
sufficient importance to warrant our assigning to the family a position much nearer to
the Moths than that which, until lately, has so generally been accorded to it.

Turning tothe Danaide and dereide, which have so much in common both in strue-
ture and habits, it is curious to find how the two families differ in their early states. The
larve of the Old-World Danaide are smooth, but provided with several pairs of long
fleshy filaments; and, from the case of Mechanitis Polymnia (mentioned by Mr. Bates,
loc. cit. p. 496), the Heliconide Danaide also present a smooth larva, which, however, is
furnished with tubercles instead of filaments. The dereide (of Africa) are, in the
larva-state, densely studded with stiff branched spines; and the true (or Acreoid) Heli-

* The larvee of some Lycenide (e. g. Thecla Quercus, L.) are stated to bury themselves in the earth before assuming
the pupal condition.

+ See Mr. Bates’s ‘Catalogue of Erycinide,’ in Journ. Linn. Soc., Zool. ix. p. 368.

+ Plate 2 of M. Guenée’s volume gives an interesting figure of the pupa of an Hphyra in its natural position.
Thais is recorded as having the chrysalis enclosed in a slight web, which is not the case with Hphyra.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 503

coniide of America are similar in this respect, both groups showing affinity to the Myin-
phalide as far as the caterpillars are concerned. The thick, blunt chrysalis of the
Danaide Butterflies also differs widely from the elongate, slender, and subangulated
pupa of Acrwa.

The African continent is very poor in Danaide, only eight species, belonging to two
genera*, being known to inhabit its vast area. Seven belong to the genus Danais; and
these, with the exception of D. Chrysippus, form a section confined to Africa, Mada-
gascar, and Mauritius, and distinguished by a peculiar coloration of conspicuous ochre-
yellow or white bands and spots on a black ground. Three of the six species appear to
be confined in their range to the western coast, between Sierra Leone and Angola; one,
with an equal range in the west, extends to the Zambesi and Natal; while two seem to
be limited to extra-tropical Southern Africa. It is most remarkable to find, in this very
limited number of Danaide, that no less than four are accompanied throughout their
range each by its closely-imitating Diadema, one of the four (Danais echeria) being
further attended by three mimicking species of Papilio, and another by one Papilio that
correctly copies it. Danais Chrysippus, as is well known, has a faithful imitator every-
where, except in Europe, in the 2 Diadema Bolina, and, in the south of Africa, finds
a second mimicker in a rare form of Papilio, apparently peculiar to that region.

The Acreide have their metropolis in Africa, about fifty species being known from
all parts of the Continent. Six Acree are distinctly the objects of mimicry by certain
Nymphalide and Papilionide+. Four of these are West African only; one extends,
under a somewhat altered form, to South Africa; and the sixth appears peculiar to the
latter region. The mimickers here, also, are constant attendants on the species imitated ;
and in three cases, where the sexes of the Acree are dissimilar, the sexes of the mi-
mickers differ accordingly.

It is worthy of notice that the mimicking Papiliones, whether imitators of Danaides
or Acree, are very frequently only the females of the species, to the exclusion of the
males, which, in such cases, wear the normal aspect of their immediate congeners?.
Among the Diademe and other Nymphalide, on the contrary, both sexes of the mimickers,

* Euplea Goudotii, well known as a native of Bourbon, is included, on the authority of Sir Andrew Smith, who
presented a specimen from “South Africa” to the British Museum.

+ The magnificent Papilio Antimachus, Drury, of which but one specimen is known to science, is very Acreiform
in habit, and is possibly an instance of special modification in imitation of some gigantic Acraa as yet unknown, or
perhaps extinct. Drury’s having “received” this Papilio from Sierra Leone is, unfortunately, no clue to its actual
habitat, the principal sea-ports in many parts of the world usually getting the credit of all natural objects, from
whatever distance inland, that are shipped from them to Europe. Antimachus has the aspect of a slow and heavy
flier; and the observation by Smeathman, quoted by Professor Westwood (Are. Entom. 1. p. 146), did not originally
apply to the great Papilio, but to Charaves Camulus, Drury (= C. Castor, Fab.), as may be seen on reference t
Drury’s 3rd volume, pp. 15-23, and pl. xxx.). Donovan, in his * Naturalist’s Repository,’ first misquoted the passage
respecting the flight of the Charaves.

+ Among fifteen Indian and Malayan imitative Papiliones tabulated by Mr. Wallace (loc. cit. p. 20), seven are
expressly mentioned as females. As regards the African species, Papilio Ridleyanus is an instance of both ¢ and &
wearing the aspect of an Acrva, and, in a less degree, the southern form of Papilio Leonidas, which is the least
accurate of the four mimickers of Danais Echeria; but in this case the majority of the specimens that most resemble

the Danais are females.

504:

as a rule *, present, with equal exactness, the appearance of the butterflies imitated.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

It

would almost appear from this that the Nymphalide imitators stood in need of a more
complete protection than the mimicking Papiliones; and I find that, in general, they
are comparatively fewer in individuals than the latter.

The following able exhibits a view of the more striking cases of mimicry that have
come under my notice, and also indicates the localities in which each model and its
copies are known to occur together :—

Tabular View of the most remarkable Mimetic Analogies among African Rhopalocera, with the Localities in which the

several Species concerned in each case are known to co-exist.

Dana”.

Danais Damocles .............

Angola. 2

Danaisphpialeaeptlels\ieverleieicece

Sierra Leone.
Ashanti.

ManaisMH\e beri avec scdeieissereisie cts =

Type
Knysna, Cape Colony.
Bashee River, Kaftraria.
Port Natal.

AV (ELT Sfeoraenate Cexeaisoa conolcrexettons istane faves

Knysna.
Bashee River.
Port Natal.

IDENENRINIEK AS oopododsondosods

Type
Sierra Leone.
Ashanti.

AViGit US UNE remtlc: herete ene serere eveyeee

Port Natal.

Danais Chrysippus
Type.

Sierra Leone.

Ashanti.

St. Helena.

Damara-land.

Bechuana-land (Motito).

Transvaal (Potchefstroom).

Cie Town a Cape
Knysna e

Ble riariberg Bay ) pen,
Bashee River.
Umquabaaba
D’Urban
Pietermaritzburg
St. Lucia Bay.
Madagascar.
Bourbon.
Mauritius.
Zambesi.
Somauli-land.

pau

Var. Alewppus, Cr... .+ 2.25...

Sierra Leone.
Var. Dorippus, Klug.
D’Urban, Natal.

Diadema Anthedon

Diadema Bolina °

NYMPHALID®.

Diadema Damoclina, n. s.
Angola.

Diadema dubia.

Sierra Leone.
Ashanti.

Diadema mima, n.s.............

Port Natal.

AtNaNe ae Ganoatioe daa ceemnae
Sierra Leone.
Ashanti.

VOU RRUSLIAr PeMar iii eedtecia ech aer
Port Natal.

rope:
Sierra Leone.
Ashanti.
St. Helena.
Damara-land.
Bechuana-land (Motito).
Transvaal (Potchefstroom).
Cape Town Woane
Knysna bs Col
Plettenberg Bay ip AoE
Bashee River.
Umquabaaba a)
D’Urban pNatal.
Pietermaritzburg. }
St. Lucia Bay.
Madagascar.
Bourbon.
Mauritius.
Zambesi.
Somauli-land.

Var.
Sierra Leone.

Var. Inaria, Cr.
D’Urban, Natal.

PAPILIoONID”.

Papilio Merope (1st form
Cenea, Stoll).

Type
Knysna, Cape Colony.
Bashee River, Kattraria.
Port Natal.

Port Natal.

Papilio Merope Q (2nd
form Hippocoon, Fab.).

Type.
Sierra Leone.
Ashanti.

Var. austr.

Port Natal.

Papilio Merope 2 (8rd
form, Trophonius, W est-
wood).

(Knysna.
Plettenberg Bay.
Bashee River.

Papilio echerioides, 9
Var.

Tsomo River, Kaf-
fraria.
Type. Papilio Leonidas.
Var. austr. (Bra-
sidas, Felder).
Bashee River.
Port Natal.

Tsomo River.
Natal.

* Diadema Bolina is a notable exception.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 505

Table (continued).

ACR#IDE.

Acreea Gea g and 2
Sierra Leone.
Ashanti.
Calabar.

Acreea Euryta

Viairsy Olereeierieitelertorersis « vsfevs o's
Calabar.

Wer, © scoscmappooocpannooe
Acrea Aganice J
Port Natal.
PACT EBMIDY,COD)s, ort) ers) eloreieie! 6 etieieress

Sierra Leone.
Calabar.
Acrea Zetes 5
Type
Ashanti.
Calabar.
Wer DE Ee ogoncedeeemeoane
Port Natal.
Acrea gina ¢ and 9

™ ,
Congo.

NYMPHALID&.

EuRryTELIDA.

PAPILIONID®.

Panopea Hirce ¢ and 9
Type.
Ashanti.
Calabar.
Pan opeasHircey eee. sss cies «cle
Var. 9.
Calabar.
Var. 9.

Congo.

Panopea Tarquinia, ¢.
Port Natal.
Panopea Lucretia.
Sierra Leone.
Calabar.
Panopea Boisduvalii.
Type.
Ashanti.
Calabar.
Var. austr.
Port Natal.

(Melanitis Phegea
Ashanti.

| Melanitis Phegea.

Pg.
9.

Calabar.

Var. 2 (Bammuakoo, W estw.

Papilio Cynorta 9
(P. Botsduvallianus,
Westw.)
Sierra Leone.
Ashanti.
Calabar.

Papilio Ridleyanus ¢
and
Congo.

|
|

In order to give a clear idea of the nature of these cases of mimicry, it is necessary to
consider them separately. I will therefore describe them seriatim as concisely as pos-
sible, only premising that, as regards Tropical Africa, so little is on record respecting the
range of individual species or their variation, to say nothing of habits and stations, that
it is at present impossible to treat satisfactorily of the instances occurring in that
immense tract of country.

1. Danats Damoctgs, Fab.
Danais Damocles, Ent. Syst. iti. 1. p. 41. no. 121; Palisot de Beauvois, Ins. Afr. et Am. Lep. t. 6.
figg. 3a, 3.

This species is rather widely spread in tropical Western Africa, examples having been
received from Sierra Leone, Ashanti, the Gaboon, and Angola (in about 8°S. lat.). It
is nearly related to D. Lgialea, Cram., but is larger, and may be recognized by the
broader central white band of the fore wings (which extends below the first median
nervule), by the more irregular subapical white bar of the same wings, and by the
smaller and whiter basal space in the hind wings. The Gaboon examples form a variety
in which the pale basal patch in the hind wings is either reduced to a small spot or
altogether absent. In Angola this Danais is accompanied by a Diadema (which also
inhabits Congo) that very closely imitates it, differmg from Diadema dubia, its nearest
ally*, precisely as Danais Damocles differs from its congener Eyialea. 'The two species

* JT propose for this butterfly the name of

Drapewa Damocrra, n. sp.
Exp. 3 in.-3 in, 5 lin.
VOL. XXVI.

Fore wing: central white bar much broader,
4A

Closely allied to D. dubia, Pal. de Beauv.

506 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

of Danais appear to exist side by side at Sierra Leone and in Ashanti; and it is probable
that the mimickers of Damocles will eventually be found in those localities from which
Diadema dubia has already been brought.

2. Danais Eetauea, Cram.
Danais Egialea, Pap. Exot. pl. 192. fig. D.

This Danais has apparently a less extensive range than D. Damocles, inhabiting
Sierra Leone, Cape Palmas, and Ashanti. It is most exactly copied by Diadema dubia,
Pal. de Beauv., the only noticeable difference being that the Diadema has three or four
additional dots near the hind margin of the hind wings. There are specimens of this
Diadema in the British Museum, from Sierra Leone and Ashanti; and in Mr. Hewitson’s
collection, from Calabar. Palisot de Beauvois records it as a native of Oware and Benin,
the latter district being situated between Ashanti and Calabar.

3. Danais Ecuerta, Stoll. (Tab. XLII. fige. 3, 7.)
Danais Echeria, Suppl. Cramer, Pap. Exot. t. 29. figg. 1, 1a.
Vaillantiana, Godt. Enc. Méth. ix. p. 183. no. 25.

D. Echeria is a widely-spread and abundant butterfly throughout the wooded parts of
South Africa, where it takes the place of its tropical ally, D. Hgialea. Like the latter,
it has an exact imitation in a Diadema, which, though nearly allied to D. dubia, appears
to be distinct*. But Hcheria has no less than three other imitators in the genus Papilio,
the two more accurate mimickers—P. echerioides, Trimen, ?, and P. Merope, Cram., 9
(= Cenea, Stoll)—being females of very dissimilar males, and the third a variety of
P. Leonidas, Fab., which seems only to occur in South Africa beyond the tropic, and
in which the pale markings are almost or wholly devoid of the green or greenish colouring
of the type-form, and several of the lesser spots generally wanting. All these mimicking
species are much rarer than the Danais—especially the Diadema, of which I met with
only two examples in Natal, and have seen but five others in collections. Papilio eche-

continuous, not interrupted on median nervure; subapical bar interrupted, narrower, more macular. Hind wing:
whitish basal space reduced to a small ovate marking, occupying discoidal cell (and extending beyond and above it),
well defined, instead of gradually fading into the ground-colour; the submarginal white spots much smaller, some
of them occasionally wanting ; ground-colour darker than in D. dubia. Unprrstpe.—aAs in D. dubia, but with the
white bars differing in the same manner as on the upperside.

Hab. Angola.

In the collection of the British Museum.

* T append a diagnosis of this species :—

DriapEMs Mima, n. sp. (Tab. XLII. fig. 7.)

Exp. 3 in. 7 lin—3 in. 8} lin. Nearly allied to D. dubia, Pal. de Beauy. Fore wing: white spot next base, in
discoidal cell, much smaller, or nearly obsolete ; subapical white band narrower, and distinctly composed of three
spots; white spots of submarginal row smaller; no trace of ochreous scaling on inner margin. Hind wing: pale
central space broader, uniformly yellow-ochreous. Unxperstpn.—Differs similarly from that of D. dubia: white spots
at bases of wings smaller. Fore wing: the bluish edging of the larger markings entirely wanting. Sexes similar.

Hab. Natal.

In the Collections of the South-African Museum, W. C. Hewitson, and R. Trimen.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 507

rioides is rather local than scarce, only occurring in woods at a considerable elevation ;
and, from Mr. Bowker’s observations in Kaffraria, the ¢ would seem to be little, if at
all, rarer than the ¢, though, in the only Natalian habitat in which I found the species
tolerably common, the females were very much scarcer than individuals of the opposite
sex. As regards the ¢ P. Merope*, however, I have no doubt of its comparative
rarity, as one may take males abundantly in the forests for days together, without once
meeting with a female.

The resemblance to Hcheria presented by the austral variety of Papilio Leonidas
(P. Brasidas, Felder, Spec. Lep. p. 19, no. 249), appears in the cabinet but a slight one
in comparison with the striking imitations just mentioned; but it is a fair-enough like-
ness in nature, especially when the insect settles +. Certain specimens (usually, but not
invariably, females), which have the spots of the fore wings reduced in number and
diminished in size, are much more like Acheria than others; and, looking at the insect
in comparison with the type Leonidas of Western Africa (which resembles no known
Danais), I have no doubt that the southern form has been, and is probably still being
gradually modified in the direction of the dominant southern Danais.

Danais Echeria presents two varieties as regards the colour of the pale spots in the
fore wings—one (the type) in which those spots are ochre-yellow (fig. 3), and the other
in which they are white (fig. 7). The former is the prevalent form in the Cape Colony,
and the latter in Natal; but I have taken both forms in each Colony, as well as inter-
mediate examples in which the spots near the costal margin of the fore wings are white
or whitish, while the rest are yellow. The Cenea-form of the ¢ Papilio Merope mimics
both these varieties, and offers corresponding intermediate specimens; while the 2 P.
echerioides, which only inhabits the eastern portion of South Africa, almost always
resembles the white-spotted variety, though I have seen, too, unusually small specimens
which copy the yellow-spotted Echeria.

With reference to Papilio Merope, 1 think it well in this place to offer some observa-
tions on what I believe to be a remarkable instance of polymorphism in the ¢ of this
species. First figured by Cramer (in 1779 and again in 1782), P. Werope has long been
known as a native of Western Africa, and more recently as also inhabiting Southern
Africa and Madagascar {. Its coloration is very conspicuous, and unlike that of any
other Papilio, the upper surface being uniformly pale sulphur, or creamy-yellow, the
fore wings with a narrow costal and broad hind-marginal black border, and the hind
wings (which are tailed) having a more or less broken black band across the disk, and
some black hind-marginal lunules. The Felders, in their ‘Species Lepidopterorum °
(1864), have separated the single species generally recognized into three, viz. (in addi-
tion to the type Merope) P. sulphureus of Palisot de Beauvois (figured by that author

* The various forms of the 9 Merope will be discussed further on.

+ P. Leonidas has the habit (rare in a Pupilio) of settling not unfrequently on the prc jjecting twig of some tree,
and there remaining motionless, with the wings closed and hanging downward, precisely after the manner of Danais
Echeria, for which species, in this position, I have, on more than one occasion, mistaken it.

+ Mr. Horace Waller has shown me a specimen taken at Mount Morambala, on the River Shire, a northern
tributary of the Zambesi.

4a 2

508 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

from a West-African specimen) and P. Meriones, n. sp., the Madagascarene form, or
“ Var. B,” of Boisduval’s ‘Spécies Général’ (p. 222). The former of the two Felderian
species appears to me of doubtful value, being founded upon some slight differences in
the outline of the wings (a very variable character in Merope) and in the width of the
discoidal cell of the hind wings*; but there is some ground for the separation of the
form inhabiting Madagascar, as will be seen immediately.

Considering for how long a time this fine Papilio has been known, and how frequently
it reaches England in collections from the various African settlements, it seems singular
that the female should be unknown. I have examined a large number of specimens in
the National Collection, the Oxford Museum, and the principal private collections, and
found them, without exception, to be males. In Southern Africa, I have been familiar
with the butterfly in its native woods for some years, have taken some scores of speci-
mens, and have examined many others from localities which I had no opportunity of
visiting : all were males.

But in various parts of the Cape, Kaffraria, and Natal there occurs, within a range
corresponding to that of P. Merope, the uncommon Papilio Cenea, of Stoll, already
mentioned as so singularly accurate a mimicker of Danais Echeria. I have reason to
believe that nearly all the specimens of this at present in collections have passed through
my hands; and among the whole number (about 25) from various localities, no male has
occurred. A black Papilio, without tailst, and marked with ochre-yellow patches and
spots, is a very different-looking butterfly from the fine pale-yellow Merope; yet there
is good reason for believing that Cenea is the female of Merope, modified in imitation of
Danais Echeria. A close inspection reveals many points in common between the two
insects. The head, thorax, and abdomen are similarly coloured and spotted. The pale
spot near the apex of the fore wings, so conspicuous in the dark border of Merope, is
almost always present (though occasionally smaller) in Cenea. On the under surface,
the ochre hue of the hind wings and border of the fore wings only differs in Cenea by
being rather darker; and the cellular and internervular streaks in the basal region, as
well as a pale space, between the radials, in the darker band beyond the middle of the
hind wings are found in both butterflies. Both in the Cape Colony and Natal, the woods
in which Merope was common were the only spots in which I met with Cenea; and on
one occasion I saw Cenea chased by Merope in the manner peculiar to a male butterfly
when in pursuit of the female. Apart from these circumstances, analogy with the case
of Papilio echerioides (the ¢ of which also copies Danais Echeria, while the 3 is
widely different) strongly supports the probability of the view here enunciated.

As Cenea occurs only in the south of Africa, its known range being from Knysna

* Dr. Felder does not use the name sulphureus to include the West-African form of Merope, expressly observing
(loc. cit. p. 77) that the agreement of Cramer’s figures, made from West-African examples, with the South-African
Merope, precludes him from considering sulphureus a geographical subspecies.

+ Cramer's earlier figures (pl. 151. figs. A, B) represent Merope without tails. This may be, as Boisduval suggests,
only the result of mutilation; but these processes are known to be a variable character in many species of Papilio,
and in Merope itself vary in size; so that the existence of a form in which they are wanting, or nearly so, need not

be thought impossible.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 509

(lat. 33° 59’ S., long. 23° 3’ E.) to Port Natal, the inquiry naturally arises, where is the
female of the tropical Merope? One might have been at a loss here, were it not for
another, and still rarer, isolated ¢ Papilio which is found in Southern Africa, viz. P.
Hippocoon, Fab. (=P. Westermanni, Boisd.), and which appears to be rather commoner
on the western coast. This butterfly closely resembles Danais Niavius, Linn., a common
species, which also inhabits tropical Africa and Natal. ippocoon is brought from the
same localities as Merope in Western Africa, but is very much scarcer in collections.
The markings already mentioned as common to Merope and Ceneu recur in Hippocoon ;
and some remarkable intermediate southern examples in my collection appear to indicate
some of the stages by which Hippocoon’s broad white markings may have been gradually
modified in the direction of the dominant southern Danais, D. Echeria, until so different
a form as Cenea was the result. In various collections, I have examined fourteen speci-
mens of Hippocoon, all of which are females.

A third remarkable Papilio must be placed as another form of the ? Merope, viz.
P. Dionysos, Doubl.*, a very rare insect inhabiting Western Africa. I had long
regarded this butterfly as probably connected with the curious series of forms under
consideration, even when I knew it only from the figure in the ‘Genera of Diurnal
Lepidoptera ;> and my opinion has been strengthened by an examination of the type
specimen in Mr. Hewitson’s collection t. That example, as well as another larger speci-
men recently received by Mr. Hewitson from Old Calabar, is a female. These two
specimens come nearer to P. Mippocoon than to any other butterfly ; but their colouring
is very singular, the larger white space of the fore wings extending the whole length of
the inner margin, and being scarcely separated from the subapical white bar by some
blackish scaling, while the hind wings are wholly warm yellow-ochreous, except the
white-spotted black hind-marginal bordering. On the underside, however, the dull ochre
margins of the wings, and the internervular streaks, are like those of Hippocoon; and
the apical spot in the fore wings, as well as the markings of the body, present no
difference. But the strongest proof of the most intimate affinity between the two forms
is afforded by a third female specimen, which accompanied that just mentioned from
Old Calabar; for in this example the characters of Hippocoon and Dionysos are
unmistakably blended, the markings of the fore wings being precisely those of the former
(except that the inner marginal white extends rather further into the wings), while the
yellow tint of the hind wings, though paler, is like that of the type Dionysos.

The fourth form of the ? Merope is that described and figured by Professor West-
wood ¢ under the name of Papilio Trophonius, and suggested by him as possibly the
female of P. Cenea§. In this form the markings quite agree in size and shape with

* Gen. Diurn. Lep. pl. 3. fig. 4.

+ I take this opportunity of recording my warmest acknowledgment of the courtesy and liberality with which Mr.
Hewitson has given me unlimited access to his fine collection. I am also indebted to Mr. Bates, Mr. Butler, Mr.
Salvin, Mr. Wallace, and Professor Westwood for similar kind assistance.

¢ Arcana Entomologica, i. p. 163, pl. 39. figs. 1, 2.

§ This view of the sexes was adopted by Doubleday, without query, in the ‘ Genera of Diurnal Lepidoptera ;” and
upon this authority I unfortunately, without investigation, perpetuated the error in Part I. of my ‘ Rhopalocera

510 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

those of the southern examples of Hippocoon; but the hind wings and the broad inner-
marginal space in the fore wings are coloured brick-red instead of white ; and the butterfly
thus becomes a very fair imitation of Danais Chrysippus*. This form of 2 is even
rarer than Hippocoon, the number of specimens that I have seen in collections being
seven only, including two that I had the good fortune to capture in the Cape Colony,
one at Knysna, and the other at Plettenberg Bay +. Professor Westwood (Joc. cit.) states
that Trophonius is a native of Guinea as well as Kaflraria.

We have thus, as it appears to me, a most remarkable case of polymorphism in the
female of Papilio Merope—three of the four forms being direct mimickers respectively
of three prevalent African species of Danais, while the fourth, differing from all the
others, yet closely related by an intermediate variety to one of them, is probably modified,
or in course of modification, in mimicry of some other protected butterfly, possibly not
a Danais ¢.

But another point remains for consideration. Papilio Merope (or its close ally) in
Madagascar, presents a female tailed and colowred like the male, and differing only in
the possession of a broad black bar on the costa of the fore wings, almost crossing the
discoidal cell.- That this is the rule in Madagascar cannot be doubted, as Mr. Plant’s
collection contained a series of females presenting little or no variation. The examina-
tion of a number of examples from the island in question leads me to think that the form
there prevalent is constant in both sexes, and entitled to rank as a distinct species §.
But whether we accord or refuse specific rank to P. Meriones matters little to those
who hold that unmistakably close alliance between two or more forms is at once the
result and evidence of community of descent. Place the males from the island side by
side with those from the continent of Africa, and perhaps few would be disposed to
regard the former as specifically distinct from the latter; and yet we find the female of

Africee Australis,’ published in 1861. Hopffer has recently (Stett. ent. Zeit. 1866, pp. 131-132) corrected the
mistake, pointing out that females of both forms are in the Berlin Museum.— Vide ‘ Zoological Record’ (1866), p. 451.

* A curious example, taken (in company with P. Merope) near St.-Lucia Bay in South-eastern Africa, by Col.
Tower, of the Coldstream Guards, is to some extent intermediate between the 7rophonius and Hippocoon forms, the
broad whitish spaces being obscured throughout with a dull-ochreous tint.

+ It is worthy of notice that on each occasion of my meeting with Trophonius, I took, in the same spot, a specimen
of the Cenea form of 9.

+ It may be objected that, in the strict sense of the term, this is not a true case of polymorphism, seeing that
intermediate varieties still occur which more or less connect the different forms. I am willing to admit that the
phenomenon is not yet absolutely complete; but the three forms that imitate the three species of Danais are already
so marked that the elimination of the few individuals of intermediate or unstable character, that serve to link to
some extent two or more of those forms, will probably be the work of no very extended period.

§ In the insular form, the black border of the fore wings forms much sharper projections inwardly on the nervules,
and the costal edging is brownish instead of black, in the g never extending below the subcostal nervure. The band
crossing the hind wings is always widely interrupted in two places, and the intermarginal black edging is wanting,
while the tails are all fuscous except the broad ochreous tip. On the wnderside, the ochreous colouring is rather
paler and more rufous in tint; and in the Q the hind wings are clouded from the base, over the disioidal cell, and the
inner-marginal region with brownish. The spots of the head and thorax are yellowish, and in the ¢ almost concealed
by a clothing of brownish hairs; and the abdomen is without the ordinary dark spots (save some very faint traces in
the ¢), being coloured almost uniformly of the same pale yellow as the upper surface of the wings.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 511

the insular race constant to one pattern, and not differing greatly from the male; while
the African female presents four distinct forms (besides certain intermediate varieties),
not one of which resembles the male. It would appear reasonable to argue from this
that Madagascar was the original starting-point of this type of Papilio, and that the
harder and more complex conditions of African life, causing a severer persecution, had
occasioned a necessity for the less active, and perhaps, as now, scarcer ¢ to assume the
protective colouring and outline of the surrounding Danaide. Yet the very wide
dispersion of this butterfly over the continent seems rather to indicate that the original
form of Merope was of African derivation, and at one time had extended to Madagascar,
possibly before that region became insulated, but that since that period, during slowly-
changing conditions of life, natural selection has induced the elimination in Africa of
all the pale, conspicuous females of the male coloration, only preserving those that
more or less resembled the protected Danaide,—while in Madagascar the female, in
the absence of any keenly persecuting agency, has retained the form and colour possessed
by the first immigrants from the continent. In the broad black costal bar of the fore
wings which distinguishes the female in Madagascar, regarded in relation to the hind-
marginal black border, it is not difficult to recognize the material upon-which natural
selection might gradually work, to the ultimate production of a Danaidiform butterfly
like Hippocoon or even Cenea; and it is remarkable that, in all the African forms of the
female, an oblique, narrow, whitish marking remains near the extremity of the discoidal
cell of the fore wings, in a position exactly corresponding to the outer border of the costal
bar, as if to record, with the other pale spots and markings, how the black of the margins
had gained upon the ground-colour as the process of increasing resemblance to Danais
was slowly wrought out.

Returning to the subject, from which this has been so lengthy a digression, it is worthy
of note that the mimicking Diadema above described seems only to occur at Natal, and
correctly copies the variety of Danais Echeria which is there prevalent, viz. that which
has all the spots of the fore wings white.

4. Danats Nravius, Linn. (Tab. XLII. fig. 6.)
Danais Niavius, Syst. Nat. ii. p. 766. no. 109 (1767) ; Cram. Pap. Exot. t. 2. figg. H, G.

This is an abundant butterfly in Tropical Western Africa; but the only special localities
that I have found recorded for it are Sierra Leone, Ashanti, and Angola. In the two
former of these districts occur two very accurate imitators of Niavius, viz. Diadema
Anthedon, Doubl., and the prevalent West-African form of the ¢ Papilio Merope (P.
Hippocoon, Fab.). The Papilio has also been received from Calabar.

It is to this striking case of mimicry that Boisduval refers in the passage which I have
quoted at the head of this paper. He mentions, it is true, Diadema dubia; but this is
owing to the confusion that has prevailed regarding the closely allied mimetic Diademe,
D. Anthedon being the species concerned, and being easily distinguished by the very
large inner-marginal white patch, and broad subapical bar of the fore wings. Cramer,
as Prof. Westwood has pointed out (Are. Ent. i. p. 152), figured the Papilio as the ¢
Niavius, in his Plate 284. fig. A; and Palisot de Beauvois subsequently did the same

512 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

(Ins. recueillis en Afrique, &c. t. vi. figs. la, 10). Danais Niavius is not confined to
West Africa, but also inhabits Natal, where it constantly presents broader white
markings, particularly in the hind wings*; and it is most interesting to find that both
the Diadema and the Papilio in that part of Africa vary in exactly the same manner
from the tropical type-form. Diadema Anthedon + has been taken at St. Lucia Bay by
Col. Tower, and is recorded from the Querimba Islands by Hopffer (in Peters’s ‘ Reise
nach Mossambique,’ p. 885); and there can therefore be little doubt of its occurrence
in company with Niavius in the intermediate Zambesi region.

I did not find this Danais commoner than the Diadema at Natal, during my visit in
the early part of 1867; but Mr. M‘Ken, the Superintendent of the Botanic Gardens at
D’Urban, informed me that it was more plentiful at another season of the year. So
close is the resemblance between this butterfly and Diadema Anthedon on the wing,
that I was never certain as to which butterfly I had captured, until close examination
had been made. The Hippocoon-form of the ¢ Papilio Merope seemed very rare at
Natal, one example only being taken, by my Kafir collector, near D’Urban. As far as
my present knowledge extends, however, this form of the female Papilio appears to range
further southward than either the Danais or the Diadema; for I have seen two examples
captured in Kaffraria proper by Mr. J. H. Bowker, and another taken near Grahams-
town by Mrs. Barber. The ¢ Papilio Merope, it should be observed, as well as the
Cenea-form of ?, occurs in both those localities, and as far to the south and west as the
Knysna River.

5. Danas Curysippus, Linn. (Tab. XLII. fig. 5.)

Danais Chrysippus, Syst. Nat. i. p. 767; Cram. Pap. Exot. t. 118. figg. B, C.
Var. A. Alcippus, Cram. op. cit. t. 127. figg. E, F.

Var. B. Dorippus, Klug, Symb. Phys. pl. 48. figs. 1-5.

This well-known species seems to inhabit the whole of Africa, ranges through southern
Asia from Syria to Hongkong, and is recorded from Java, Ceram, and Timor, in the
Malayan Archipelago. It even extends into Southern Europe (Greece and Turkey),
and, according to Godart, has been taken at Naples. It presents two varieties,—one
(Alcippus, Cr.) in which the disk of the hind wings is more or less suffused with white ;
the other (Dorippus, Klug) in which the black apex of the fore wings, and its oblique
white bar, are obliterated by the brick-red ground-colour }. Both these varieties seem
to be most frequent in Africa, Alcippus being known from four, and Dorippus from three
different parts of that continent. It is needless to dwell upon the very striking

* An example of this southern variety was brought from the Zambesi by the Rey. H. Rowley, and is now in the
Hope Museum at Oxford.

+ In my ‘ Rhopalocera Africe Australis,’ pt. ii. p. 838, I have given Anthedon as a synonym of dubia. This is an
error, into which I was led, in the absence of specimens of dubia, by Boisduval’s description (Faune Ent. de Mad. &e.
p- 40), and by his mention of dubia as having a special analogy with Danais Niavius.

= The type of Klug’s Dorippus, figured in ‘ Symbolwe Physics’ (Joe. cit. figs. 1-4), consists of examples of both sexes,
from New Dongola and Ambukhol, on the Nile in Lower Nubia, which present both the red suffused apex of the fore
wings and a broad white suffusion over the disk of the hind wings. The “variety” of the male, however (fig. 5),
wants the white in the hind wings, but has a dull fuscous shade over the basal half of both wings.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 513

resemblance borne to Danais Chrysippus by the female Diadema Bolina, Linn., as this
is one of the most generally known eases of mimicry in existence. The Diadema
accompanies the Danais throughout its range, with the single exception of the European
shore of the Mediterranean

and is even recorded from Australia and South America *,
regions in which Chrysippus does not occur. It is very seldom that one receives a
collection, however small, containing Chrysippus, in which Bolina is absent. I have been
careful to note all the recorded localities of specimens of both butterflies that have come
under my notice, and find that the two coexist in twenty-two different localities, in
addition to which there are seven instances of their occurrence in closely adjacent
districts +. Still more interesting is the fact that the 2 Diadema presents two varieties
exactly corresponding with the varicties of Chrysippws mentioned above, viz. one in
which the hind wings are more or less clouded with white, and another (Jnaria, Cram.)
that has the apical black and white of the fore wings replaced by the brick-red ground-
colour t. In each of these cases, I have not succeeded in finding more than one locality
where the variety of Chrysippus is known to be accompanied by the corresponding
variety of Bolina, viz. Sierra Leone, where the white-clouded form of both butterflies
occurs, and D’Urban, Port Natal, whence I have both Dorippus and Inaria; but
there seems little reason to doubt that they are to be found together in many other
places, when we consider how widely the varieties of both insects range.

In nature, the imitation of Chrysippus by the ¢ Bolina is singularly deceptive, as well
when the butterfly is settled on flowers as when it is on the wing; and it requires a keen
eye and close observation to distinguish one insect from the other. I am disposed to
imagine that the closeness of the mimicry even deceives the male Chrysippus; for, on
one occasion, at Port Natal, in a spot where the Danais was abundant, I was for some
time watching two females of Bolina that I had carefully marked, and was as mucli
surprised as interested to observe a Chrysippus pertinaciously chase one of the Boline
about the place. The female Diadema naturally gave Chrysippus no encouragement,
and, being more active on the wing, repeatedly evaded her pursuer$. Waiting to see
the close of this singular chase, I unfortunately lost sight of the Chrysippus among the
other specimens floating about; but I captured the ¢ Bolina, and have no doubt that
the pursuer was of the male sex.

I have already noticed the rare form of the female Papilio Merope (P. Trophonius,
Westw.), which mimicks D. Chrysippus. This imitation is not by any means as close

* Regarding the latter region, it seems to be doubtful whether D. Bolina has succeeded in naturalizing itself in
any of the localities mentioned by authors; but the insect certainly appears to have been taken in Guiana (Boisduval
and Doubleday), Surinam (Cramer and Boisduyal), Cayenne (Godart), and Para (Hopftter).

+ Eyen in the oceanic islands to which Chrysippus extends, such as St. Helena (Coll. Burchell), Bourbon, and
Mauritius, Bolina appears as its constant companion ; but I have not heard of the latter's occurrence in Teneriffe, where
Chrysippus has been taken.

+ Similarly to Klug’s type of Dorippus, the Inaria-form of the Q Bolina is sometimes found with a white suffusion
on the hind wings.

§ The ¢ Bolina, when pursuing the female, keeps a little below her, with his wings constantly and rapidly
quivering, while the female slowly rises, with little motion of the wings, towards the summit of some adjacent trec.
Mr. J. H. Bowker observed this in Kaffraria, in the year 1863; and I have since noticed he same in Natal.

VOL. XXVI. 4B

514 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

as that by Diadema Bolina, but it is sufficiently near to deceive the collector when the
butterfly is on the wing. Until they settled on flowers, when the tremulous motion of
the wings at once betrayed their disguise, I mistook both the examples of this Papilio
that I captured for Chrysippus.

In Western Africa (Ashanti) occurs a fine Nymphalide, Romaleosoma Eleus, Dru.,
the colouring and pattern of the female of which show a strong, but far from exact
resemblance to those of D. Chrysippus, the principal element of difference being the
possession by Eleus of a broad black band, containing conspicuous white spots, which
borders the hind wings. There is, however, in the British Museum, a variety of the
% Lleus, from Congo (in 6° 8. lat.), which more nearly approaches the aspect of Chry-
sippus. Compared with the type-form, it is smaller, and with more elongated fore wings ;
the ground-colour is redder and clearer; the apical black of the fore wings occupies a
smaller space, while its white bar is broader; and the border of the hind wings is nar-
rower, though still broad and conspicuous*. Congo is one of the known habitats of
Chrysippus.

6. AcrkmA GBra, Fab.

3 Papilio Gea, Fab. Sp. Ins. u. p. 32.

P. Epea, Cram. Pap. Exot. pl. 230. figs. B, C.

9? Papilio Iodutta, Fab. Ent. Syst. mi. 1. p. 175.
3,2. Acrea Gea, Godt. Enc. Méth. ix. p. 238.

I have no doubt that Godart rightly considered the Jodutta of Fabricius to be the
2 of that author’s Gea, as the difference in colour of the pale bands is the only distinction,
and there are several instances of allied Acree in which the fulvous or yellow markings
of the male are replaced by white in the female. Doubleday (‘Gen. Diurn. Lep.’
p- 141) gives the Limandra of Jones’s ‘Icones’ as the female of Gea, while Godart
refers it to 2. Huryta, Linn. The specimens named Timandra in the National Collection
agree well with Fabricius’s description of Jodutta, and are evidently females of Gea.

The male has been received from Ashanti, Calabar, and Congo; the female from Sierra
Leone and Calabar. Two other butterflies inhabiting both Ashanti and Calabar are
close mimickers of this Acrea, viz. Panopea Hirce, Dru., and the female Papilio Cynorta,
Fab. (=P. Boisduvallianus, Westw.)+. In the Panopea, the imitation is twofold, the
differing male and female of the Acrea being copied by the corresponding sexes of the
mimicker; but in the Papilio it is the female only that copies (very exactly) the female
Gea, the male being of a very different pattern as regards the fore wings. In addition to

* Danais Chrysippus is not without its mimickers among the eastern Mymphalide, the most remarkable of which
are the Javan Cethosia Penthesilea, Cr., and the female of Argynnis Niphe, Linn., inhabiting India and China.
The Cethosia differs widely from Chrysippus on the under surface, but the upperside is a very close copy, both in
pattern and colours, of that of the Danais, differing only in the possession of a submarginal row of spots in the hind
wings. The case of the 2 Argynnis is the more interesting from the fact of the sexes being so dissimilar, the male
being of the ordinary colouring of the genus.

+ I have elsewhere discussed the grounds for considering P. Boisduvallianus to be the female of Cynorta (see
Trans. Ent. Soc.).

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 515

the localities named, the Papilio has also been found at Sierra Leone and the Gaboon,
at the former of which places the Acrea occurs.

There is a difference in the outiine of the wings between the male and female of Aera@d
Gea, as in many other species of the same genus; and this discrepancy is reproduced in
Panopea Hirce, the female of which has the fore wings blunter and broader than those
of the male. So deceptive is the mimicry of the ¢ Gea, by the ¢ Hirce, that Godart
(loc. cit.) quotes Drury’s figure of the latter as a representation of the Acrea. In the
Linnean collection there is a specimen of the Panopea labelled “ Acrea Gea, Fab.” ;
and I found an example associated with a specimen of the same Acre@a in the Banksian
Collection at the British Museum. ‘The figure of “ Huryta” in Clerck’s ‘Icones’
(t. 31. f. 180), to which Linné refers in the twelfth edition of the ‘Systema Nature,’ is
evidently drawn from a female of the same Panopea*.

This species of Panopea further presents several varieties of the female, which agree with
no known examples of Acrea Gea, but, strangely enough, are very fair imitators of certain
varieties of an allied species, 4. Huryta, occurring in the localities (Calabar and Congo)
which they inhabitt. In the British Museum there is an interesting specimen of the
female Hirce, in which the bands, though paler, are coloured like those of the male.
This example only bears the label “ West Africa,’ and I am therefore unable to state
whether this apparently rare form of the female occurs in company with that which is
white-banded.

Most of the examples of Ielanitis Phegea, Fab., a member of the Hurytelide, are
mimickers of Acrea Euryta; but a female specimen, from Ashanti, in the National Col-
lection (which is, I believe, the type of IZ Bammakoo, Westw. Gen. D. Lep. pl. 68. f. 3)
bears a nearer resemblance to the female 4. Gea in the position of the subapical bar of
the fore wings, and in the extension of the white band of the hind wings over the inner
margin of the fore wings.

7. Acra#A Evuryta, Linn.
Acrea Euryta, Syst. Nat. ii. p. 757. n. 69; Cram. Pap. Exot. pl. 233. figs. A, B.

This is a most variable species in both sexes. Mr. Hewitson has recently (October,
1867) devoted two plates of his ‘Exotic Butterflies’ to the delineation of the principal
varieties. It is not unlikely that a knowledge of the stations and habits of these but-

* Mr. Butler, who kindly pointed this out to me, has suggested that the Punopea should stand as P, Eurytus, Clerck,
this name being older than Drury’s. But it seems clear that Clerck figured the insect in the belief that it was Linne’s
Euryta, which is an Acreea; and it thus appears to me desirable, especially with the view of avoiding confusion in
names, to retain the appellation of Hirce for the Panopea. Linne’s description, too, as Mr. Hewitson remarks (‘ Ex.
Butt.’ Oct. 1867), accords with the Acraa, notwithstanding that he quotes Clerck’s figure.

+ Mr. Hewitson, who has already, in his ‘ Exotic Butterflies’ (part for October, 1867) delineated the varieties of
Acrea Euryta, and in whose collection I saw these singular varieties of P. Hirce, is about to publish the latter also ;
and I therefore refrain from more particularly describing the imitations in question.

+ After examining Mr. Hewitson’s fine series of this butterfly, I am disposed to agree with him that it is at present
impossible to separate the numerous forms which he has figured, with the exception of the female shown in the
second plate, fig. 29, which appears to be a distinct species.

4 B2

516 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

terflies would enable us to distinguish several local races, if not distinct species; but,
with the scanty information at hand, the attempt to do so would prove of little value.

As already mentioned, certain varieties of this abundant Aerea (which is known to
inhabit Sierra Leone, Ashanti, Old Calabar, the Gaboon, and Congo) are the objects of
imitation by varieties of Panxopea Hirce, ¢, received from Congo and Old Calabar. The
particular varieties of the 4cr@a that are so imitated are known to inhabit the same
localities as the mimickers.

Another imitator is the scarce Eurytelide, Ielanitis Phegec, Fab. This butterfly,
like others of its family, is marked on the underside of the wings with numerous short
transverse lines; and it is interesting to observe how, at the base of the hind wings,
several of these lines are confluently grouped, in manifest imitation of the spots which
occupy the same position in the dAcre@a. The fulvous-marked examples of Jf, Phegea,
of which I have seen two, appear to be males; but the variety of Huryta which they
most closely resemble is a female, figured by Mr. Hewitson (/oc. ei/. f. 30), in which the
fulvous bar of the fore wings is rather narrow and with an ochreous tinge, and the inner
margin coloured with fulvous. ‘The white-banded females of the Melanitis copy the 9?
Acrea figured on the same plate (f. 31), which has the band of the fore wings rather
broader than usual, and that of the hind wings, with the inner margin of the fore wings,
slightly tinged with yellow. IZ Phegea has been brought from Old Calabar and Ashanti,
as well as from other West-African regions not specially recorded.

§. Acr#A AGANICE, Hewits. (Tab. XLII. fig. 2.)
Acrea Aganice, Hewits. Exot. Butt. i. pl. 6. £. 3; Trimen, Rhop. Afr. Austr. i. p. 109. n. 69.

This Acrea is closely related to A. Huryta, Linn., differing chiefly in the smaller
size and distinct coloration of the male, which has the pale bands yellowish, or yel-
lowish white, instead of fulvous. It is only known to occur in the South of Africa,
inhabiting Kaffraria proper and Natal, and is accompanied in the latter district by a
mimicking Panopea, which is nearly allied to P. Lucretia, Cram., and which I have
recently described, in the Transactions of the Entomological Society of London, as P.
Tarquinia. 1 mentioned (loc. cit.) the fact of this Nymphalide flying in the same woods
with Acrea Aganice, and have noted its rarity *, and how completely in general appearance
and habits it resembles its model. When, however, the insects are closely compared,
the mimicry is not so striking, as the Panopea possesses an additional small whitish bar
near the apex of the fore wings; but this is a subordinate feature, not noticeable when
the butterfly is on the wing.

9. Acrama Lycoa, Godt.
Acrea Lycoa, Godt. Ene. Meth. ix. p. 239. n. 27.

A range of some extent is recorded for this Acrea, viz. Sierra Leone, Ashanti, Calabar,

* Since the paper referred to was written, I have seen two other examples of Tarquinia in Mr. Hewitson’s col-
lection—one from Natal, and the other from the Zambesi. The former of these has much yellower bands than those
of the two that I met with in Natal, and evidently copies the yellower examples of Aganice.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 517

and Congo. The species may at once be known from its allies by the peculiar pattern of
the fore wings—the inferior pale marking running almost parallel with the subapical
bar, and nearly to the posterior angle, instead of forming an inner-marginal space
adjacent to the band crossing the hind wings. Panopea Lucretia, Cram., appears to
mimic this derea; but the resemblance is not so accurate as that between P. Turquinia
and 4. Aganice. Sierra Leone, the Gold Coast near Ashanti, and Calabar are the
known localities inhabited by P. Lueretia, which seems as rare as P. Targuinia, one
specimen in the British Museum, two in Mr. Hewitson’s collection, and two in the pos-
session of Mr. Swanzy being all the examples that I have seen.

10. Acr#a Zerss, Linn. (Tab. XLII. figs. 8, 9.)

Acrea Zetes, Syst. Nat. ii. p. 766. n. 110.

P. Menippe, Dru. Ilustr. N. H. iii. pl. 13. f. 8, 4; Stoll, Suppl. Cr. Pap. Ex. pl. 28. f. 1.
Var. Acrea Acara, Hewits. Exot. Butt. iii. pl. viii. f. 14, 15.

Acrea Caffra, Felder, Reise der Novara, ii. p. 369, pl. xlvi. f. 10, 11.

North of the Equator, the type-form of this species has an extensive range on the
western coast, but it does not appear to occur further to the south than Fernando Po.
Specimens in various collections have been received from the island in question, as well
as from Calabar, Ashanti, Cape Palmas, and Sierra Leone. In Southern Africa, the
species is represented by a well-marked variety, 4. Acara, Hewits.*, which differs, on
the upper surface, in having all the markings of the fore wings strongly defined (the red
ground-colour being wholly free from the almost universal fuscous suffusion so constant
in the type), and in possessing a conspicuous subapical ochreous bar. Both in the type-
form and in the southern variety, the colouring of the female is universally very much
duller and fainter than that of the male.

From Old Calabar and Ashanti a rare and handsome Nymphalide, Panopea Boisdu-
valii, Doubl. (Gen. D. Lep. pl. 37. f. 3, ¢), which closely imitates the type Acrea
Zetes, has been received. A male from the former district, in the collection of Mr.
Hewitson, and a female from the latter, in the British Museum, are the only West-
African specimens that I have seen; but these two examples respectively resemble in
their differences the dissimilar male and female of the Acrea, the female exhibiting an
incomplete subapical whitish ray, answering to that of the 9 Zeées.

In Natal Boisduvalit reappearst, in company with, and evidently mimicking (in the
red and black colouring of the fore wings and their ochreous subapical bar) the Acara
form of 4. Zetes ; and here, again, each sex of the Acre is copied by the corresponding
sex of the Panopea. <A singular example of the male Panopea, taken at Port Natal by

* Acrea Caffra of Felder. Mr. Hewitson notes (loc. cit.) that Acura is “perhaps only an extraordinary variety of
A. Menippe.”

+ [have examined eight Natalian specimens in collections, five males and three females. Six of these were taken
by Mr. M‘Ken, the Superintendent of the Botanic Gardens at D’Urban, who has kindly contributed specimens to the
South-African Museum and to my own collection, and who informed me that this beautiful insect is one of the
rarest of the native butterflies, but haunts the same spots as Acrwa Zetes. I saw but one individual (a female) on the

wing during my stay in Natal, but did not succeed in capturing it.

518 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

Mr. M‘Ken, and in the collection of the South-African Museum, is interesting from its
entirely wanting the ochreous subapical bar of the fore wings, all the apical region being
simply semitransparent fuscous grey (as in the tropical form), though the basal and
inner-marginal region has the red ground and black spotting as conspicuous as usual in
South-African specimens. This example thoroughly links the southern Boisduvalii
with the West-African type-form, and seems to indicate that the process of assimilation
to the southern form of the Aerea has not been fully completed.

Both sexes of the southern 4. Zetes, but especially the male, exhibit a tendency to
whitish suffusion about the discoidal cell and median nervules of the hind wings. Even
this slight variation is imitated by the Panopea; for the only example, a male, brought
from the Zambesi by the Rev. H. Rowley* has some faint whitish clouding in the same
part of the hind wings; and a female from Natal, in the South-African Museum, is
marked in the same manner.

It is not only on the upper surface that Botsduvalit successfully copies Zetes; the
underside is very effectually imitated, as well as such Acrveoid characters as the yellow
palpi and the spotting of the thorax and abdomen. FE ven in outline (the female Acrea
having less produced and blunter fore wings than the male) Boisduvalii is in both
sexes a faithful imitator.

11. Acrma Eeina, Cram.
Acrea Egina, Cram. Pap. Exot. pl. 39. f. F, Gt.
A. Zidora, Godt. Enc. Méth. ix. p. 237.

The principal characters distinguishing this species from its near ally, Ze¢es, Linn., are
as follows, viz.: in the fore wings the rufous marking near the posterior angle is much
enlarged and conspicuous; the innermost of the three black spots in the discoidal cell,
the customary ochreous marks along the hind margin, and the whitish-ochreous clouding
beyond the costal black bar (all marked characters of Zetes) are wanting, and the costal
bar itself is much narrower; while, in the hind wings, the black spots are larger, more
numerous, and grouped nearer to the base, and there are no red spots in the hind-
marginal border. These differences prevail in both sexes; but the ? is much duller and
obscurer than the d, and with faint markings.

Like A. Zetes, Egina inhabits Sierra Leone and Cape Palmas; but it also occurs in
Congo. From the latter region only there has been received Papilio Ridleyanus, White,
a very rare butterfly t, which, in pattern and coloration, stands alone in its great genus,
though apparently belonging to the Leonidas and Cyrnus group. The likeness which
this Papilio bears to A. Egina is very striking; and on the wing it must with difficulty

* This specimen is in the Hope Museum at Oxford, where Professor Westwood kindly pointed it out tome. I have
not found Acrea Zetes recorded as a native of the Zambesi country ; but the presence of P. Boisduvalii (of the southern
form) renders it highly probable that Zetes inhabits that region.

+ The Hyina of Stoll (Suppl. Cramer, t. 25. f. c, 3c) is distinet, being a smaller insect, with only a narrow reddish
line along the inner margin of the fore wings and a very narrow black hind-marginal edging to the hind wings.

+ I know of but three examples, viz. a male and female in the British Museum, and a male in Mr. Hewitson’s
collection.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 519

be distinguishable both from that Acrea and from A. Zetes. I have been doubtful
whether the Papilio better mimicked Zetes or Egina, but, on close comparison, believe
the latter to be the special object of imitation, in consequence of the red discal band in
the fore wings of Ridleyanus evidently copying the corresponding rufous space presented
by Hgina—though the red spots in the border of the hind wings resemble those of Zetes,
and of A. Perenna, Doubl., a native of Ashanti*. The female P. Ridleyanus resembles
the female Hgina, not only in its coloration being much duller than that of the male,
but (as in the case of Panopea Boisduvaliti and Acrea Zetes) in the blunter, more
rounded outline of the fore wingst.

The palpi of the Papilio are yellow, like those of the der@a; and the hinder portion
of the abdomen of the male is almost wholly yellowish beneath, and marked laterally
with spots of the same colour, increasing in size to the extremity, so as to approximate
to the colouring of the same part in the male Hgina, while the paler and more distinct
spots on the distended abdomen of the female present a nearer resemblance to the mark-
ings of the female Eyina.

I have now passed under review the most remarkable instances, eleven in number, of
mimetic analogies known to occur among the butterflies of Africa?. It has been shown,
(1°) that the mimicking butterflies invariably occur in districts inhabited by the species
mimicked, and in six cases (of southern species or varieties) are found in the very same
localities; (2°) that, in eight cases, the mimickers are known to be very much scarcer
than the species which they copy; (8°) that, in five cases, where the Danais or Acrea
presents local forms, or merely slight varieties, even these are imitated by individuals of
the mimicking species; (4°) that, in three cases, where the sexes of the insect mimicked
differ remarkably from each other, the sexes of the mimicker present corresponding dif-
ferences ; and (5°) that, in four cases observed by me in nature, it was next to impossible
to distinguish the living mimicker from the species which it imitated. In the instance
of Danais Echeria, I have demonstrated how, in addition to a Diadema, no less than
three species of Papilio copy that butterfly, the two closer mimickers in Papilio being
females of wholly dissimilar males, and belonging, in fact, to different sections of the
genus! In the very remarkable case of the polymorphic Papilio Merope, it has been
my endeavour to point out how three of the four forms of female in Africa are protected

* 4, Perenna also has the rufous space near the posterior angle of the fore wings, but it is a considerably smaller
insect than either Eyina or the Papilio, and has the fore wings remarkably narrow and elongated.

+ In the dull-fulvous suffusion of the hind wings, the 9 Ridleyanus more resembles some female examples ot
A. Zetes.

+ The vastness of the African regions that have never been entomologically explored renders it almost certain
that many other equally striking cases of mimicry remain to be discovered. No part of Equatorial or even Tropical
Africa has yet been searched by a competent lepidopterist—the collections transmitted to Europe from the western
coast and other parts having been formed by persons who collected at random, with little, if any, care to record special
localities or stations. Some idea of the Rhopalocerous riches of the region within a few degrees of the equator may
be formed from an inspection of the third yolume of Mr. Hewitson’s ‘ Exotic Butterflies,’ in which plate after plate is
filled with figures of new species from Old Calabar, the fruit of the labours of a single collector, who, but for a short

time and in a limited area, entomologized in that district.

520 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

by wearing the aspect, respectively, of three prevalent species of Danais; while the
Madagascarene Papilio Meriones, so closely allied to P. Merope as to be regarded by
most authors as merely a local race of the latter, presents but one form of female, not
ereatly differing from the male.

Such results as these, confirming and supplementing, as they do, the similar series of
facts observed by Mr. Bates in Equatorial America, as well as those tabulated by Mr.
Wallace in reference to India and the Malayan archipelago, are of the deepest signi-
ficance. Inexplicable as they must ever remain when regarded on the theory of the
independent creation of all organic beings precisely as we now behold them, they become
clearly intelligible when viewed as the natural consequences of the innate variability of
species, and the preservation and development by inheritance, through all time and under
all changes of surrounding conditions, of every successive variation advantageous to the
organisny originating it. In the infinitely complicated “struggle for life,” any advan-
tage, however slight, inevitably has its effect; and the individuals possessing it will not
only hold their ground to the exclusion of less fortunate competitors, but will transmit
the precious quality to some at least of their descendants. The process by which natural
selection gradually effects those innumerable protective resemblances of which these
mimetic analogies among butterflies form but a small portion, has been so ably traced by
distinguished naturalists, that it is unnecessary for me to attempt what could at best be
but a repetition of arguments already adduced; but I may be permitted, in conclusion,
to express my conviction of the harmonious relation in which the theory of the muta-
bility and gradual origin of species stands in regard to what is now universally admitted
respecting inorganic nature. No one disputes, at the present day, that the crust of the
earth has undergone profound changes both structurally and chemically; but who is
now found attempting to account for these mutations on the exploded theory of vast
convulsions of nature or general cataclysms? It is conceded on all hands that the
known forces and agencies at work under our eyes are sufficient, in the lapse of ages, to
account for all the past changes, however stupendous, of which the records remain ; but,
with a strange inconsistency, any suggestion of gradual change in the incomparably
more plastic and variable organic world is too often rejected with incredulity or disdain ;
and many men of science still cling to the theory of successive whosesale destructions
and as sudden creations to account for the extinct organisms revealed by paleontology.

Tt may, with confidence, be predicted that the day is not far distant when such dis-
cordant views of Nature will be unknown in the scientific world; for the progress of dis-
covery proclaims, with ever-increasing force, that the famous axiom Natura non facit
saltwm, is a truth of universal application, and that it is as impossible to sever the life of
the present from that of the remotest past as to interpose a barrier between the present
and the unknown future.

MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES. 521

EXPLANATION OF THE PLATES.

The Plates partially illustrate the most striking cases of Mimicry among South-African
Butterflies,—Plate I. consisting chiefly of the species of Danaide and Acreide that are
the objects of imitation, and Plate IT. of the imitating forms belonging to other families.

Puate I.

Fig. 1. Papilio Meriones, Felder, 2, from Mada-
gascar.

The black costal bar in the fore wings indicates
how the extreme disparity between the sexes of
the very closely allied Papilio Merope of the
African continent may have originated.

Fig. 2. Acrea Aganice, Hewitson, 3, from D’Ur-
ban, Natal.

This species is mimicked by the following, a
member of the Family Nymphalide, viz. :—
Fig. 4. Panopea Tarquinia, Trimen, 6, from

D’ Urban, Natal.
Fig. 3. Danais Echeria, Stoll, ¢, from Knysna,
Cape Colony.

The type-form, prevalent in the Cape Colony,
but scarcer to the eastward of it, and rare in
Natal.

Fig. 7. Danais Echeria, Stoll, var., 2 , from D’ Ur-
ban, Natal.
This variety prevails in Natal, almost to the
exclusion of the type-form.

Fig. 6. Danais Niavius, Linn., var., ¢, from
D’ Urban, Natal.

A Southern form, differing from the Tropical

West-African type in the much larger white
space of its hind wings.

VOL. XXVI.

Puate II.

Fig. 1. Papilio Merope, Cramer, ¢, from Knysna,
Cape Colony.

Fig. 3. Papilio Merope, Cramer, 1st form of 9,
from Knysna, Cape Colony. (P. Cenea,
Stoll.)

Mimics the type-form of Danais Echeria.

Fig. 4. Papilio Merope, Cramer, 1st form of 9°,
from D’Urban, Natal. (P. Cenea, Stoll.,
var.)

Mimics the Natalian variety of Danais Echerio.

Fig. 7. Diadema mima, Trimen, n. sp., ¢, from
D’ Urban, Natal.

Also mimics the prevalent variety of Danais
Echeria in Natal.

Fig. 2. Papilio Merope, Cramer, 9, variation,
from Tsomo River, Kaffraria.

Intermediate between the lst form of 2 (P.
Cenea, Stoll.) and the 2nd form (P. Hippocoon,
Fab.), and indicative of the yet unstable cha-
racter of the polymorphic 2 Merope.

Fig. 6. Papilio Merope, Cramer, 2nd form of ¢,
from Graham’s Town, Cape Colony.
(P. Hippocoon, Fab., var.)

Mimics the Southern form of Danais Niavius,
and, like the latter, differs from the West-African
Hippocoon in the much larger white space of its
hind wings.

4C

522 MR. TRIMEN ON MIMETIC ANALOGIES AMONG AFRICAN BUTTERFLIES.

Puate I.

Fig. 5. Danais Chrysippus, Linn., 9, from Cape
Town, Cape Colony.

Fig. 8. Acrea Zetes, Linn., var., 3, from Pieter-
maritzburg, Natal.

A Southern form, A. Acara of Hewitson,
differing from the Tropical West-African type
in the bright, unsuffused colouring of the upper
surface of the fore wings.

Fig. 9. Acrea Zetes, Linn., var., 2 ,from D’ Urban,
Natal.

The 2 of the Southern form, Acara, Hew-

itson.

Puate II.

Fig. 5. Papilio Merope, Cramer, 4th form of 9,
from Knysna, Cape Colony. (P. Tro-
phonius, Westwood.)

Mimies Danais Chrysippus, which is abun-
dant at Knysna and throughout the Colony.
Fig. 8. Panopea Boisduvalii, EB. Doubleday, var.,

3g, from D’Urban, Natal.

Mimics the ¢ of the Southern form of Acrea
Zetes, and, like that insect, differs from the
Tropical West- African type in the bright colour-
ing of the fore wings.

Fig. 9. Panopea Boisduvalii, K. Doubleday, var.,
2, from D’Urban, Natal.

Mimics the ° of the Southern form of Acrea
Zetes.

A.G Butler, del.et lith 1868

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