TT HOPE REPORTS |
VOEMTH |

1902

THE STRUGGLE FOR EXISTENCE
PROGECIIVE RESEMBLANCE
‘ MIMICRY, WARNING COLOURS
AND SEASONAL CHANGES IN
wr RIAN INSECTS

EDITED BY

EDWARD B. POULTON, M.A., D.Sc.
Hon VL DS PRINCETON, FUR:S:,.PiLtS.,) Fev. S508.G.S:

HUPE PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF OXFORD
FELLOW OF JESUS COLLEGE, OXFORD
PRESIDENT OF THE ENTOMOLOGICAL SOCIETY OF LONDON
CORRESPONDING MEMBER OF THE ACADEMY OF SCIENCE, NEW YORK, AND
THE SOCIETY OF NATURAL HISTORY, BOSTON

‘Semper Africa novi aliquid apportat’

OXFORD
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1902

THE STRUGGEE FOR EXISTENCE
PROTECTIVE RESEMBLANCE
MIMICRY, WARNING COLOURS
AND SEASONAL CHANGES IN

AFRICAN INSECTS

EDITED BY

EDWARD B. POULTON, M.A., D.Sc.
Hon. LE.D: PRINCETON, F.R.S., F.L.S., F.Z.S., F.G.S.

HOPE PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF OXFORD
FELLOW OF JESUS COLLEGE, OXFORD
PRESIDENT.OF THE ENTOMOLOGICAL SOCIETY OF LONDON
CORRESPONDING MEMBER OF THE ACADEMY OF SCIENCE, NEW YORK, AND
THE SOCIETY OF NATURAL HISTORY, BOSTON

‘Semper Africa novi aliquid apportat’

OXFORD
ERIN TED FOR PRIVATE CIRCULATION

BY HORACE HART, PRINTER TO THE UNIVERSITY

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IT is now ten years since the death of the veteran
entomologist—the most distinguished of all students of
Insect systematics—who was the first occupant of the
Hope Chair of Zoology. At the end of a period which
must form a. considerable fraction of the active life of
any man, it is appropriate that I should lay before the
University and those who desire to promote scientific
research in our land, a brief statement of the work which
has been done and of hopes for the future.

The two main lines of work which are suggested by
the presence of the Hope Collection in Oxford, were
clearly explained to the electors to the vacant chair by
the late Professor Huxley.

‘There are two courses open, he wrote, on Jan. 15,
1893, ‘for either of which there isa good deal to be
said. lt may be considered that the chief business of
the Professor is to be an efficient Curator of the great
collection of Insects under his charge, to sort it out
into so-called species, and catalogue them in such a
manner that the Collection’ may be made as available
and useful as possible to systematists and students of
Geographical Distribution. . . . The other course is to
elect a Professor, . . . who will use the Collection as
a means to the elucidation of the larger problems which
now press upon us. This is the alternative which would

4 PREFACE

commend itself to me if I had anything to do with the
appointment.’

A very natural inquiry may be raised as to why a
collection of /zsects, above all other animals, should be
so especially valuable for the elucidation of the larger
problems which deal, not only with the species of a single
group, but with every one of the innumerable and
infinitely varied forms, vegetable no less than animal,
in which life manifests itself. The answer is to be found
in the large number of offspring produced by each pair
of Insects, and the rapidity with which the generations
succeed each other, many cycles being completed in a
single year in warm countries; in the severity of the
struggle for life which prevents this remarkable rate of
multiplication from becoming the cause of any progressive
increase in the number of individuals; and finally in the
character of the struggle itself, which is precisely of that
highly specialized kind between the keen senses and
activities of enemies and the means of concealment or
other modes of defence of their Insect prey, which leads
by action and answering reaction to a progressive raising
of the standard in both pursuer and pursued. This is
why itis that Insects mean so much to the naturalist
or the philosopher who desires to look beneath the surface
for the forces which have moulded existing forms of life
out of earlier and very different forms. The wings of
butterflies, it has been said, ‘serve as a tablet on which

. H. W. Bates, quoted by A. R. Wallace in ‘ Natural Selection,’ London,
1875, p. 132. A more extended quotation is much to the point. H.W. Bates
was writing in ‘The Naturalist on the Amazons’ (London, pp. 347, 348 of
the 1879 edition), on the abundance and variety of the butterflies at Ega on
the Upper Amazons. (A few of the actual specimens captured by him in this

locality are in the Hope Collection.) ‘I paid especial attention to them,’ he
writes, ‘having found that this tribe was better adapted than almost any

PREFACE 5

Nature writes the story of the modifications of species.’
But the careful study of Insects tells us even more than
this; for it gives us the clearest insight we as yet possess
into the forces by which those modifications have been
brought about. Light is thrown upon the causes to which
organic evolution is due no less than upon the course
which organic evolution has pursued.

In issuing the third volume of Hope Reports it may be
claimed not unjustly that these insistent ‘larger problems’
have received during the past ten years that special atten-
tion which Huxley advocated. The subject which more
than any other has claimed the attention of the Hope

other group of animals or plants, to furnish facts in illustration of the
modifications which all species undergo in nature under changed local
conditions. This accidental superiority is owing partly to the simplicity
and distinctness of the specific characters ‘of the insects, and partly to the
facility with which very copious series of specimens can be collected and
placed side by side for comparison. The distinctness of the specific
characters is due probably to the fact that all the superficial signs of change
in the organization are exaggerated and made unusually plain by affecting
the framework, shape, and colour of the wings, which, as many anatomists
believe, are magnified extensions of the skin around the breathing orifices
of the thorax of the insects. ‘These expansions are clothed with minute
feathers or scales, coloured in regular patterns, which vary in accordance
with the slightest change in the conditions to which the species are exposed.
It may be said, therefore, that on these expanded membranes nature writes,
as on a tablet, the story of the modifications of species, so truly do all
changes of the organization register themselves thereon. Moreover, the
same colour-patterns of the wings generally show, with great regularity, the
degrees of blood-relationship of the species. As the laws of nature must
be the same for all beings, the conclusions furnished by this group of
insects must be applicable to the whole organic world; therefore the study
of butterflies—creatures selected as the type of airiness and frivolity—
instead of being despised, will some day be valued as one of the most
important branches of biological science.’ In this case the seer may be
said to have brought about the fulfilment of his own prophecy by the
inspiration given in his epoch-making theory of mimicry.

6 PREFACE

Department has been that very study of Warning Colora-
tion and Mimicry from which, as an American naturalist
has recently stated, ‘the theory of natural selection as
applied to Insects receives its strongest support'!.’ Indeed
the reference to Insects might safely be omitted; for the
facts brought to light in the pursuit of this study furnish
what is probably the most convincing of all evidence in
support of the Darwinian hypothesis.

In the meantime the systematic side of the Department
has occupied a large amount of time and care. It has
not been found that the two courses indicated by Professor
Huxley are by any means mutually exclusive. The
measure of success which has attended the attack on the
larger -—problems has been ‘due in great part tothe
existence in Oxford of a splendid general collection—
taking all the groups of Insects into account, the second
in the British Empire. |

Such success as the Hope Department has achieved
in these two directions has been due to many causes :—
to the energy and sympathy with which my Assistants,
Mr. W. Holland and Mr. A. H. Hamm, have entered into
the various researches which we have undertaken together ;
to the important investigations carried on by Dr. Dixey ;
to the recognition of Oxford as a centre where these
problems are studied. The result of this latter influence
has been the continual and ever-increasing accession of
material and observations from all parts of the world.
The inflow from Africa has been so large that the present
volume of Reports is entirely occupied with the problems
of Ethiopian zoology. This solid contribution to the

* A. G. Mayer, Science Bulletin of the Brooklyn Museum, Vol. I, No. 2,
Oct. 1902, p. 36.

PREFACE 7

natural history of the great continent which occupies so
dominant a position in our thoughts is mainly owing to
a great Rhodesian naturalist, Mr. Guy A. K. Marshall;
but a fruitful correspondence is also being carried on with
English naturalists in British East Africa, Uganda, British
Central Africa, Southern Nigeria and the Soudan. It is
a pleasure to reflect that years before we knew of the
great ideas for Oxford which were maturing in the mind
of Cecil Rhodes, the Hope Department was steadily making
itself recognized as a centre for the study of African
natural history. There is also great satisfaction in the
knowledge that so large a proportion of those who are
in touch with the Department are Oxford men, receiving
help and advice long after they have ceased to reside—
glad on their part to think that they are helping one of
the institutions of their University.

The amount of work done in ten years is undoubtedly
encouraging, but there is an aspect of it which is the
reverse. ‘The steady increase in the collections, due to the
work of the Department becoming more and more widely
known, has gradually occupied a larger and larger share of
my time and that of my Assistants, until finally we have
reached a point at which the accessions of each year can
barely be brought into a fit condition for cataloguing and
incorporation, while nothing is left for the vast mass of the
old collections, which imperatively demand a large amount
of attention. It is essential that the Department should
have further assistants with the mechanical skill necessary
for the manipulation of old and brittle specimens. I do
not wish to make too much of the demands upon my own
time and energy for work of a more or less mechanical
kind; but it is the fact that I wrote over 1,000 letters for
the Department in the course of 1902. A considerable

8 PREFACE

proportion of the correspondence would, I trust, always fall
to my share, but I hope that the time could be spent with
greater profit to the University if it were possible to obtain
some relief. I must also repeat what I have often said
before, that for the library to be of any real value the
catalogue must be completed; while for the safety of the
books a large sum must be spent on binding. The
University accepted over forty years ago one of the most
complete and valuable of then existing libraries which
dealt with the material of the Hope Collections. It is not
creditable that no steps have been taken to complete a
catalogue which is now so imperfect as to be useless, and
that an immense number of valuable monographs should be
endangered, and many should be injured, for want of binding.
And, as regards the Collections themselves, there is the
continual and pressing need for more accommodation in
safe and well-made cabinets, and the removal of all that
are unsafe and indeed extremely dangerous.

A great deal has been done during the past ten years.
In addition to the endowment of the Hope Chair, which
has now sunk to under £380, the grant. of £150 a year
paid by the University for salaries, the Hope Department
endowment of about £48 a year, and the Spilsbury en-
dowment of about £4 Ios., in addition to these yearly
payments a sum of nearly £1,850 has been spent on the
Department, a very solid help towards the making good
of some deficiencies mentioned above. Of this sum
rather over half has been contributed from various
University sources—from the Common University Fund
£515, from the Delegates of the Museum £266, from |
Convocation £150; and £800 has been spent on cabinets,
450 on book-binding, and the remainder on miscel-
laneous needs. A sum of £900, nearly half the whole

PREFACE 9

expenditure on these deficiencies, has been raised by friends
of the University and the Hope Department, and chiefly
by my own College, which has come to the assistance of
a hard-pressed University institution, electing me to a
Fellowship, with the full knowledge that the yearly income
would be applied to University purposes. Of this £900,
exactly three-quarters, £675, has been spent on cabinets,
4120 for special work upon the Collections, and the
remainder in adding to the Collections.

Much has been done, but increased activities have
brought increased needs. It is impossible, at any rate for
some time, to expect adequate help from the University.
I must, therefore, look for assistance to those who
sympathize with the aims of the Hope Department, and
value the work of which these volumes of Reports are
the evidence. |

EDWARD-B. POULTON:

HOPE DEPARTMENT OF ZOOLOGY,
UNIVERSITY MUSEUM, OXFORD.
february 9, 1903.

‘Semper Africa nove aliquid apportat.’

Notr.—In reference to the Latin quotation on the title-page and cover
of this volume I have adopted the form given by Erasmus in the Adagza
(ed. 15328 [Basle], p. 767). His note runs: ‘Huic simillimum est illud
Plinianum, quod in historia. mundi refert, Libyam semper aliquid novi
adferre. Quod quidem ideo dicebatur, quod in siticulosa regione ad unum
aliquem rivum plurimae ferarum species bibendi gratia convenire cogantur :
inibique varia mixtura violentiae Veneris, varias monstrorum formas, sub-
indeque novas nasci. Porro Plinius sumpsit ab Aristotele, apud quem
refertur libro de generatione animalium secundo capite quinto. Anaxilas
apud Athenaeum libro decimo quarto festiviter ad rem detorsit.’ The
preceding ‘adagium’ is Semper adfert Libya mali quippram.

_ There does not appear to be any authority for the form in which the

quotation usually appears—‘ Ex Africa semper aliquid novi.’ My friends,
Mr. C. E. Doble and Mr. W. G. Pogson Smith, have kindly helped me to
trace the quotation. 1 Se 2

CON PEN Fs OR VOL rt

1. Preface.

2. Notes on some cases of Seasonal Dimorphism in Butterflies, with an

account of Experiments by Mr. G. A. K. Marshall, F.Z.S. By
Frederick A. Dixey, M.A., M.D., F.E.S., Fellow of Wadham College,
Oxford. (From ‘Transactions of the Entomological Society of
London,’ 1902, p. 189.)

3. Five Years’ Observations and Experiments (1896-1901) on the

Lionomics of South African Insects, chiefly directed to the Investi-
gation of Mimicry and Warning Colours, by Guy A. K. Marshall,
F.Z.S. _ With a Discussion of the Results and Other Subjects
suggested by them, by Edward B. Poulton, M.A., D.Sc., F.R.S.,
Hope Professor of Zoology in the University of Oxford, Fellow of
Jesus College, Oxford; and an Appendix containing Descriptions
of New Species by Ernest E. Austen, William L. Distant, Colonel
Charles T. Bingham, F.Z.S., Guy A. K. Marshall, and Jules Bourgeois.
(From ‘ Transactions of the Entomological Society of London,’ 1902,

p. 287.)

4. Abstract of the above, by Edward B. Poulton, Hope Professor. (From

7

.‘ Proceedings of the Entomological Society of London,’ March 5, 1902.) -

. The Protective Resemblance to flowers borne by an African Homopterous

Insect, Ylata nigrocincta, Walker. By Sidney Langford Hinde.
Communicated by Professor Edward B, Poulton, M.A., D.Se., F.R.S.
(From ‘ Transactions of the Entomological Society of London,’ 1902,

p. 695.)

. Report of the Hope Professor of Zoology for rg00. (From the ‘ Oxford

University Gazette.’)

. Report of the Hope Professor of Zoology for rg01. (From the ‘ Oxford

University Gazette.’)

é 189° 5)

XI. Notes on some cases of Seasonal Dimorphism in
Butterflies, with an account of Experiments by
Mr. G. A. K. MarsHatt, F.Z.S. By FREDERICK
A. Drxey, M.A., M.D., F.ES., Fellow of Wadham
Colfege, Oxford.

[Read March 19th, 1902.]

PuaTe IV.

CONTENTS.
PAGE
1, Seasonal Dimorphism in Catopsilia pomona, Fabr.. . . 189
2. Seasonal Dimorphismn in Catopsilia pyranthe, Linn. . . 193

3. Irregularities of Seasonal Dimorphism in various Genera 194

4, Experiments and Observations in Seasonal Dimorphism
conducted by Mr. G. A. K. MarsHatt, F.Z.S., in the
Wears S96 OOM ee eae a cg ae eee oy 29D

DOM Peps ey tein ss Ey ap Boy ieee ae OIG

1, SEASONAL DiMoRPHISM IN Catopsilia pomona, Fabr.

I HAVE long been of opinion, from the examination
of many hundred specimens, that no line of specific
demarcation can be drawn between Cuatopsilia pomona,
Fabr., and C. crocale, Cram. This conclusion was based
mainly on the fact that, distinct in appearance as typical
examples of the two forms undoubtedly are, it is easy
to arrange a series of examples showing every possible
gradation between the two. The relation between C.
pomona and C. crocale so much resembles that between
forms which there is reason for regarding as cases of
seasonal dimorphism, that I was led to suspect that the
dimorphism of C. pomona-crocale might also have a
seasonal significance. In 1898 I mentioned my suspicion
to Mr. Trimen, showing to him at the same time a good
series, including many transitional forms, of C. pomona,
which had been captured near Brisbane in 1897 by T.
Batchelor, and presented to the Hope collection by Mr.
G. C. Griffiths. This series was noticed by Mr. Trimen in
his Presidential Address to the Entomological Society of
London, delivered on January 18, 1899, and was con-
sidered by him as “lending some probability to the view

TRANS. ENT. SOC. LOND. 1902.—PART II. (JUNE)

190 Dr. F. A. Dixey on

that C. crocale and C. pomona (including C. catilla) will
prove to be seasonal forms of one species.” * Direct
evidence on the point was, however, lacking; and 1]
therefore welcomed a statement made later by Batchelor
in a letter from Brisbane, and kindly communicated to
me by Professor Poulton, that C. erocale and C. pomona
were one species, “crocale being the summer brood and
pomona the autumn one.” It does not appear that any
observer has as yet actually bred one form from the other,
so that it cannot even now be said that their specific
identity is proved with absolute certainty. Nevertheless,
the opinion of a collector who has taken large numbers of
both forms is of weight,and may safely be held to indicate
a strong probability that, at all events in part of their
range, C. pomona, Fabr. and C. crocale, Cram. are seasonal
phases of the same species.

It is, however, evident that the case with regard to
C. pomona is not quite a simple one. In the autumn of
1900, a series of eighteen specimens of Catopsilia was
received by the Hope Professor at Oxford from the late
Mr. L. de Nicéville, who stated that they were all caught
nearly at the same time in the Kangra Valley, Western
Himalayas, by Mr. G. C. Dudgeon. Of these eighteen,
sixteen were taken on August 11, and the remain-
ing two on August 13, 1900. Two of the captures on
August 11 were Catopsilia pyranthe, Linn.; and of the
remainder, eight were C. crocale, Cram., and six were
C. pomona, Fabr. Those caught on August 13 were
C. crocale f and C. pomona 2 taken in copulé. In two
private letters to the Hope Professor, Mr. de Nicéville
appeals to this series of specimens in support of the
view that C. pomona+ and C. crocale constitute one

* Proc. Ent. Soc. Lond., 1898, p. Ixxvi. It is hardly necessary
to recall the fact that this address of Mr. Trimen’s contains an
excellent account of nearly all the experiments and observations
that had been made on the subject of seasonal dimorphism in
butterflies up to the time of its delivery.

+ De Nicéville calls it C. catilla, Cram.; but the latter name,
under which Cramer figures the form with brownish-crimson patches
on the under-surface (see Cramer, Pap. Exot., III. t. 229, D, E), is
later than that of Fabricius. Fabricius’s type still exists in the
Banksian cabinet, where I have examined it in concert with Dr. A.
G. Butler. The six specimens of C. pomona caught on August 11
include two ©. catilla, Cram. The British Museum contains six
specimens of C. crocale and seven of C. pomona caught by Mr.

Seasonal Dimorphism in Butterflies. 191

variable species, the variation not being due to seasonal
causes. This view was published by de Nicéville in
1894, and was reiterated by him on several subsequent
oceasions.* There can be no doubt that de Nicéville’s
opinion receives support from the present series of speci-
mens. The fact of C. crocale f pairing with C. pomona $
tends to show their specific identity, while the simul-
taneous occurrence of the two forms in presumably
equal numbers seems adverse to the supposition that the
dimorphism of this species has a seasonal significance.
With regard to the first point, that of specific identity,
I think there can now be no reasonable doubt that the case
is made out. I have already mentioned my own conviction
on the matter, which was arrived at independently, and
on different grounds. Batchelor’s observations here coin-
cide with de Nicéville’s; and it may be added that
Piepers,t who has bred the species in large numbers, is
strongly of the same opinion. On the other hand, Dr. L.
Martin, writing of the butterflies of Sumatra (Journ.
Asiat. Soc. of Bengal, LXIV, ii, p. 490, 1895), considers
C. crocale and C. catilla (pomona) distinct, on the following
grounds :—C. crocale, the far commoner form, occurs on
roads, near houses and gardens, and is never found in the
forest. C. catilla is found only in the forest. The antenne
of C. crocale are black in both sexes, those of C. catilla are
red. The underside of the males in @. crocale is unspotted,
and the tuft of hair on the inner margin is whitish. In
C. catilla the males, like the females, have reddish spots
on the underside of both wings, and the tuft of hair is

Dudgeon on the same occasion (August 12) as those mentioned
above. They are stated to have formed part of a migratory flight
which lasted all day.

* Gazetteer of Sikkim, p. 166, 1894; Journ. Asiat. Soc. Bengal,
LXIV, ii, p. 490, 1895 ; Journ. Bombay Nat. Hist. Soc., xi, p. 586,
1898; Journ. Asiat. Soc. Bengal, LX VIII, ii, p. 211, 1899. The
first two of these are cited by Mr. Trimen, loc. cit., p. 1xxvi, note.

+ “Die Farbenevolution bei den Pieriden,” Tijdschr. der Neder-
landsche Dierk. Vereenig.; (2) Deel V, p. 119, 1898. Piepers gives
thawrwma, Reak., as a synonym; the latter, however (from Mada-
gascar), is unquestionably distinct. ‘“ Pomona, Cram.” (tbid.) is a
slip; the name was bestowed by Fabricius. Piepers’s view was first
published in 1891—“ Observations sur des vols de Lépidoptéres”—
Natuurkundig Tijdschrift voor Ned.-Indié, Dl. L, 1891, pp. 205,
222. In the same periodical, Dl. LVII, 1898, he repeats it, but
speaks, rather curiously, of “ Gnoma, Feld.,” as a form of ‘ Catopsilia
pomona, Cram.” (loc. cit., p. 111).

192 Dr. F. A. Dixey on

distinctly yellow. The females of both forms are variable,
but the range of variation is distinct in the two. De
Nicéville, however, rightly remarks that “the distinctive
characters on which Dr. Martin relies are all quite incon-
stant, and entirely break down” when large numbers of
both forms are examined. It may be added that the
difference of habit alleged to exist between C. crocale and
C. pomona is no disproof of specific identity, imasmuch as
a similar difference, witnessed to by both Trimen* and
Marshall,+ obtains between Precis sesamus, Trim. and the
southern representation of P. ectavia, Cram. (called by
Marshall P. octavia-natalensis). The form natalensis,
according to Marshall, frequents high, open spots; sesamvus
is shade-loving, though it occasionally flies with natalensis,
especially at the change of seasons. Seswmus is more
wary than natalensis ; it is more often found in gardens,
and occasionally enters human habitations. It also con-
trasts with natalensis in being at times gregarious. But
in spite of these well-marked divergencies of habit, the
two forms, as is well known, have been absolutely proved
to be seasonal phases of the same species. Hence, in the
case of C. pomona and C. crocale, Dr. Martin’s objection
on the score of habit cannot be held any more conclusive
than that founded on the difference in aspect.

With regard to the second point, that of the seasonal
relations of the two forms, it seems that the utmost we
can at present allege is that in part, at all events, of its
range the dimorphism of C. pomona is associated with the
change of season. That this is not the case everywhere
is evident from de Nicéville’s observation, as quoted by
Trimen,t that “the innumerable varieties which are found
in both sexes occur at all times;” and, more particularly,
from the statement that “both true C. crocale and the
dimorphic form, C. catilla, Cram. occur commonly in
Mussoorie from July to October, and in Dehra Dun
throughout the warmer months of the year.Ӥ On the
other hand, we have Batchelor’s categorical assertion from

* South-African Butterflies, vol. I, 1887, pp. 230, 233.

+ Ann. Mag. Nat. Hist., 1898, vol. II, pp. 33, 34.

¢ Proc. Ent. Soc. Lond., 1898, p. Ixxvi.

§ Mackinnon and de Nicéville, Journal of Bombay Nat. Hist.
Soc., vol. XI, 1898, p. 586. Piepers also denies absolutely that the
dimorphism of C. pomona is seasonal (“‘ Notes from the Leyden
Museum,” vol. XXII, 1899, note 1, p. 13, ibique cit.).

Seasonal Dimorphism in Butterflies. 193

Brisbane, given above; while the fact recorded by Dr.
Martin (/oc. cit.) that among many hundreds of both sexes
of C. crocale, all presumably belonging to one emergence,
taken by him near Bindjei, there was not a single C.
catilla (pomona), may possibly have a similar significance.*

It is not a little remarkable that although there are
forty-three specimens of C. pomona and C. crocale in the
Hope collection duly labelled with locality and date, they
cannot be said to throw much light on the question of
seasonal dimorphism. What is wanted is a long series of
observations carefully carried on in one locality, and
accompanied, if possible, by breeding experiments.

If, as is probable, it should eventually be shown beyond
doubt that the different forms of C. pomona, though
related to the seasons in some part of its range, occur
indifferently at all times in others, the case would by no
means stand alone. I propose in the next place to notice
very briefly several statements that have been made by
different authorities with regard to other species, which
statements tend to show that in many cases where the
existence of seasonal modification has been reasonably
presumed, or even actually demonstrated, the seasonal
relation is far from being rigidly fixed in all parts of the
area of distribution.

2. SEASONAL DIMORPHISM IN Catopsilia pyranthe, Linn,

The first instance that may be taken is that of Catopsilia
pyranthe, Linn. This butterfly grades imperceptibly into
C. gnoma, Fabr. just as C. erocale does into U. pomona.
Here again, in the absence of breeding experiments, the
absolute proof of specific identity is still lacking; but de
Nicéville had no doubt, from his own observations, that the
two forms represent a single species. In this case he is
able to assign a seasonal value to the two forms,—C.
pyvanthe being in his opinion the wet-season, and C. gnoma
the dry-season phase of the species. But the point of
special interest, in view of the irregularity that appears to
obtain in the seasonal relations of C. crocale and C. pomona,

* It should, however, be noted that “N.-E. Sumatra does not
possess a well-marked dry and wet season, such as is found over
most of the continent of India, there being no month in the year
when it does uot rain.” Journ. Asiat. Soc. Bengal, LXIV, 1895,
pt: li, p. 362. See below, p. 196. ,

194 Dr. F, A. Dixey on

is the fact that, as recorded by de Nicéville himself, the
different forms of C. pyranthe, though corresponding to the
seasons in some parts of its range, are independent of them
in others. Thus, in speaking of this species under the
name of C. chryseis, Drury, he notes that “it is not seasonally
dimorphic in Sumatra as it is in India.” * Again, he
remarks under C. pyranthe, “Moore in the ‘ Lepidoptera
of Ceylon’ gives four forms of this species as separate
species; C. gnoma, Fabr., C. tlea, Fabr., C. chryseis, Drury,
as well as typical C. pyranthe. Manders notes that as far
as his observations go these four forms are not dependent
on season, but appear indiscriminately nearly throughout
the year, those flying in the dry season from February to
April being a little smaller than those found during the
rest of the year.” + On the other hand he says, “True
C. pyranthe is not very common in Mussoorie in the rains;
the dry-season form, C. gnoma, Fabricius, even less so.
In the Dun both forms are common in their respective
seasons.” ¢

If then we are to trust the observations that have been
cited, we are led to the conclusion that in these Catopsilias,
viz., U. pomona and C. pyranthe, we have to deal with two
polymorphic species, each of which has no doubt several
geographical forms, and each of which shows, in most
localities, a special tendency to cleavage into two well-
contrasted types. These latter phases in each case are in
some parts of the range of the species dependent on
seasonal changes; in other parts, however, they show no
such connection.

We may now pass on to the consideration of similar
irregularities as shown in other groups.

3. IRREGULARITIES OF SEASONAL DIMORPHISM IN
VARIOUS GENERA.

It has been recorded by most of those who have ex-
perimented on the subject, that there are individual
differences in the reaction of members of the same brood
to what appear to be identical conditions of the environ-
ment. A conspicuous instance of this is the well-known

* Journ. Asiat. Soc. Beng., LXIV, 1895, ii, p. 490.
+ Ibid., LX VIII, 1899, i, p. 211.
t Journ. Bombay Nat. Hist. Soc., XI, 1898, p. 586.

Seasonal Dimorphism im Butterflies, 195

experience of Mr. Marshall, who in April 1898 bred a
specimen of Precis sesamus and another of P. octavia-nata-
lensis from two eggs, laid on the same day by the same
mother, and reared under precisely similar conditions.*
Dr. Butler has also put it on record that Captain Nurse
bred Teracolus yerburw, Swinh., and 7. nowna, Luc.,+ from
a batch of similar larvee, the perfect insects presumably
emerging at the same season. Many cases have been
observed where, although each of the two forms of a
species is on the whole confined to its own time of year,
there is yet a considerable amount of overlapping at
the change of seasons; this overlapping showing itself
both by the simultaneous occurrence of freshly-emerged
specimens of both phases, and also by the appearance of a
more or less complete series of “intermediates.” A good
instance of the simultaneous occurrence, in the field, of
ditferent phases believed on strong grounds to be seasonal,
is afforded by the capture of all three forms (“ wet,’ “dry,”
and “intermediate ’’) of Precis sesamus by Mr. Crawshay at
Nairobi within little more than a week during the month
of April.{ Many records of this kind are in existence ;
and are often, no doubt, to be ranked as examples of the
seasonal overlapping that has just been mentioned.

It is however evident that there are numerous cases of
simultaneous occurrence which cannot be brought under
this head. Besides the definite statements of de Nicéville
with regard to two species of Catopsilia, we have now a
considerable bulk of evidence, with regard to many species,
of the appearance side-by-side, at all times of year, of
forms closely analogous with what are now well established
as seasonal phases. Thus, again according to de Nicé-
ville, the ocellated and non-ocellated forms of MJelanitis
leda, Linn., which he has shown to be related in India to
the wet and dry seasons respectively, both occur in North-
East Sumatra all the year round. In Java it has been

* Ann. Mag. Nat. Hist., 1898, vol. ii, p. 30.

+ More accurately, perbaps, 7. evagore, Klug. T. nowne is the
dry-season phase of the African form 7. daira, Klug. Capt. Nurse’s
larvee were found at Shaik Othman, and no doubt belonged to the
Arabian form, of which 7. yerburii, Swinh., is the wet, and 7.
evagore, Klug, the dry-season phase. This is pointed out by Butler
in Ann. Mag. Nat. Hist., 1897, vol. ii, p. 460. The original record
is in Proc. Zool. Soc., 1896, p. 247.

t Proc. Zool. Soc., 1900, p. 916.

TRANS. ENT. SOC. LOND. 1902.—PART HI. (JUNE) 14

196 Dr. F. A. Dixey on

noted by Piepers * that the non-ocellated form, though on
the whole belonging chiefly to the dry season, is also to be
met with during the rains. It is true, as Piepers says, that
in Java, as in the Malayan Islands generally, the distinc-
tion between dry and wet season is not so sharp as on the
Indian mainland; so that a certain amount of inter-
mingling of the two forms might perhaps have been
antecedently expected. It does not appear, however, that
all dimorphic species are affected by these or the lke
conditions in the same way. De Nicéville points out, in a
passage quoted by Trimen, that with this exception of
Melanitis leda there are no dry-season forms in North-east
Sumatra; and Doherty mentions analogous facts in refer-
ence to localities with a generally moist climate, like
Ceylon and Singapore, and also, mutatis mutandis, to dry
countries like Sind.t The prevalence of wet-season
forms in the equatorial forest region of West Africa is
another phenomenon of the same kind. Instances such
as these show that a generally damp country may be
characterized by a greater abundance of “ wet-season ”
forms, and vice versd. But these cases of the prevalence
of “dry” or “ wet-season ” forms respectively, according to
the general climatic conditions of a given locality, are, as
we have just seen, accompanied by others which seem to
prove that in certain districts, especially perhaps dry ones,
the phases that are usually associated with the seasons
occur indiscriminately at all times of the year.

Many such instances are recapitulated by Butler in his
late revisions of the genera 7'eracolus and Terias. Teracolus
eupompe, Klug, for example, has a wet, an intermediate
and a dry phase. “The two latter undoubtedly fly
together, and in Aden it is tolerably certain that all the
phases emerge at the same time as mere variations.” ¢
With regard to 7. halimede, Klug, Butler observes, “ 7.
acaste represents the wet-season phase, 7. halimede the

* “Die Farbenevolution bei den Pieriden,” Tijdschr. der Nederl.
Dierk. Vereenig; (2) Deel V, 1898, pp. 179—185, etc. The value of
the theoretical considerations based by Piepers on the facts that he
has evidently observed with much care, appears to me to be greatly
diminished by his refusal to admit the influence of selective
adaptation, even as a provisional hypothesis.

+ Proc. Ent. Soc. London, 1898, p. Ixvili. Compare Watson ;
Journ. Bombay Nat. Hist. Soc., 1894, vol. viii, p. 489, ete.

~ Ann. Mag. Nat. Hist., 1897, vol. ii, p. 497.

Seasonal Dimorphism in Butterflies. 197

intermediate, and 7’. cw/estis the dry-season phase of the
species; but they are none of them confined to seasons,
but occur (as is the case with other species in very arid
countries) as mere coexistent variations.” * Of 7. proto-
media, Klug, he remarks, “ At Aden all three [seasonal]
types occur together as mere variations.” + Other species
of Teracolus of which similar statements are made are 7’,
protractus, Butl., 7. phisadia, Godt., T. pucllaris, Butl., 7.
vestalis, Butl., 7. evagore, Klug, and 7. pleione, Klug.
With regard to Zerias Butler also notes that, “as in
Teracolus, those countries which have no wet season never-
theless produce the three phases of a species as coexistent
varieties.” | There is reason to think that in the New
World, at any rate, there may occur a similar intermingling
of forms which is not confined to “ countries having no wet
season.” Thus, Messrs. Godman and Salvin write as
follows: ‘“ Many of these forms [of Z’evias] are said to be
due to the season of the year at which they appear, wet-
season and dry-season broods having each their peculiar
characteristics. These observations have been made chiefly
in the east. In our country we have not noticed any
phenomenon of this kind.” § Mr. G. C. Champion again,
if my memory does not deceive me, in the discussion ‘that
followed the exhibition of certain specimens of Callidryas
referred to by Mr. Trimen (doc. cit.), many of which were
collected by himself, stated that according to his experience
of these butterflies, the varying forms of the same species
from the same locality had no definite relation to the
seasons. Colonel Swinhoe, besides recording the fact that
he has taken all the seasonal forms of certain eastern
Teracoli flying simultaneously at Karachi, has also averred
that he has captured Byblia simplex, Butl., the supposed
dry-season form in India of B. ilithyia, Drury, practically
all the year round. Some doubt has been thrown by de
Nicéville and by Marshall on the latter observation ; the

* Ann. Mag. Nat. Hist., 1897, vol. ii, p. 502. The dates of Col.
Yerbury’s captures at Aden clearly prove the simultaneous occurrence
of different “seasonal” forms, but they do not seem incompatible
with a certain amount of correspondence on the part of these phases
with the time of year. See, e.g., the dates given for Teracolus
celestis and T. acaste; Proc. Zool. Soc., 1884, pp. 489, 490.

7 Ann. Mag. Nat. Hist., 1897, vol. ii, p. 507.

t Ibid., 1898, vol. i, p. 57.

§ Biologia Centrali-Americana. Rhopal. ii, p. 154.

198 Dr. F. A. Dixey on

former, however, says Butler, is a fact that can be proved
from the data on the Museum specimens.*

Statements of this kind, the list of which could be largely
extended, go far to show that the case of Catopsilia pomona
and C. crocale is by no means an isolated one, and that just
as there are regions in which more than one geographical
form of a widely-ranging species may be found flying
together,t so there are districts of a greater or smaller
extent where diverse forms of a species, confined for part of
its range to definite seasons, may all occur simultaneously.
No doubt the data are as yet insufficient for a complete
explanation of these phenomena. It seems, however, clear
that the forms or phases which are usually called “seasonal”
may occur under many diverse conditions and in many
different proportions. It appears further that they do
not fall mto a regular system of succession, except in the
presence of regular alternations of season, and not always
then. I still venture to think that a probable view con-
cerning many of them is that briefly expressed by me
some years ago in “ Nature” (Vol. lx ; 1899, p. 98), viz.,
that polymorphism, however it may have arisen, is capable
of being brought more or less into relation with locality
and season under the influence of natural selection. On
the other hand, it 1s conceivable that in some cases at all
events the forms in question may have first arisen as
adaptations to the seasonal changes, and afterwards, in
consequence of extending their range, or of some other
alteration of conditions, may have partly or entirely lost

* Ann. Mag. Nat. Hist., 1897, 11, p. 386 ; [bid., 1896, 11, p. 335. The
following instances may be added from specimens with data in the
Hope collection :—(1) Australian form of Terias hecabe, Linn. (T.
sulphurata, Butl.) ; the dry, wet, and intermediate seasonal forms, all
taken by Mr. J. J. Walker, R.N., on June 19, 1890, at Port Darwin,
North Australia. (2) Teracolus phlegyas, Butl. (1. difficilis, E. M.
Sharpe) ; a wet-season male taken paired with a dry-season female,
both in good order, by Mr. G. A. K. Marshall, May 3, 1899, at
Salisbury, Mashonaland. (3) Teracolus vestalis, Butl. ; the wet and
dry-season forms both taken at Karachi on May 10, 1888, by Mr.
W. D. Cumming. (4) Belenois severina, Cram. ; wet and dry-season
forms both taken on Feb. 13, 1897, at Karkloof, Natal ; a wet-season
male paired with a wet-season female, and another wet-season male
with a dry-season female on Feb. 24, 1897, at Malvern, Natal. All
these by Mr. G. A. K. Marshall.

} E.g. the various forms of L. chrysippus, Linn., which are all
found together at Aden. See Butler in Proce. Zool. Soc., 1884, pp.
478-481 ; and Col. Yerbury in Journ. Bombay Nat. Hist. Soc., 1892,
p. 209.

Seasonal Dimorphism in Butterflies. 199

their correspondence therewith. These are questions that
must, I think, for the present remain unanswered ; though
whatever the solution may be, there seems no need to
anticipate that it will weaken the case for selective
adaptation.

4. EXPERIMENTS AND OBSERVATIONS IN SEASONAL DI-
MORPHISM CONDUCTED BY Mr. G. A. K. MARSHALL,
F.Z.8., IN THE YEARS 1896—1901.

In ae “Annals and Magazine of Natural History,”
1901, uu, p. 403, Mr. Marshall writes as follows:—“ Two
years ago I made a few experiments in applying moist
heat to the pup of several species of TZeracolus. Un-
fortunately all my notes on the subject have been lost,
but, so far as I can recollect, the results were almost
entirely negative, which I then attributed to insufficient
heat. The resulting specimens were, however, sent to
the Oxford University Museum with full data.” There
are also in the Hope collection several other specimens,
collected by Mr. Marshall in 1896 and following years,
which are of considerable interest in their bearing on the
subject of Seasonal Dimorphism. By the kindness of the
Hope Professor, I am permitted to give Mr. Marshall’s own
comments on both series of specimens. These are contained
in private letters to Professor Poulton, and have not hitherto
been published. I propose to arrange the notes in chrono-
logical sequence; but it will be seen that the experiments
fall into two main groups, which are more or less inter-
mingled in order of time. The first group of experi-
ments includes cases where one form of a species was
reared under normal conditions from eggs laid by another
form of the same species. In the second group of experi-
ments, the pup, or sometimes the larvz in their later
stages as well as the pups, were subjected to artificial
conditions in order to see whether any effect could be
thereby produced on the following emergence. It is well
known that very striking results have been brought about
by artificial conditions of temperature in the case of
dimorphic butterflies in Europe and North America. The
names of Dorfmeister, W. H. Edwards, Weismann, Merri-
field and Standfuss, to say nothing of other s, will occur to
every one as those of the authorities to whom we owe nearly

200 Dr. F, A. Dixey on

the whole of our knowledge in this particular. In view of
the great difference between the temperate and tropical
seasons, it was natural to suppose that the seasonal forms
of tropical butterflies would be found to stand im relation
to quite other meteorological conditions than those re-
sponded to by the Nearctic and Palearctic species which
had previously formed the subject of experiment. So far
as I am aware, the only factor found generally operative
in these latter cases is a raising or lowering of the
temperature; the direct effect of humidity has been tried,
but almost always with negative results. Mr. Marshall,
on the other hand, has successfully used heat in combi-
nation with both moisture and dryness, and has also
employed moisture unaccompanied by heat. By all these
means, as will be seen, he has secured results analogous
indeed with those of the European observers, but as a
rule far less complete. It is possible that there may still
be discovered some factor or combination of factors which
will produce, in dimorphic tropical species, equally strik-
ing results with those to which Merrifield and Standfuss
have now accustomed us. Most, however, of the species
so farinvestigated by Mr. Marshall have proved compara-
tively resistent to this kind of treatment, and he has no
instance of artificial modification which can be ranked
with the Araschnia levana of many experimenters, or the
Selenia tetralunaria of Mr. Merrifield.

Mr. Marshall’s initials are here appended to each separate
extract from his correspondence.

“ Estcourt, Natal ; Dec. 14, 1896.—I only succeeded in
getting three eggs of Teracolus topha,* of which I send
you one of the resulting specimens, which is undoubtedly
T. auxo, being of the early wet-season form with the upper
side black markings not yet fully developed. The eggs were
laid within five minutes of one another, and they hatched
simultaneously, but one larva pupated a day later than the
other two and emerged a day later. The first two examples

* The result of this experiment was communicated by Mr. Marshall
to the “ Entomologist’s Monthly Magazine,” 1897, p. 52, and is
referred to by Mr. ‘l'rimen in his address above quoted (Proc. Ent.
Soc. Lond., 1898, p. Ixxii). It should be noted that the name
T. topha, Wallgrn., which is now used by both Mr. Marshall and
Mr. 'Trimen to designate the dry-season form of 7. auxo, is con-
sidered by Dr. Butler to be applicable rather to an intermediate
form between 7. awxo and T. keiskamma, Trim., the latter being the
true dry-season phase. (Ann. Mag. Nat. Hist., 1897, ii, p. 453.)

Seasonal Dimorphism in Butterflies. 201

(of which yours is one) are quite similar, but the third has
the black edging to the apical patch of the forewing a trifle
heavier, and also shows a trace of the black line along the
inner edge of the patch characteristic of the full summer
form. As the eggs were all laid by the same female, and
the larvee were reared under absolutely similar conditions,
it would seem at first sight that the heavier markings
could only be due to the longer larval stage, but this seems
highly improbable. I was astonished at the rapid develop-
ment of this species; egg-stage, three days; larval stage,
twelve to thirteen days; pupal stage, eight days. Total,
twenty-three to twenty-four days. From this I should
estimate that there must be from nine to ten broods in
the year.”—G, A. K. M.

The above-mentioned specimen, a male, is now in the
Hope collection, and entirely bears out Mr. Marshall’s
description. It is a well-marked, but not extreme example
of the “wet-season” form 7. auxo, Luc. Mr. J. Mansel
Weale’s experience of the same species is well known ; *
and it may be noted that of five bred examples sent to the
Hope collection by Mr. Weale in 1878, there isa pair each
of the auxo (wet) and topha (dry) form, together with a
single female of an intermediate phase. Mr. Marshall’s
experiment removes the subject of the specific identity of
these several forms from the region of probable conjecture
to that of proof.

“ Estcourt, Dec. 14, 1896.—While staying with Mr. Burn,
at the junction of the Blaauwkraantz River with the Tugela,
I tried to see whether the black markings of the early wet
brood of Zeracolus anne could be intensified by damp
surroundings, so as to resemble those of the full wet form.
For this purpose I had a tin half filled with wet sand,
in which I stuck the pupz on thin sticks, covering it
over with a cloth on which was a wet sponge. Into this
I put five freshly-turned pupz, of which I kept three in
for seven days and two for nine. Only one specimen
emerged out of each lot, and so far as I can see there
is absolutely nothing unusual about either of them.
Although the results of the experiment are negative, they
are interesting, in that they éend to show that cold moisture
cannot accentuate the black markings of the wet-season
form, and also that cooler surroundings (induced by evapor-

* Trans. Ent. Soc. Lond., 1877, p. 273. See also Mr. Barker’s
comments ; Ibid., 1895, p. 422. '

202 Dr. F. A. Dixey on

ation) do not tend to cause a reversion to the dry-season
form. The first lot of 7. annx I bred (under ordinary
conditions) were in pupa during fine warm weather, and
took nine days to emerge. Those placed in the damp tin
took in both cases twelve days. Three other pupe kept
under ordinary conditions were also twelve days in pupa,
the last six days being cold, wet weather; these however
were all of the full wet form, one female being even blacker
than usual. With this species I observed that the bred
specimens were nearly always more advanced in coloration
than freshly emerged captured specimens.”’—G. A. K. M.

Eight of the specimens of 7. annx, Wallgrn. above
referred to, are now in the Hope collection. One of these
emerged on Nov. 17, 1896, after a pupal stage of twelve
days, “during seven of which it was kept in the damp tin
jar, as above stated. It is an ordinary wet-season male,
not extreme in character. A well-marked wet-season
female, also in pupa twelve days, but under usual condi-
tions, emerged on Nov. 11. This may be the female
mentioned above. The only other bred specimen is a
well-marked wet-season male, decidedly darker than the
first. It emerged on Nov. 18, but there is no note as
to its duration in the pupal state. The remaining five
specimens were caught in the open. A female taken
on Nov. 6 is wet-season ; a pair on Noy. 12 are intermedi-
ate, as are two males alken on Nov. 14 and Nov. 16
respectively.

“ Estcourt, Dec. 14, 1896.—On my return here I attempted
a small test experiment as a converse of the former one,
viz., Submitting pupze to dry warm conditions. My modus
oper andi was as follows: on’a tripod stand I placed a round
tin contaiming a little water; on the mouth of the tin was
a china saucer filled with dried sand, in which were placed
the pupz beneath an inverted glass, the water being
warmed by a spirit-lamp. Into this I put a suspended
larva of Byblia iithyia, a pupa seven days old, and another
two days old. I applied too much heat at first, keeping
the water at a boil, which killed the larva. I then turned
the lamp as low as possible, keeping the tin just hot
enough for the hand to bear. The older pupa emerged
in three days (normal pupal stage, thirteen to fifteen days)
and presents no marked peculiarity, as you may see, being
of the early wet-season form, which was the only form
occurring at that time in the natural state. The last

Seasonal Dimorphism in Butterflies. 205

pupa emerged after six days’ heating (eight days in pupa);
unfortunately it had a difficulty in emerging, and I arrived
too late to help it. But such as it is, it seems to me
a very interesting specimen, for it is clearly intermediate
in colouring, being therefore a step backwards towards the
dry form. — Its intermediate character is shown on the
underside of hind wings, in the deeper ground-colour and
more accentuated white bands, and on the upper side by
the broad interruption about radial nervules of the sub-
marginal black line in forewings, a character which only
occurs in the dry or intermediate form of the female, and
never in the early wet form of that sex.’—G. A. K. M.

The two specimens here mentioned are both in the
Hope collection. The difference between them is marked,
the one which emerged on Nov. 27, from the pupa which
was already seven days old before ‘being exposed to dry
warmth, being a wet-season male of the ordinary kind ;
while the other, which was only two days old when sub-
jected to the same dry warmth, emerging on Nov. 30, is
a crippled female, distinctly of the dry- season form, not
extreme, but quite unmistakable, and entirely differing from
specimens captured in the same locality at the same time
of year.

“ Malvern, Natal ; Feb. 21, 1897.—I have been trying to
find some reason to account for the occurrence of the marked
varieties of Biblia ilithyia. This again is a widespread
and common species, and comparatively conspicuous, so
that there must be some sort of protective agency
work. I can only explain it by the fact that DB. edithyia
strongly suggests an Acrva on the wing. Its general
coloration, somewhat elongated wings and flapping flight
(so different from that of its congeners), all tend to suggest
this. That the typical form does not actually resemble
any species of Acrva is of course plain, but I certainly
regard the variety acheloia as a marked stage of incipient
mimicry. On the underside, the hindwing of this variety,
in its wet-season form, differs from that of the type in
having lost the whitish bands, which gives it a very marked
resemblance to Acrwa serena-buxtont. Again, the loss of
the discal row of spots on the upper side of the hindwing
points the same way, and it is interesting to note that,
so far as my experience in South-east Africa goes, where
A. serena-buxtoni occurs, there acheloia prevails over
the typical form. Again, the chief difference between

204 Dr. F. A. Dixey on

the Central African serena and its southern sub-species
is that in the former the black band near the apex of
the forewing is continuous, but broken in the latter.
If I remember right, there is asomewhat similar difference
between acheloia and its Central African form goetzius,
which, if correct, would further bear out my idea. Now
as to the winter form; the underside of this is of course
quite unlike that of any Acrexa, and I can only suppose
that it is a case of protective resemblance on the principle
of the zebra’s or tiger’s stripes, for the insect always
roosts on grass. It is interesting to note however that that
part 1s undergoing modification in the variety acheloia,
as the marginal white line in both wings has already done.
It would be interesting to know whether there is any
likeness between this species and the Indian Acrwas.”—
G. A. K. M.

As I have elsewhere stated, I consider that Byblia
gotzius, Herbst, which Mr. Marshall here speaks of as
£. wlithyia var, acheloia, is entitled to distinct specific rank
beside SB. tlithyia, Drury. Mr. Marshall’s observation with
regard to the continuity of the apical black band of the
forewing in the Central African form of B. gétzius is borne
out on an examination of specimens in the Hope collec-
tion and the British Museum. It was remarked by me
some time since, in discussing the modifications of B.Jithyia
and its allies, that “the Socotran B. boydi resembles most
specimens of B. gdtzius from the West African subregion —
in having the dark costal bar of the forewing continued
rather heavily across the wing to join the submarginal
band. This is also more or less the case with two females
of L, gotzius from Abyssinia, and specimens of the same
from Somaliland and Aden in the British Museum; but in
examples from South and East Africa the connection be-
tween the costal and the submarginal dark bands is often
slight or absent.” * It is worth noting that the marginal
white line spoken of by Mr. Marshall, on the under-
side of both wings in the dry-season form of B. ilithyia,
has disappeared from the dry-season B. gotzius, but persists
in B, boydi, of which only the dry-season form is at present
known. This is another indication of the intermediate
position of the latter insect, which, though nearer to
B. gotzius, yet shows several points of resemblance to
B. ilithyia.

* Proc. Zool. Soc., 1898, p. 378.

Seasonal Dimorphism in Butterflies. 205

On the whole Mr. Marshall’s view as to the incipient
mimicry of Acrea serena, Fabr., by Bb. gotzius seems a
very probable one. The underside of the wet-season
B. ilithyia perhaps recalls slightly that of the Indian Acrwa
(Telchinia) viole, Fabr., but the likeness in this case is of
a remote kind.

“ Malvern, Natal; May 14, 1897.—EHaperiments on
submitting pupx to conditions of moistwre or dry heat.
The apparatus used for dry-forcing was a covered tin (into
which was poured a little water) placed on a tripod over a
spirit-lamp. On the lid of the tin was placed some dried
sand, into which was stuck a stick bearing the pupze, which
were covered with an inverted glass. The ‘damp tin’
contained very damp sand, the pup being separated
from it by a grating of perforated zimc; and the mouth
of the tin was covered with a cloth, on which was placed a
wet sponge.

“ EXPERIMENT WITH Acrva cabira.

1897

March 26. Two larvge (a and 6) pupated this morning ;

I put them in the dry forcer in the evening.

, 28. A larvee (c) pupated, and was left in the
breeding-cage.

» 931. Two larve (@ and e) pupated; d put in the
forcer, ¢ left in breeding-cage.

April 6. c¢ emerged, being a normal male.

- 8. eemerged, a normal female; d not yet
emerged, but still alive; @ and 0 probably
dead.

, 9 devidently too weak to emerge, so I helped
it out, but it was only just alive, and
wings did not expand. Its colouring was
apparently normal. a and 6 never
emerged at all, but shrivelled up.

“ Result.—Acrea cabira apparently unable to exist in a
very dry, hot climate, as might be supposed from its dis-
tribution. Itis noteworthy that two pupz of Zerias brigitta
emerged satisfactorily in forcer during the same period.

“EXPERIMENT WITH Pinacopteryx pigea.

1897
April 2. Seven larvee (a to g) pupated.

206 Dr. F. A. Dixey on

1897
April 3. Put two pupee (a and 0) in dry forcer; two
more (c and d@) in damp tin ; and left three
(e, f and g) in breeding-cage.
» 9. Took ¢ and d out of damp tin, as they
showed signs of emergence.
» 10. a,¢ and e emerged in the morning. «@ was
a female of the yellow form, showing an
approach to the dry-season form in a
slight reduction of all the black spots and
borders, especially the discal spot in fore-
wings; ¢ was a female of the white form,
and had all the black spots well marked ;
é was a white female, intermediate in the
development of black markings between

a and c¢.
Removed 0 from forcer to breeding-cage.
, ll. 0b,d,fandg emerged. b was a white female
5) ? g fo)

in which the black markings were not
quite so light as those of a, but noticeably
lighter than those of ¢; d was a normal
wet-season male; / and g were yellow
females intermediate in markings between
the extreme forms @ and e.

“ Result.—The differences exhibited are slight, but so far
as they go they apparently tend to show that the effect
of dry heat is to reduce the black markings, and that of
cool moisture to enhance them. It is to be observed that
yellow and white forms of the female occur at both seasons,
the deeper yellow specimens are however more prevalent in
winter. Reliable seasonal distinctions are greater or less
development of the marginal black spots and discal spot
in forewing, combined with less or greater acuteness of
forewing.

“EXPERIMENT WITH Crenis boisduvalit.

1897
Apml 9. Twenty-two larvee of C. boisduvalit pupated.
» 10. Put six pupze into dry forcer; six into damp
tin; and left the rest in breeding-cage.
» 14, Six pupze in forcer emerged; there were
four male and two female, but two of the
former were deformed.

Seasonal Dimorphism in Butterflies. 207

1897.

April 15. Three males and three females emerged in
damp tin; one male escaped and another
was deformed. Three males and six
females also emerged in breeding-cage.

“On comparing the three sets of specimens the difter-

ences were found to be remarkably slight, all the specimens
being of a more or less intermediate character between
the wet and dry season forms (as might be expected
during this month for those bred under normal conditions).
But such slight differences as do exist appear to be fairly
constant. In the females the black patches on the under-
side of the forewings are constantly best developed in
those from the damp tin and least in those from the forcer.
Those reared under normal conditions are much nearer the
former in this respect, being all rather lighter, except
examples which are hardly separable from those reared
under moist conditions. The differences in the hindwings
are too slight to be taken into account. In the males
those from the forcer show a slight difference from the
rest in having the black mark on the underside of
the forewing somewhat reduced, and a greater suffusion
of ochreous scales on the upper side of the hindwing. The
others are practically inseparable. The seasonal differences
in this species are very clearly defined as a rule.

“SECOND EXPERIMENT WITH Pinacopteryx piged.

1897.

April 8. Six larve pupated (a to /).

» 9. Two larve pupated (g and h). Put a, b and c
into dry forcer, and d and e into damp
tin.

, 10. Two larve (j and k) pupated. Put g into
damp tin.

, 15. Removed @ and 6b from forcer to breeding-
cage ; ¢ was dead; cause unknown.

, 16. a@and b emerged; both females.

, 17. f and h emerged in breeding-cage; both
females.

» » @& emerged in damp tin; female.

, 18. e emerged in damp tin ; female.

» » Jj emerged in breeding-cage ; female.

, 19. g ands emerged in damp tin and breeding-
cage respectively ; both males.

208 Dr. F. A. Dixey on

“No notes were kept of individual markings, but on
comparing the three sets it was noticeable, as in previous
experiments, that considering the disparity of conditions,
the markings showed wonderfully little difference. It is
however indisputable that, taking the specimens in con-
junction with those of the previous experiment, all those
subjected to dry heat had the black markings appreciably
less developed than those whose pupze were kept in a cool,
moist atmosphere. Those that were reared entirely in the
breeding-cage are mostly of an intermediate type of
colouring, though two are quite as bright as the heated
specimens, but none of them resemble those that were
kept damp.

“Tt is noteworthy that in Crenis boisduvalit the speci-
mens reared under normal conditions showed just the
opposite tendency.

“ Although the experiments are on far too small a scale
to prove anything one way or the other, yet to my mind
they appear to lend more support to the theory that the
heavier development of black markings in South African
butterflies during the summer is probably more dependent
on the prevalence of moisture than on the action of heat:
though the very small effects shown by these agents in the
above experiments suggest the supposition that the absence
or presence of black markings alone cannot be referred
entirely to climatic agency, as I had been previously
inclined to think, but have been developed by natural
selection, for some purpose not at present apparent, which
has worked on the slight tendency to variation caused by
climatic influence.” —G. A. K. M.

In 1896 Mr. Marshall had exposed some larvae of Acrwa
anacreon to “dry-season” conditions just before pupation,
but they all died in consequence, as he believes, of over-
heating (Estcourt, Oct. 15, 1896). On Oct. 7, 1897, he
writes from Malvern: “The experiment in which I found
that the pups of Acrwa cabira were killed by dry heat
which did not affect Terias brigitta, leads me to think
some of these highly-developed nauseous species may have
suffered in hardness of constitution, which would account
for their not spreading more widely than they do.”

Of the specimens referred to by Mr. Marshall in the two
series of experiments on Pinacopteryx pigea, Boisd., a, ¢ and
e of the first series, and a, 0, d, ¢, f and h of the second
series are in the Hope collection, The divergences noted

Seasonal Dimorphism in Butterflies. 209

as the result of the different treatment are more easily
visible in the first series than in the second.

The Hope collection also possesses seven specimens of
the above-mentioned series of Crenis boisduvalii, Wallgrn.
These are a pair of the “dry heat” emergence on April 14;
a pair of the “damp tin” emergence on April 15; and a
male and two females which emerged under normal con-
ditions, also on April 15. There is no doubt that the
“dry heat” female is considerably lighter on the upper
surface, and has the dark marks on the under surface of
the forewings less distinctly marked than any of the
others. The differences between the males are of the
same kind, but somewhat less apparent.

“ Aug. 29, 1899.—I am sending you by this mail a small
lot of butterflies, including the bred P. sesamus and archesia,
and twenty-one bred specimens of Teracolus omphale and
T’. achine, with their respective parents. . The Teracoli
will be valuable as actually proving seasonal dimorphism
in these species. I must admit that I was much surprised
to find that the warm, damp atmosphere had no effect on
LT. omphale (D1—4) whatever.* The apparatus I used was
a very deep circular tin (uncovered), which was partially
filled with water, in which was placed a stand ; to this
the pup were pinned, they being about four inches above
the water. In the case of 7, omphale (D1—4) I képt the
spirit-lamp with only a tiny flame, so as to keep the water
just hot, and so that a faint warmth could always be felt
on placing the hand above the mouth of the containing tin.
On account of the negative results thus obtained, I came -
to the conclusion that the heat applied was perhaps in-
sufficient in all these cases. Unfortunately, I had not
enough material left to test this properly, but in the case
of 7. achine (C1 and C2) I kept the water at about
180° F., still keeping the tin uncovered, and, as you will
see, this has undoubtedly had a more decided effect, espe-
cially in the case of C2, which was put in before actual
pupation. J was, however, surprised that with Cr the
protectively coloured under side should have been affected,
rather than the black markings of the upper side. In
view of this result I think the previous experiments must
not be taken as conclusive. Among the Teracoli there

* It appears to me to have had a slight effect, as can be seen on

comparing D2, D3 and D4 with D5, Do ‘and D7. _ See pp. 211-13.—
Be ASD:

210 Dr. F. A. Dixey on

is a highly interesting female omphale (KE, No. 15).”—
GaAn i MM.

The specimens of T'eracolus here spoken of were all ob-
tained.at Salisbury, Mashonaland. They are as follows :—

1. Teracolus achine, Cram.

X. A “wet-season” female (Figs. 5, 5a). Captured
March 26, 1899. Laid one egg.

X1. Offspring of X. From egg laid March 26; hatched
March 31; pupated April 23; kept under normal
conditions; emerged May 9. A “dry-season ”
female, not extreme, corresponding to the form
described by Trimen (South African Butterflies,
vol. i, 1899, p. 136) as; 7. antevippe, Boisd., 9.
(Figs. 6, 6a.)

B. An “intermediate” female. Captured April 28,
1899 ; laid 15 eggs.

Br. Offspring of B. Egg laid April 28 ; hatched April
29; pupated June 12; kept under normal con-
ditions; emerged July 20. <A dry-season male,
corresponding to 7. untevippe, Boisd., as described
by Trimen, Joe. cit.

B2. Offspring of B. Egg laid April 23; hatched April
29; pupated June 15; kept under normal con-
ditions ; emerged July 22. A well-marked dry-
season male, the pink of the hindwing under side
more pronounced than in Br. The left hindwing
is not completely expanded,

C. An intermediate female, verging towards “ dry.”
Captured April 26, 1899; laid 17 eggs.

Cr. Oftspring of C. Egg laid April 26; hatched May
3; exposed to damp heat from 10 p.m., June 22,
to 8 am., July 4. Emerged July 7. An inter-
mediate male, on the under side resembling the
wet-season form.

C2. Offspring of C. Egg laid April 26; hatched May 3;
exposed to damp heat from 10 p.m., June 22, to
8 am., July 4; pupated 8 am., June 23; emerged
July 8. An intermediate male, like C1, but some-
what more closely approaching the wet-season form
on the upper surface.

C3. Offspring of C. Egg laid April 26; hatched May 3;
pupated June 22; kept under normal conditions ;
emerged July 29. A male, intermediate on the

bo

Seasonal Dimorphism in Butter flies. 21d

upper surface, but with the under side decidedly of
the dry-season type.

C4. Offspring of C. Egg laid April 26; hatched May 3;
pupated June 28; kept under normal conditions ;
emerged Aug. 3. A dry-season female.

. Leracolus omphale, Godt.

D. A wet-season female (Figs. 7, 7a). Captured April
26, 1899. On the same day laid 19 eggs, which
hatched on May 3. Seven of the resulting butter-
flies are in the Hope collection, as follows :—

Di. Exposed to damp heat from 6 p.m., June 17, to
11 pm. June 25; pupated 11 pm, June 17;
emerged June 27. <A dry-season male, crippled.

D2. Pupated 2 p.m., June 17; damp heat 6 p.m., June
17, to 11 pm., June 25; emerged June 27. A
yellow dry-season female, imperfectly expanded.

D3. Damp heat 6 pm., June 17, till emergence;
pupated 8 pm., June 17; emerged June 28. A
yellow dry-season female.

D4. Pupated 2 p.m., June 17; damp heat 6 p.m., June
17, to 11 p.m., June 25; emerged June 29. A dry-
season male, not extreme.

D5. Pupated June 10; normal conditions; emerged
July 12. A white dry-season female, more ad-
vanced than D2 and D3; as shown by the dimin-
ution of the dark markings on the upper surface,
and the disappearance of the transverse bar and
orange-shaded discoidal spot on the under side of
the hindwing, traces of these being visible in both
the females D2 and D3, which had been exposed
as pupze to damp heat.

D6. Pupated June 10; normal conditions; emerged
July 14 (Figs. 8, 8a). A white dry-season female,
still more advanced than D5.

D7. Pupated June 16; normal conditions; emerged
July 17. A dry-season male, more advanced than
D4.

EK. A yellowish wet-season female. Captured April 30,
1899. Laid 15 eggs the same day. Offspring :—

Ki. Hatched May 8; pupated June 28; normal con-
ditions; emerged July 25. A white dry-season
female, with dark markings on disc of forewing
greatly reduced, and with a yellowish shade re-

TRANS. ENT. SOC. LOND. 1902.—PART Ul. (JUNE) 15

212 Dr. F. A. Dixey on

placing the orange in the centres of the apical
interspaces. This is the specimen referred to by
Mr. Marshall as E, No. 15 (p. 210).

E2 and 3. Hatched May 8; pupated June 28; normal
conditions; emerged July 26. Two dry-season
males.

F. A wet-season female. Captured May 3, 1899. Laid
2 eggs, which hatched on May 9. Offspring :—

Fr. Pupated June 27; normal conditions; emerged
July 26. A dry-season male.

F2. Pupated June 28; normal conditions; emerged
July 26. A yellow dry-season female, not extreme.

G, A wet-season female. Captured May 10,1899. Laid
10 eges. Offsprmg :—

Gr. Hatched May 18; reared under normal conditions ;
emerged July 31. A white dry-season female, not
extreme.

G2. Hatched May 18; normal conditions; emerged
Aug. 3. A dry-season male.

In all the above cases, the “ dry-season ” offspring of the
parent Zeracolus omphale corresponds generally with the
form described by Mr. Trimen (South African Butterflies,
vol. iii, 1889, p. 145) as 7. theogone, Boisd. The specific
identity of these two forms had long been suspected, and
by the above series of specimens is placed absolutely
beyond doubt.

In 1898 Mr. Marshall sent home a collection of butter-
flies from Salisbury, Mashonaland, which was described by
Dr. Butler in Proc. Zool. Soc., 1898, pp. 902-912. In an
accompanying letter to Dr. Butler he says: “I am some-
what in doubt as to the Zeracoli I have sent you labelled
pallene, for they are practically indistinguishable from the
extreme dry form of omphale; yet the wet form is cer-
tainly not omphale, which I do not remember ever to have
seen here, but seems referable to pallene.” Dr. Butler (oe.
cit., p. 911) “has not the least doubt that these examples
are ordinary 7’. omphale.’” An examination of similar
specimens sent to the Hope collection by Mr. Marshall as
T. pallene, led me independently to the same conclusion as
Dr. Butler; and it is worthy of notice that while several
of the bred examples just described are not separable from
Mr. Marshall’s specimens of 7’. pallene, the four parents,
all of which were captured at Salisbury, are identified by
Mr. Marshall himself as 7. omphale. The inference seems

Seasonal Dimorphism in Butterflies. 213

clear that there is no reason for considering Mr. Marshall’s
“7 pallene” from Mashonaland as specifically different
from 7. omphate.

It will be seen from the above descriptions that the
damp heat to which many of the specimens of 7. omphale
were exposed was not entirely without effect; though the
changes in the direction of the wet-season form are no
doubt less marked than those produced in the case of 7.
achine, where the heat employed was greater.

3. Teracolus phlegyas, Butl.* (7. dificilis, K. M. Sharpe).

One specimen: Salisbury, Mashonaland. Larva sus-
pended June 6, 1898; placed in damp forcer June 7;
pupated same day; removed June 30; emerged June 31.
An intermediate female, on the whole nearer to the dry
than to the wet-season form. The dark markings on the
upper surface of the forewings, including the discoidal
spot, are, however, somewhat strongly developed for a dry-
season form ; and there is a well-defined grey basal patch,
but no dark inner-marginal bar. Beneath, the hindwings
have lost the definite spots of the wet-season phase, but
have not assumed the dry-season colouring in its full
development.

This completes the list of specimens of Z'evacolus men-
tioned by Mr. Marshall in his letters. The succeeding ex-
tracts bear reference to the African forms of the genus
Byblia, Hiibn.

“ April 25, 1899.—I have a few authentic eggs of Byblia
ilithyia and acheloia, which may perhaps decide the justice
of Dixey’s contention as to the specific validity of the
latter.

“ April 19, 1901.—I hope to be able to get some definite
evidence as to Byblia, as I have now five pup and three
larvee bred from authenticated eggs of ilithyia, and one
pupa and six larve from those of vulgaris, i.e. the wet-
season form of B. acheloia (=. gotzius). The resulting
butterflies will also prove the seasonal variation in the
two forms. So far as my present material goes, I find
that there is a very slight colour-distinction between the
two larvee in the last stage only.

“Sept. 27, 1901.—The specimens resulting from my
damp experiments, together with those already sent, might

* This, though belonging to an earlier series of experiments, is
inserted here for convenience

214 Dr. F. A. Dixey on

form the nucleus of a most interesting and instructive
series to show the experimental evidence as to the proxi-
mate causes of seasonal dimorphism.... . You will
find some of the specimens from my Syblia experiments.
The few that emerged all bred true to their parents, but
the principal evidence consists in a slight, though constant,
colour-distinction which I found in the larvee of the two
insects, thus proving them to be distinct species.”—
Gi As KM

It is satisfactory to me to find that in consequence of
his latest experiments, Mr. Marshall now holds the view
as to the specific distinction between the two continental
forms of Byblia which I felt justified in putting forward in
1898.* The specimens recently forwarded by him to the
Hope collection from Salisbury, Mashonaland, are as
follows :—

A. B. ilithyia, Drury. A worn wet-season female.
Captured March 17, 1901. Laid 3 eggs. Off
spring :—

At. Hatched March 21; pupated April 11; emerged
April 28. A wet-season female.

A2. Hatched March 21; pupated April 11; emerged
April 29. An intermediate male.

B. B. ilithyia, Drury. A worn wet-season female (Fig.
1). Captured March 17, 1901. Laid 5 eggs. Offt-
spring :—

Br. Egg laid March 24; hatched March 28; pupated
April 11; emerged April 29. An intermediate
male.

B2. Ege laid March 24; hatched March 28 ; pupated
April 11; emerged April 30. An intermediate
female (Fig. 2).

B3. Ege laid March 24; hatched March 28; pupated

55

April 11; emerged May 1. An intermediate

the form vulgaris, Staud. (Fig. 3). Captured March
24,1901. Laid 6 eggs. Offspring :—
C1. Egg, March 24; hatched March 28; pupated April

* Proce. Zool. Soc., 1898, p. 376. The current number (Feb. 1902)
of the “Entomologist’s Monthly Magazine” contains the first instal-
ment of a paper by Mr. Marshall in which he gives a detailed account
of his experiments in the breeding of Byblia ilithyia and B. gétzius,
with descriptions of larve and pupe.

Seasonal Dimorphism in Butterflies. 215

22; emerged May 8. A dry-season male, of the
form acheloia, Wallgrn. (Fig. 4).
C2. Egg, March 24; hatched March 28; pupated April
24; emerged May 13. A dry-season male, lke
C1.
E. B. ilithyia, Drury. A wet-season female. Captured
March 24,1901. Laid 2 eggs. Offspring :—
Er. Egg, March 24; hatched March 28; pupated April
27; emerged May 19. An intermediate male.
These specimens supply complete proof, if proof were
wanted, of the specific identity of B. vulgaris, Staud. with
B. acheloia, Wallgrn., and also of B. ahithyia, Drury, with
the African form corresponding to B. simplex, Butl., of
India. It is to be noted that none of the bred B. aithyra
are of the full dry-season form, One or two of them, how-
ever, approach it so closely as to leave no manner of doubt
that later in the year the typical “dry-season ” colouring
would be developed.
The following specimens of Zerias sent home by Mr.
Marshall are also worthy of note :—

Terias brigitta, Cram.

A. Malvern, Natai. Pupa in dry heat 6 days; emerged
April 4, 1897. A wet-season male.

This is no doubt one of the two 7. brigitta mentioned
above (p. 205) as having withstood an amount of heat
which proved fatal to Acrewa cabira.

B. Malvern, Natal. Pupa under normal conditions ;

emerged April 9, 1897. A wet-season female.

Terias senegalensis, Boisd.

A. Salisbury, Mashonaland. Captured April 7, 1901.
(Figs. 9, 9a.) Laid 3 eggs. A wet-season female
of 7’. senegalensis, Boisd. Offspring -——

At. Egg laid April 7; hatched April 11; emerged June

10. (Figs. 10, 10a.) A dry-season male, of the
form 7 ethiopica, Trimen.

These two specimens are of great interest, as showing
that a 7. hapale-like form (7. xthiopica) may be bred from
a 7. hecabe-like parent (7. senegalensis) ; and as thus tend-
ing in some respects to confirm Mr, Marshall’s view
expressed to Dr. Butler in 1898 as follows :—

“ You will notice among the Zerias that I have poimted

216 Dr. F. A. Dixey on

out that 7. xthiopica and butlert of Trimen are respectively
dry and wet forms of the same species, and thus, taking the
synonymy given in your revision, hapale must fall as a
seasonal form of senegalensis. I have not actually proved
the case by breeding, but I think you can take my obser-
vations on trust now.”* | may mention that I had some
time ago come independently to the conclusion that the
T. hapale forms could not be specifically separated from
the 7. senegalensis assemblage, and had arranged the
examples in the Hope collection in accordance with that
view. But I do not think that even now the seasonal
relations of these forms are quite clear.

In addition to the series just described, Mr. Marshall has
also presented to the Hope collection the greater number
of the specimens resulting from the experiments recorded
by him in the “ Annals and Magazine of Natural History,”
1901, vol. u, p. 898. They exemplify the very slight effect
produced on the early dry-season broods by subjecting the
larvee and pupee to conditions of moisture without heat.
In Mr. Marshall’s opinion, the amount of occasional inclin-
ation towards the wet-season form shown in this series is
no more than might have been met with in examples of
similar dates caught in the open. These specimens need
no further notice here, having been fully dealt with by Mr.
Marshall in his paper above referred to.

5. SUMMARY.

The main points of the present paper may be sum-
marized as follows :—

1. Catopsilia pomona, Fabr. (including C. catilla, Cram.),
and C. crocale, Cram. are phases of a single species. In at
least one part of its range, these phases appear to be in
relation with the seasons ; in other parts there seems to be
no such connection.

2. In like manner Catopsilia pyranthe, Linn. is conspeci-
fic with C. gnoma, Fabr. Here the association of each
form with its own season is better recognized, but there is
reason to think that even in this case the relation by no
means obtains universally.

3. There are many other instances on record of the
simultaneous occurrence in a given locality of forms of a

* Proc, Zool. Soc., 1898, p. 909.

Seasonal Dimorphism in Butterflies. 217

species which are either known to be characteristic of the
seasons in other parts of the range of the species, or which
at least are analogous with proved cases of seasonal
dimorphism.

4. Some of these cases of simultaneous occurrence are
undoubtedly due to an overlapping at the change of
seasons. In other instances the intermingling of the
different forms takes place indifferently all the year round.
This is perhaps more especially apt to occur in regions
where the climate does not show very well-marked
alternations between wet and dry.

5. Mr. Marshall has proved the specific identity of the
following pairs of forms by actually breeding one from the
other :—(a) Teracolus topha, Wallgr., and 7’. auxo, Lue. ;
(6) Teracolus achine, Cram., and T. antevippe, Boisd.; (¢)
Teracolus omphale, Godt., and 7. theogone, Boisd.; (d) Terras
senegalensis, Boisd., and 7. #xthiopica, Trim.; (e) Byblia
gotzius, Herbst (vulgaris, Staud.), and B. acheloia, Wallgrn. ;
(f) Byblia wlithyia, Drury, and the African form of JB.
simplex, Butl. In each of these cases it was already
known that the different forms were respectively associ-
ated with different seasons, but the actual proof of specific
identity afforded by “breeding through” had _ hitherto
been wanting.

6. The final stage can in many cases be influenced by
the artificial application of heat or moisture during the
pupal condition. Thus, Mr. Marshall has found that dry
warmth may cause the early wet-season form of Byblia
ilithyia to approach the dry-season type of coloration ;
while the intermediate or early dry-season forms of
Pinacopteryx pigea and Crenis boisduvalii were slightly
affected in the same direction. Warmth in conjunction
with moisture produced in early dry-season forms a
tendency to revert to the garb of the rains. This was well
seen in Teracolus achine, and to a less extent in 7’. phlegyas
and 7. omphale. On the other hand, neither the early
wet-season form of 7’. annx (1896), nor the early dry-season
forms of several other species (1901) seem to have been
attected by the application of moisture without the addition
of heat, though a tendency towards the wet-season form
made itself apparent under these conditions in Pinacopteryx
pigea and, to a slighter extent, in Crenis boisduvali.

7. Mr. Marshall has now detected constant differences in
the respective larvee and pup, which prove that Bybla

218 Dr. F. A. Dixey on Seasonal Dimorphism, etc.

gotzius (including B. vulgaris) is specifically distinct from
B. whithyia.

In conclusion, I wish to thank the authorities of the
British Museum of Natural History, particularly Dr.
Butler and Mr. Heron, for help that has always been
courteously and readily given. My obligations to Professor
Poulton are still more weighty; I owe to him the enjoy-
ment of complete facilities for work in the Hope Depart-
ment at Oxford, the loan of Mr. Marshall’s letters, and the
photographs of specimens that illustrate this paper. With
regard to Mr. Marshall himself, I should wish to place on
record my sense of the high value of his work as collector,
experimenter and observer. He has had good opportunities,
which he has known how to use in a thoroughly scientific
manner. — Moreover, what he has already achieved justifies
us in looking for still greater results from his persevering
labours.

EXPLANATION OF PLATE IV,

Fig. 1 (underside). Byblia ilithyia, Drury. A wet-season female.

2 (underside). Offspring of the above. An intermediate female,
approaching the ‘‘ dry ” form which corresponds to the Indian
B. simplex, Butl. ;

8 (underside). Byblia gotzius, Herbst. A wet-season female, of
the form vulgaris, Staud.

4 (underside). Offspring of the above. A dry-season male, of the
form acheloia, Wallgrn.

(See pages 214, 215.)

Or

, 5a (underside). Teracolus achine, Cram. A wet-season female.
6, 6a (underside). Offspring of the above. A dry-season female
of the form described by Trimen as antevippe, Boisd.
7, 7a (underside). Zeracolus omphale, Godt. A wet-season female.
, 8a (underside). Offspring of the above. A dry-season female,
of the form described by Trimen as theogone, Boisd.

(See pages 210-211.)

co

9, 9a (underside). Terias senegalensis, Boisd. A wet-season female.
10, 10a (underside). Offspring of the above. A dry-season male,
of the form xthiopica, Trimen.
(See page 215.)
In the actual specimens, owing to the presence of colour, the difference

between the wet- and dry-season forms of the same species is more striking
than appears in the Plate.

Pies NW

Trans. Ent. Soc., Lond., 1902.

Fre. 9.

=.)
yy . ' 2 .
Fic. 10

ip

tae
f y 2
be
= Ye
Y f = to § “A
4 “E >
y * 7A & Ld
w ; i J
A \ >
a bs ove

* y, } ‘ \
ys ee Cay, Fic. 10a.

André & Sleigh, Limited.

Dixey.
All the figures ave about % of the natuval size.

Seasonal Dimorphism in Butterflies.

XVIT. Five Vears’ Observations and Experiments (1896—
1901) on the Bionomies of South African Insects,
chiefly directed to the Investigation of Mimicry and
Warning Colours, by Guy A. K. MARSHALL, F.Z5.
With a Discussion of the Results and Other Subjects
suggested by them, by Epwarp B. Poutton, M.A,,

f=)

D.Se., F.R.S., Hope Professor of Zoology in the
University of Oxford, Fellow of Jesus College,
Oxford; and an Appendix containing Descriptions
of New Species by Ernest E. AUSTEN, WILLIAM
L. Distant, Colonel CHARLES T. BINGHAM, E.ZS.,

Guy A. K. MARSHALL, and JULES BOURGEOIS.
[Read March 5, 1902. ]

PRArHs DXe To) Oxi.

CONTENTS.

1. IntRopuction. By Guy A. K. Marswaun and E. B.
POULTON NS pee eek een Das eee aglaw aes

2. EXPERIMENTS ON MAntipa: IN NATAL AND RHODESIA,
KCrap As Wee Mls) resents) eer ied tne Gs eee Seek hn eal

3. CoNCLUSIONS FROM EXPERIMENTS ON Manvipa&. (EK, B. P.)

4, EXPERIMENTS ON SPIDERS IN THE Karkuoor, (G, A.
IGM IVIG ete ee grants pee ee sn On ep afc. “icly S

5. REesuLTS oF EXPERIMENTS ON SPIDERS AND THE EARLIER
EXPERIMENTS ON MANTIDA : ONE PROBABLE MEANING
oF THE TENACITY OF Lire IN DistasteruL INsEcTs.
(Ga Ac Kea): RR ce ir home oes comme

6. THe ATracks or PREDACEOUS INSECTS OTHER THAN
MaNTIDZ UPON CONSPICUOUS SPECIALLY-DEFENDED
LEPIDOPTERA, ETC. (E. B. P.)

A. Predaceous Hymenoptera and Newroptere .

B. Predaceous Coleoptera.

C. Predaceous Diptera. ea neater ee eat. ba 5

7. LEPIDOPTERA WITH WARNING COLOURS SPECIALLY LIABLE
TO THE ATTACKS OF Parasitic Insects. (G. A. K. M.)

8. EXPERIMENTS ON LizARDS AND Froes. (G. A. K.M.) .

9, EXPERIMENTS ON CAPTIVE KESTRELS (Cerchneis rwupi-
coloides and C. nawmanni). (G. A. K. M.) .

TRANS. ENT. SOC. LOND. 1902.—PART III. (NOV.)

288 Mr. G. A. K. Marshall on

10,

wile

12,

14,

15.

16.

Wie

28.

EXPERIMENTS ON A TAME GROUND HoRN-BILL (Bucorax
caffer). (G. ARKS M.)e see nade eo Odd
THE INSECT-FOOD OF WILD SouTH Armtre any Bieps. (G.
AK. M) 5: 9G oe See
Records oF ATTACKS ON LEPIDOPTERA, ESPECIALLY
BUTTERFLIES, BY WILD SoutH AFRICAN Birps. (G.
A. K. M.) ; :
RECORDS OF ATTACKS ON anes BY WILD irae
IN INDIA AND CrYLon. By Cou. J. W. YERBURY,
R.A. conan ane a
RECORDS OF Anmugera. ON + [Eyam san, ETC., BY WILD
BurMEsE Birps. By Con. C. T. BinaHam
Guy A. K. MARSHALL’S INDIRECT EVIDENCE OF THE
ATTACKS UPON ButtTeRFuies. (E. B. P.)
EXPERIMENTS ON A CAPTIVE MUNGOOSE WITH INSECT-FOOD,
(GicAy KM). ty i:
EXPERIMENTS ON A CAPTIVE Mekeges: WITH ‘im FOOD,
WARNING CHARACTERS AND DISTASTEFUL QUALITIES
IN SoutH AFRICAN Birps. (G. A. K. M.)
EXPERIMENTS ON Cercopithecus pygerythrus. (G. A. KK. M.)
EXPERIMENTS ON CAPTIVE Bapoons. (G. A. K. M.)

. CONCLUSIONS FROM EXPERIMENTS ON CAPTIVE BABOONS,

MuUNGOoosE, AND Kerstrets. (G.A.K.M.) .

. CONCLUSIONS FROM EXPERIMENTS ON Bapoons. (E. B. P.)

THE CHIEF CONSPICUOUS SPECIALLY-DEFENDED GROUPS IN
THE COLEOPTERA INFERRED FROM G, A, K. MARSHALL’S
EXPERIMENTS. A COMPARISON BETWEEN COLEOPTERA
AND LEPIDOPTERA IN THIS RESPECT. (HE. B. P.).

. EXPERIMENTAL EVIDENCE OF THE VALUE OF THE TERRI-

FYING MARKINGS IN Cherocampa LARVa&, (G. A. K. M.)

. EVIDENCE OF A SUPERSTITIOUS DREAD OF THE LARVA OF

Cherocampa elpenor. (K. B. P.) :
EXPERIMENTAL. EVIDENCE OF TERROR CAUSED BY THE
SQUEAK OF Acherontia atropos. (G. A. K. M.)

3. INSECT STRIDULATION AS A WARNING OR INTIMIDATING

CHarRactER, (G. A. K. M.). ae

HumMAN EXPERIENCE OF THE ‘TASTE AND SMELL OF ie
SECTS AFFORDS UNTRUSTWORTHY EVIDENCE OF THE
EFFECT UPON THE SENSES OF INSECTIVOROUS ANIMALS,
(EBB) iis :

Guy A. K. MarsHa.u’s Panee OF SpASONAL Ouandne
IN Sourn ArricaAN BurTeERFLIES OF THE Genus Precis,
(EN BSPs)

A, Introduction .

347

348

353

359

361

366

376

378

379

380

387
388

392

397

399

402

403

405

414
414

The Bionomics of South African Insects.

B. Historical oat:

C. The Demonstration by Gor Ae K. Mine Het (pases
simia is the Wet Phase of P. antilope . :

D. The Habits of the two Seasonal Phases of the ‘South
African Species of the Genus Precis, and the Stations
they respectively occupy . AS, AMEE page <

E. Hvidence of Adaptation in the conspicuous wnder-sides a
the Wet-Season Phases almost equal to the ee of it
an the cryptic Diy Phases :

F. The severity of the Struggle for Gentens among Tisects
tm the African Dry Season as compared with the Wet.
The relation of the Seasonal Changes in Precis to those
of other Butterflies aie ae

G. The succession of the two Seasonal eines of Precis
sesamus in Nature Pai ‘ .

H. The attempt to control the Phases of P. sesamus an
P. archesia by the artificial application of Moisture
and Heat to the earlier stages. Re lines of
Experiment . ne

I. The Bearing of the Seas oiead Biles of Prec upon ‘the
Science of Insect Systematics 5 ;

29. THE GREGARIOUS INSTINCT IN HYBERNATION AND Er
GRATION oF Insects. (E. B. P.) oo

30, DEscRIPTION AND Discussion oF MATERIAL BEARING ON

Mimicry In SourH AFRICAN RHOPALOCERA COLLECTED

BY Guy A. K. MARSHALL, AND THE RECORD OF

OBSERVATIONS MADE BY HIM. (E. B. P.)

Black-and-White Amauris-like Group

Limnas chrysippus-like Growps :

The Origin and Meaning of the Three Oni For ms oR

Limnas chrysippus .

D, A study of Mimetic Forms may able Ws to Sones ier
the Lost Stages through which the Older Model has
passed .

nF

E, Amauris echeria-like Croan marked Secondary Resem-
blances between the Forms mimicking echeria s

F. The Origin of the black-marked golden-broun Tr “opie
at the base of the Hind-Wing under-side in many
Ethiopian Butterflies, ‘

G. Compound Group containing Bopresenianioes of all ‘the
three previously described. Species probably entering
two Groups $ eee ee NS

H. Groups of Synaposematic Hiotieas sped al the same
Place und Time

289

415

450,

458

460

466
466
468

488

490

492,

TRANS, ENT. SOC, LOND. 1902. =D ARM SnnE “(ov.) 20

290 Mr. G. A. K. Marshall on

I. Mimetic Species of South African Lycenide and Hes-
peride captured with their Models

J. Mimicry in Lycenide and to a less extent in Hesper de
a Character of the Ethiopian a Possible Inter-
pretation .
K. Mimicry in the Nuanahedane Batonune or Miilleri tan 2
L. Miscellaneous Observations on Mimicry in South ks ican
Rhopalocera. (G. A. IKK. M.) .
31. WARNING CoLOURS AND MIMICRY (ALMOST WHOLLY Win

LERIAN) IN SourH ArricaN CoLEopTEeRA. (G. A. K.

Mand) HB. RP.)
A. Peculiar Warning eaten ana Directive Maris in
Carabide and Cicindelide. (EK. B. P. and G. A. K. M.)
B. Mutilloid Coleoptera: Cleridx, Carabide, and Cicinde-
lide: Primary and Secondary Synaposematic and
Pseudaposematic Associations, (EK. B. P.)
C. Lycide as Models for other Coleoptera and Insects of
many Orders. (G. A, K. M.). nin et eg ee
D, Miillerian (Synaposematic) Groups in South African
Coleoptera. (G. A. K. M.)
a. Cantharid Group one
B. Intermediate Group senneetny ae Gantt
and Coccinelloid Groups .
y- Coccinelloid Group .
6. Group of small pale elton aad fad Phatonhate
with their Melyrid and- Cureculionid Mimics
(CHEE) ee ee de ge ez
E. Comparison between certain Coleopterous Groups in
Borneo and South Africa, with respect to Mimicry,
Common Warning Colours, ete. (G. A. K. M.)
F. Note on Rhynchophora with Preeryptic Colowring as
Models for Mimicry. (E. B. P.) mee
32. Common WARNING CoLouRS IN SouTH Awroan Hy-
MENOPTERA AND THE MIMICRY OF THEM BY INSECTS
OF OTHER ORDERS. (G, A. K. M.) “
A. Group with Black Bodies and Dark Blue Wings, ety
Fossores
. Mimicry of aentens by Resouces ana tate of oer
Orders. (EK. B. P.) AR
C. Group with Black Bodies and Y lees Tails, chee
Diploptera
D. Group with Dark Bodies, Genus WI ae edi an Red.
Brown Tails: Megachile the Models Spee
E. Group with Black Thorax and Yellow Abdomen, all
Hymenoptera

ee)

493

The Bionomics of South African Insects.

F. Group with Black Bodies, Blue Wings, and Yellow or

Red Thorax.

G. Group with Black Bodies, Blue Wings, and Red or

Yellow Heads . Pe oe RE ES Mid y, PSS
H. Group with Black and Yellow-Banded Bodies, all

Hymenoptera ER: Ge cee Ae ae 35:
I. Group with Dark Wings and Black-and- Yellow Legs:

Ichneumonid Models. 3 :
J. Black and Yellow-Barved Braconid Ghote aa ante one
K. Black and Red Braconid Group and Mimics .

L. Diptera mimicking Single Species of Hymenoptera father

than the General Type of a Group
t. Asilid Fly mimicking Xylocopid Bie:
2. Syrphid Fly mimicking a Wasp .
y. Bee-like Group
M. Group of Ant-like Insects puplued Posgetliae :
33. Mimetic RESEMBLANCE OF MANTISPIDZ TO Hyew-
OPTERA. (KE. B. P.).
34. CONVERGENT GROUPS oF SoUTH AvRIGAN Heurprena,
(GAL KE M.)) 4 ‘
A. Black and Red Lyqeoid Group
B. Group of Yellow Hemiptera with Black Vie ana one or
two Black Bars.
35. MISCELLANEOUS OBSERVATIONS ON Sour AFRICAN Ty.
sects. (G. A. K.M.) . ; :
A. Note on the Courtship of Limnas Ge YSippus .
B. The possible meaning of the Suc of Female Acreine .
C. A Rhodesian Muscid Fly Parasitic on Man .
APPENDIX
EXPLANATION OF Pranns
INDEX .

5382
532
533

533
533
534
534
539

536

292 Mr. G. A. K. Marshall on

1. INTRODUCTION:
A. By Guy A. K. MARSHALL.

THE observations and. experiments which form the
groundwork of the present memoir were originally under-
taken by me at the imstance of Prof. Poulton, and such
interest as they may possess is largely due to his valuable
suggestions and advice. Moreover he has been good
enough to undertake the entire clerical work in connec-
tion with the publication of the paper, and he alone is
responsible for the numerous excellent plates with which
it is illustrated. The utility of experiments such as here
recorded depends almost entirely upon the manner in
which the results may be treated. The mere accumula-
tion of facts of this kind has little real value, unless these
facts are properly classified and co-ordinated, and their
bearing upon current theories adequately considered and
discussed. This portion of the work has been left almost
entirely in Prof. Poulton’s hands, and I feel that I am
fortunate in having obtained his hearty co-operation ; for
his wide experience in this particular line of research
insures a thorough treatment of the subject.

In carrying out the experiments I have always en-
deavoured, so far as in me lay, to record the results as
impartially as possible. But on reviewing my experiences
as a whole I cannot escape the conclusion that they lend
very strong support to the theories of Mimicry and
Warning Colours as enunciated by Bates, Fritz Miiller,
and Wallace; I feel convinced that were naturalists more
ready to carry out extensive experiments of this nature
there would be much less of the prevalent @ priori eriti-
cism of these valuable theories which throw light upon a
vast number of facts which must otherwise remain for us
mere meaningless coincidences. It is especially important
that experiments should be made by as many different
observers as possible, for in this way alone can the errors
due to unavoidable personal bias be eliminated; and if
the present publication only has the effect of inducing
other entomologists in South Africa, or elsewhere, to turn
their attention to the interesting problems involved, it
will have fully served its purpose.

G. AWK Me

The Bionomics of South African Insects, 293

BY By EB) Pownron:

The following memoir has been written upon and
around the oveat mass of valuable material supplied by
Mr. Guy A. “K. Marshall’s observations, experiments, and
nad from 1896 to 1901. So far as this material
consists of specimens it is open to the study and criti-
cism of all naturalists; for it has been placed by the
generosity of Mr, Marshall in the bionomic series of the Hope
Department in the Oxford University Museum, The
paper itself has been gradually growing during these years,
not only by the accumulation of specimens, but by an
uninterrupted correspondence between Mr. Marshall and
myself. Extracts from Mr. Marshall’s letters form a very
important part of the whole work, and it is only right to
point out that they were not written for publication, and
that any want of co-ordination or continuity is entirely
due to this cause. At the time when they were selected
and arranged for publication there was no prospect of Mr,
Marshall’s return to England, and I was anxious that as
many naturalists as possible might have the opportunity
of reading the observations and discussions from which I
had learnt so much and received such great pleasure ; and
when eventually he did return the paper had been
read. Although no attempt was made to alter or re-write
these extracts, Mr. Marshall’s presence in England has
made an immense difference in the work. We have been
able to discuss the general arrangement and illustration as
well as the details of many obscure and difficult subjects.
On several points he has written paragraphs which give a
far higher value to the paper. Where the experience of
the naturalist on the spot has been specially required it
has become available. ‘The sections of the paper under my
own name have also greatly benefited by his kind assist-
ance, and the opportunity of discussing points of special
difficulty or uncertainty. It will be clear to all who read
the paper that Mr. Marshall and I do not entirely agree in
the interpretation of many facts, especially those connected
with the seasonal phases of Precis, and in the extent and
predominance of Miillerian mimicry as compared with
Batesian in Lepidoptera. For these and other reasons it
is necessary to state explicitly that I am solely responsible

294 Mr. G. A. K. Marshall on

for the opinions and considerations set forth in the sections
to the titles of which the initials “ E. B. P.” are appended (in
both the contents and the text). Mr. Marshall’s numerous
and important contributions to these sections are always
acknowledged and placed between inverted commas. The
titles of Mr. Marshall’s sections are indicated by the initials
“G. A. K. M.,” and my contributions to these are always
placed between square brackets, and are furthermore
indicated by my initials.

Colonel J. W. Yerbury has kindly contributed one section
and Colonel C. T. Bingham another, and both have given
much help in other parts of the work. Some of the most
strange and interesting insects were undescribed species,
and would have been comparatively valueless for the
purpose of this memoir, were it not for the kind assistance
of the naturalists who have written the Appendix. Dr.
F. A. Dixey kindly read the proofs and made many valuable
suggestions and corrections. Mr. C. J. Gahan has given
much kind assistance in the sections dealing with Cole-
optera and in the identification of species. The number of
species sent by Mr. Marshall is so large that the work of
identification has been very laborious and prolonged, and
we desire warmly to thank Sir George Hampson and the
whole of the staff of the Insect Department of the British
Museum, every one of whom has been consulted at one
time or another. We also wish to thank heartily Colonel
C. T. Bmgham, who has named the whole of the
Hymenoptera; Colonel J. W. Yerbury, who has worked
out the majority of the Diptera; Mr. M. Jacoby, who has
named many Phytophaga; Monsieur Jules Bourgeois, who
has named the Lycidv, and Mr. W. L. Distant, who has
named the Hemiptera. Much other kind assistance has
been given and is acknowledged in the text of the work.

Valuable material -with excellent data, comparing in a
most interesting manner with that sent by Mr. Marshall,
was contributed from British East Africa by my kind
friends Mr. and Mrs. 8. L. Hinde.

The thirteen uncoloured plates are reproduced from
excellent negatives taken from the actual specimens by
Mr. Alfred Robinson in the Oxford University Museum.
The two coloured plates are reproduced from Mr, Horace
Knight’s drawings of the specimens.

A brief abstract of some of the chief results here recorded

The Bionomics of South African Insects, 295

in detail was communicated to the Zoological Section of
the British Association at Bradford (Report 1900, pp.
793-4), and an abstract of the present paper is printed in
the Proceedings of the meeting at which it was read (Proc.
Ent. Soc. Lond., 1902, pp. x—xiii). Some of the observa-
tions were also brought before the International Zoological
Congress at Berlin, 1901 (Verhandlung, p. 171). Lists of
the specimens presented to the Hope Department and a
brief statement of the principles which they illustrate
have been published yearly in the “Report of the Hope
Professor of Zoology” communicated to the “Oxford
University Gazette.” Allusion to some of the material
and the problems it illustrates, has also been made by the
present writer in Linn. Soe. Journ. Zool., vol. xxvi, 1898,
p- 558, and Report Brit. Assoc., 1897, p. 689. Much has
been written upon the work on seasonal dimorphism in
the genus Precis, but full references will be found in this
section of the present paper.

The first part of the following work, occupying just
half of it, deals with experiments and observations upon
insectivorous animals, and the conclusions and considerations
arising out of this work. The experiments on Mantide,
Kestrels, and baboons will be found to be especially
numerous and important. A table shows all the examples
of Asilidx and the species forming their prey which could
be found recorded or preserved in the British Museum
and Hope Collection. The direct and indirect evidence
of the attacks of birds on butterflies meets objections
which are often raised, and indeed nearly the whole of
this part of the paper is an effective reply to those who
ask for facts rather than hypotheses. One very important
side of the work is the employment of Coleoptera on a
large scale, and the clear evidence of aposematic and
synaposematic colours in the group. A comparison between
the Coleoptera and Lepidoptera in this respect is attempted.
The first half of the memoir ends with a section discussing
and criticizing the conclusion that there is any great signifi-
cance or value in human experience of the taste and smell
of insects.

The second half of the work is more heterogeneous.
Its first section attempts to supply an interpretation of
the startling seasonal phases of butterflies of the genus
Precis. In this section Dr. A. G. Butler’s convenient

296 Mr. G. A. K. Marshall on

terms “ wet phase” and “dry phase” * are generally used
in preference to “ form” or “variety,” while Mr. Marshall’s
useful sign €) to indicate the former and © to indicate the
latter are freely employed. The remainder of the paper is
chiefly devoted to the description of an immense mass of
material illustrating mimicry and common warning colours
in Rhopalocera, Coleoptera, Hymenoptera, and to a less
extent Hemiptera. Many interesting conclusions emerge
and are discussed.

I entirely agree with Mr. Marshall’s opmion that an
unbiassed consideration of the facts presented in this paper
yields a very strong measure of support to the classical
theories of Bates, Wallace and Fritz Miiller. I would go
further and maintain that Mr. Marshall’s observations and
experiments here recorded, place Africa in the first position
as the region which supplies stronger evidence than any
other of the validity of these theories. But Iam even
more impressed by the strong support yielded to the
modern developments of Fritz Miiller’s theory of mimicry.
Where has Professor Meldola’s Miillerian explanation in
1882 of the common facies of specially-protected sub-
families of butterflies received such illustration as in the
groups of synaposematic Acrwinw captured in one place
and at one time; or the extension in 1887 by the
present writer of the same interpretation to the types of
insect colour and pattern which are common to a country,
received such support as in the marvellous group of
Mashonaland insects of many Orders with an appearance
founded upon that of the distasteful Coleopterous genus,
Tycus? And the most recent developments of all, the
discovery (1894-7) of the principle of “reciprocal mimicry ”

or “diaposematic resemblance,” and of the specially close
See resemblance of the females in Miillerian mimicry
no less than in Batesian by Dr. Dixey, together with his
Miillerian interpretation of resemblances between mimics
overlying their resemblance to a common model, all these,
founded on the study of Neotropical forms, have supplied
the explanation of numerous instances in the Ethopian
Region although applied to very different families and

* The term “phase” is advantageous inasmuch as it is conveniently
applicable to the whole of the winter or summer generations of a
species, as well as to single individuals of either seasonal form,

The Bionomies of South African Insects. 297

sub-families of butterflies, to Coleoptera as well as to
Lepidoptera.

T cannot conclude without warmly thanking my friend
Mr. Marshall for the pleasure [I have enjoyed in the work
which we have done together.

ii Bi Be

2. EXPERIMENTS ON MANTIDH IN NATAL AND RHODESIA.
(G. A. KK. M.) Natal, February 1897.

I. Gave a wingless Acrva horta to a Mantis. It seized
it and threw it away. On a second presentation it felt
the butterfly carefully with its antenne, then took it and
began eating first the haustellum, then the palpi, and
finally the whole head with apparent relish. On biting
at the thorax, however, it threw it down with evident
disgust and began wiping its mouth on its fore-legs as
though to take away the taste. I again presented the
butterfly, but the Mantis at first only ran away from it.
At last it took it again and began eating the thorax, but
quickly threw it down and would have nothing more to
do with it.

II. Experiment ¢.—Caught a full-grown Mantis and put
it in a large green gauze bag. In the afternoon put in
a house-fly, which was not eaten that day, but was gone
next morning. Then put in a wingless male A. horta (a
bitter yellow juice exuded from the wing stumps). On
perceiving it the Mantis ran towards it, seized it and made
a bite at the back of the thorax, but started back as if in
great surprise, and wiped his mouth on his front legs. He
exhibited both fear and curiosity ; for as the Acrva ap-
proached he edged away, just keeping far enough off to
be able to touch it with the end of his long antenna, and
when the Acrva walked away he followed, still feeling it
over. At this pot I was called away, and on my return
found that the Acreva had been eaten all except the head and
apical half of the abdomen. Afterwards put into the bag
the Amauris echeria which had been rejected by spider C
(Experiment 13), and which was half dead. As the
Mantis took no notice of it I left, but on inspection in
the evening I found that this butterfly had been entirely
devoured, only a few small fragments of wings and legs
being left. ;

Experiment ).—Gave the Mantis a perfect male A.

298 Mr. G. A. K. Marshall on

horta. He tackled it at once, seizing it from above and
biting the thorax, but he quickly let go and began wiping
his mouth as before. A few minutes later be made a
second attempt with the same result. After this he ap-
peared to avoid it. I then put in a wingless Amauris
echeria and left him for some time. On my return I
found it had been entirely eaten, whereas the A. horta
was still untouched. Took the hovta out, cut off its wings
and replaced it. The Mantis eyed it with suspicion when-
ever it came near him, and felt it cautiously with his
antennz; when it came too near him, he backed away
and would not attempt to touch it. Later on I tapped
the gauze so that the horta fell close by the Mantis. He
eripped it at once, and began eating away at the underside
of the abdomen, but soon threw it down again, and would
not touch it although I gave him no other food for twenty-
four hours. After that I put ina male Belenois severina,
which he devoured readily.

Experiment «.—After starving the Mantis for twenty-
four hours I gave him a JL. chrysippus. On seeing it
fluttering he came down to it eagerly and soon caught it.
The large wings prevented him for some time from getting
at the body, and he therefore ate away almost half a hind-
wing. He then went on and ate the whole insect except
the limbs.

Experiment d.—Gave the Mantis a Papilio demodocus.
He had some difficulty in catching it at first, owing to its
size and strength, but eventually seized it from below and
devoured it. ,

Experiment ¢e.—Gave an entire female horta to my
captive Mantis. He caught it, bit the thorax and started
back with disgust, just as in the previous experiment, but
his efforts to get rid of the nasty taste were more prolonged,
For over five minutes he continued cleaning his mouth on
his fore-legs or rubbing it from side to side on the gauze.
I then put in an entire Amawris echeria, but he seemed
too scared to attempt to touch it. However, he caught it
during the night (while there was a light in the room) and
ate all the abdomen, leaving the head and thorax.

EXPERIMENTS ON Jantide at MALVERN, NATAL.

III. On March 11, 1897, I captured a large female
green Mantis [probably Polyspilota caffra (Westw.) or very

The Bionomics of South African Insects. 299

near it]. On the 12th I gave her an A. petrwa, which she
devoured entirely. On the 13th I gave her an A. serena ;
she seized it and ate a good piece out of one of the hind-
wings. She then attacked the thorax, but after a few
bites threw down the insect and began ejecting a brownish
liquid from her mouth on to a leaf, and also wiped her
mouth with her legs in the usual manner. A few moments
after I put in a male Hypolimnas misippus, which she
soon caught and ate. Later on I put in another A. serena,
but she paid no attention to it. I then put in a P.
demodocus, with the same result, so I removed them both.
On the 14th I gave her no food. On the 15th I put in
one A. encedon and one female H. misippus, but no atten-
tion was paid to either. I eventually removed encedon,
leaving misippus. Later on put in Hurytela hiarbas, and
left both in all night and through the next day, but
the Mantis would not touch them. As it was beginning
to show signs of weakness I released it.

1V. Experiment «.—March 25. Caught another female
Mantis [probably the same species as the last], and gave
her an A. cabira, which she quickly caught. She began
by eating part of the fore-wing, but as she reached the
base of the costa dropped it suddenly. A little later,
while I was not watching, she took it up again and ate
all the body except the head and anal segment. I[ then
gave her a Charaxes varanes and a P. demodocus, which
were both eaten immediately in succession.

Experiment ).—March 26. Gave the Mantis an A.
cabira. The day being cloudy and cool, she was sluggish,
and it was some time before the butterfly was caught.
She missed the first two strokes, catching it at the third
and eating it entirely.

Experiment c—March 27. Gave one A. encedon to
Mantis. It immediately flew right on to her, which
seemed to frighten her considerably, and she did not
attempt to catch it, but edged away when it approached.
This continued for a quarter of an hour, so I took the
encedon out and put in a P. deimodocus, which was soon
caught and eaten. Later on I put in a Neptis agatha ;
the Mantis seemed rather suspicious of it, but eventually
caught and ate it. I then gave her a Pentila tropicalis,
which she ate, including the whole of the two fore-wings.
I then tried her again with the same specimen I had
given her in the morning; she caught and ate it without

300 Mr. G. A. K. Marshall on

any signs of distaste. Subsequently gave her Mylothris
agathina, which was also eaten.

Experiment @.—March 28, I gave a male A. serena to
Mantis. After a few moments she caught it and ate a bit
out of the wings, but soon threw it down. The butterfly
at once walked straight back to her and was promptly
caught again, and after a single bite was again rejected.
On looking a few hours afterwards I found it had been
eaten. I then gave her a P. tropicalis and an Alena
amazoula, both of which were eaten, the latter wings
and all.

Experiment ¢.—March 29. I gave Mantis one Hurytela
hiarbas, one Pyrameis cardui, one Junonia clelia. All
were eaten.

Experiment /#—March 30. I put one male A. serena,
one P. demodocus, one N. agatha, and one P. tropicalis into
the Mantis’ cage at the same time. They were caught
and eaten in the order mentioned without any sign of
distaste. Immediately after she had finished I put ina
brilliant dark-blue moth with orange markings (Lgybolis
vaillantina), which has astrong smell. To my surprise she
completely demolished it, and then ate a second P.
demodocus.

Experiment g.—March 31. Gave Mantis a P. demodocus
in the morning, which she ate: in the afternoon gave her
one L. chrysippus, which she ate without any ado, and
immediately afterwards a female HH. misippus. I then
gave her an Acrxa natalica, which she quickly seized,
but on biting the thorax dropped it at once. For some
time she paid no attention to it, but later on tried it
again, biting a little out of the wings and then dropping
it again ; after which she had nothing more to do with it.
Subsequently put in a Papilio brasidas, which was promptly
eaten.

Experiment /4.—April 1. In the morning gave Mantis
an A, serena. She caught it, and after eating the apex of
one fore-wing threw it down, but a few seconds after she
caught it again, nibbled a bit out of the costa of fore-wing
and again threw it down. After a short interval the
butterfly walked past her, she seized it, bit at the thorax
and at once rejected it. A few moments later she made a
fourth attempt, this time eating half an antenna, but again
found the taste too much for her. I then removed the
butterfly and put in an A, encedon, but after nibbling a

The Bionomies of South African Lisects. B01

small bit out of the wing she would have nothing more to
do with it. In the afternoon I tried her with an A. cabira,
which she also refused; I removed it and put in one J.
clelia and one P. brasidas, but apparently the continued
disappointments she had undergone disheartened her, for
she would have nothing to do with either of them, but
avoided them, and only tried to escape through the glass
of the cage. About an hour after she ate the lrasidas,
but had not touched the clelia by sundown.

Experiment 7.—For three days I fed the Mantis only on
clearly edible species. On April 5, after eating two P.
demodocus I gave her L. chrysippus, which she soon caught,
but after eating a small portion of a hind-wing, she threw
it down. <A few seconds after, however, she caught and
consumed all except the wings. She then ate a male /7.
misippus, and immediately after I put in a male and
female A. cabira. The male was eaten at once; she then
caught the female and ate a piece of the wing, but threw
it down after the first bite at the thorax. The butterfly
remained for a long time at the bottom of the box feign-
ing death, so I put in another L. chrysippus. The move-
ments of the latter disturbed the cabira, which was
promptly seized by the Mantis, the abdomen being eaten
but the thorax rejected. Shortly afterwards the chin ‘Yysippus
was caught and eaten from head to tail. Next morning
as a sequel to this feast I found the Mantis in an ap-
parently half-dead condition. The abdomen was much
distended and no feces had been passed for twenty-four
hours. I therefore gave it no food whatever for two days.
On the third day it seemed better and faeces were passed
freely, but it still seemed very weak and refused food.
Next day I found that it had lost all power of gripping
with its fore-legs, so I fed it by hand on edible species.
This I continued to do for several days, but it never
properly recovered its strength, so I killed it.

V. EXPERIMENTS ON Pseudocreobotra wahlbergi, Stal.

1897.

Sept. 3. Lower Umkomaas River, Captured a male
P. wahlbergi, and gave him an Aerxa
cabira; he nibbled a bit out of the wings,
then ate the whole abdomen, but on

302

Sept.

”

>

”

12.

13.

Mr. G. A. K. Marshall on

reaching the thorax rejected it, the but-
terfly having still sufficient vitality to
flutter about.

In the morning gave him an J. safitza, which
he ate at once. In the afternoon put in
an Acrewa encedon, which he seized twice,
but on eating a bit of the wing rejected ;
however, towards evening it was eaten.

Gave him an <Acrea serena; he seemed
frightened at first and avoided it, but ate
it about an hour afterwards. Put in
another later, which remained untouched.

The serena of yesterday was left uneaten all
the morning. I therefore removed it and
put im an A. cabira, which was also
refused.

In the morning removed the cabira, and put
in another serena. As the Pseudocreohotra
had not eaten it towards evening, I gave
him an JZ. safitza in addition. He seemed
to detect the difference, watching it 1m-
mediately it was put in, and as soon as it
came within striking distance, he seized
and ate it, but still paid no attention to
the serena.

The serena remained uneaten all day, though
from its appearance it had evidently been
seized. In the afternoon put in a Neptis
agatha. The Mantis avoided it at first
just like the Acrwva, but about an hour
later I found it had been eaten.

Wet days and no Acrwas procurable.

Gave Mantis an J/. sajitza and an A. serena
at the same time; he seemed very fright-
ened of both, avoiding them, or else strik-
ing at them in order to drive them away.
Some hours later I found the safitza had
been eaten entirely and a small piece out
of the abdomen of the Acrwa, which, how-
ever, was still quite lively.

Gave Mantis two A. serena during the day,
both of which were seized at once and
eaten entirely from head to tail without
any sign of distaste.

Sept.

The Bionomies of South African Insects. 308

15.

—

pn
CO

19:

20.

bo
bo

Put in an A. serena. It was eaten after a

short interval. Later gave him an A.
encedon. At first he seemed only fright-
ened, but subsequently caught it, and after
taking a bite at the thorax threw it down
and paid no further attention to it.

Brought Pseudocreobotra from Umkomaas to

Malvern.

Put a Teracolus omphale and an A. encedon

into his box, but they were not touched
all day, owing to their imactivity and the
large size of the box. The Mantis was
also more sluggish in its movements than
in a natural state.

Caught the encedon and offered it to Mantis

in my fingers. He objected strongly at
first, but eventually took a small nibble
but would not try another bite. Offered
him the omphale in the same way, but
being suspicious he refused it also, but at
last took a bite, and, finding it all right,
ate it all, On again putting the encedon
near his mouth he only felt it with his
palpi but would not eat.

Left the same encedon in all day in hopes

that he might be compelled to eat it by
hunger ; but he did not do so.

Hncedon untouched, so removed it and put

Pseudocreobotra into a smaller box with
the specimens of A. serena, but he seemed
to take no interest in them. On holding
one of the butterflies to his mouth, he felt
it persistently with his palpi and seemed
almost as though he were trying to eat
but could not. He was certainly weaker
on his legs.

One of the encedon died during the night,

and in the course of the morning I found
the Mantis apparently eating at its head
as it lay at the bottom of the box, without
using his fore-legs, which were held out
on each side. However, on taking up the
butterfly I found he had made no impres-
sion on it. I then placed a Zerias brigittu

304 Mr. G. A. K. Marshall on

close to his mouth, and he mumbled at it

in the same manner without eating. It

therefore seemed evident that his mandi-

bles must have been paralyzed in some

way, and on examination this proved to

be the case, for they could be moved

easily with a pin backwards and forwards,

the insect clearly having no control over

them whatever. The grip of the fore-

legs though noticeably weaker than nor-

mal was not completely lost, as in the

previous experiment with a “ Charaxes-

eating ” Mantis. I am inclined to think

that the insect became at last partially

blind, both from its actions and from the

appearance of a small discoloured patch

in the left eye, a symptom which also

occurred in the “ Charaxes-eating” Mantis.

The results of Experiment VI. practically negative the
supposition that any of the above symptoms might be due

to insufficient food.

VI. EXPERIMENTS ON LEAF Mantis (Phyllocrania
insignis, Westwood).

[One of Mr. Marshall’s specimens was compared with
the type of the above-named species in the Hope Collection. |

At the Lower Umkomaas River, during September, I
kept two specimens of this insect for twelve and fourteen
days respectively without a particle of food, and neither
their vitality nor activity were in any way impaired at the
end of that period. When captured they were in their
pupal instar, and the final change took place on the 10th
and 7th days respectively, both insects casting their skins
in a normal and healthy manner in spite of their long fast.

VII. EXPERIMENTS ON “ CHARAXES-EATING ” MANTIS
(Polyspilota caffra, Westwood, or very near this species).
1897.
Sept. 23. Caught, at Malvern, Natal, a “Charaxes-
eating” Mantis (in the pupal stage), and
gave him a 7. achine and an <Acrea

The Bionomics of South African Insects. 305

serena at the same time. He was some-
what wild at first, paying no attention to
them but only trying to escape. Eventu-
ally he took the achine from my fingers
and ate it, and later caught and ate the
Acrea.

Sept. 24. Mantis ate a Belenois severina.

25. Mantis ate two <Acrewa encedon without
showing any signs of distaste.

», 26. Gave him two A. encedon, but they were not

eaten.

» 27. Mantis ate one encedon.

, 28. The second encedon dead. Put in four Par-
dopsis punctatissima, but no notice was
taken of them.

29. Mantis still refused to eat. Two P. puncta-
tissima dead. Put in one A. encedon.

, 30. One more P. punctatissima dead, and the
remaining one was three-parts eaten, the
encedon being left.

Oct. 1. Gave Mantis one P. punctatissima and one A.
serena in addition, but he made no attempt
to catch any of them, even when they
settled quite close, merely feeling towards
them with his antenne; if they came
too near he only ran away or else drove
them off by striking out straight with his
fore-legs. The discoloured patch in the
left eye made its appearance on this day,
and the sight on that side was evidently
somewhat impaired. The legs also seemed
to be weakening, and the grip of the front
pair was not so strong as in normal speci-
mens.

-, 2. No butterflies eaten, though I tried several
times. I think that he may have been
preparing for the final change of skin,
which would account for his refusal to eat.

3. Mantis attempted to perform the final ecdy-

sis during the night, but owing to his. bad
state of health could not free himself
properly from the old skin, being per-
manently deformed in-a doubled-up
TRANS. ENT. SOC. LOND, 1902.—PART III. (NOV.) 21

be)

306 Mr. G. A. K. Marshall on

attitude when I first looked at him. I
therefore killed him.

Judging from Experiment VI. the inability to change
can hardly be attributed to weakness caused by want of
food.

[In relation to the above-recorded experiments it is im-
portant to know the habits and natural food of the Mantis,
and if possible to determine the species. Mr. Marshall
kindly sent a specimen of an identical, or at any rate very
closely-allied species, together with the following notes.
—E. B. P.]

“Umkomaas Mouth, Natal ; Sept. 3, 1897.—The Malvern
species of Mantis is one of the largest out here, and I
selected it as I knew it to be almost entirely a butterfly
feeder. It frequents chiefly Acacias and their allies, and
catches the Charaaes which come to suck the gum. Un-
fortunately they are scarce at Malvern, and I could not
procure a single specimen during the winter, for I had
hoped to make the very experiment you suggest, viz.
feeding exclusively on Acreeas or L. chrysippus. However,
T caught two small Mantises on my arrival here (Umkomaas
Mouth) yesterday, but I have not as yet even seen an
Acrea.”

“ Malvern, Oct. 7, 1897.—I am not quite certain whether
the Charaxes-eating Mantis sent is specifically identical
with the one that died from Acrva diet. I thought it
was the same in the pupal stage, but the imago of the
latter has the upper wings entirely green, with a small
yellowish spot about the middle. Unfortunately I have
not been able to get one.”

[The Mantis sent (captured at Malvern, Sept. 1897)
was Polyspilota caffra (Westwood), of which the type is in
the Hope Collection, Oxford.—kK. B. P.]

VIII. ExXperIMENtTS with Pseudocreobotra wahlbergi, Stal,
female.
1897.

Sept. 26. I captured at Malvern a full-grown female
Ocellated Mantis, which ate a specimen
of Acrea encedon during the day. |

7. Gave Mantis two A. encedon. She ate the
thorax of one, rejecting the remainder.

, 28. The remaining encedon died to-day. Re-

moyed it, and put in one Aerva neobule

bo

The Bionomics of South African Insects. 307

and five Pardopsis punctatissima, but the
Mantis would not touch any of them.

Sept. 29. Mantis still refused to eat, one P. punctatis-

sima dead.

30. Three more punetatissima dead. Mantis

ate the neobule entirely.

Oct. 1. The remaining punctatissima dead. Put in
two <Acrvxa encedon and one A. serena.
The Mantis seemed more keenly alive
to their presence than usual, eagerly
following them in their flight with sharp
turns of the head lke a cat watching a
swallow. At last the serena gave her a
chance, and was immediately seized and
eaten. No more were eaten during the
day, and towards evening I put in another
A. serena.

The day being dull the three butterflies
were quite mactive. I therefore placed
the serena near the Mantis, which soon
seized it, and ate nearly the whole of one
fore-wing and part of the other; finding
this unsatisfactory she dropped it. I then
offered her an encedon, which was promptly
taken and devoured entire, and immedi-
ately afterwards the serena was eaten.

ie 3. Wet day: no Acrzeas procurable. Remain-
ing encedon dead.

4. Gave Mantis two A. encedon. She was
evidently hungry, on two occasions making
futile jumps towards the butterflies as
they fluttered past, instead of waiting
for them to come. within striking dis-
tance ; there was however a noticeable
decrease in her general vivacity. Eventu-
ally she caught both butterflies in quick
succession, and devoured them completely.
After the head of the second one was
eaten, a large drop of yellow liquid oozed
from the thorax. On tasting it she drew
back quickly and seemed uncertain
whether to go on or not, but finally put
her mouth to it and sucked it all up,

»

bo

»>

308

—6,

~

3:

2.

Mr. G. A. K. Marshall on

though it appeared to me as if it were
done under protest.

No Acreeas procurable.

Put four P. punctatissima into her box at
the same time. During the short time
I was watching I saw her catch one or
other of them no less than seven times,
but on each occasion after the first nibble
or two she threw it down with evident
disgust.

I was absent all day, but all the butterflies
had evidently been further attacked by
the Mantis, and small pieces had been
eaten out of the fore-wings, but in no
case had the bodies been damaged.

Removed all the punetatissima and put in
two A. cabira and one A. encedon, which
were consumed entirely in quick succes-
sion. The Mantis appeared to show no
decided symptoms of ill-health at present.
I was unable to complete the experiment.

IX. First EXPERIMENT WITH MANTIS. SALISBURY.

1898

March

9

9

ae

6.

~I

Caught a pair of large green Mantis in
copuld [Sphodromantis lineola, Burm. }.

Gave them one A. caldarvena, one A. halali,
and one A. neobule, but they were all
untouched.

The caldarena had been caught and dis-
carded, the thorax and one wing being
partly eaten; removed the butterflies.

Female Mantis ate the male. Put in two
caldarena and one induna; Mantis tasted
one of the former but quickly threw it
down. During the day the other two
were evidently caught and tasted, as they
were both more or less damaged about
the head and thorax.

Mantis ate one caldarena and the induna ;
remaining caldarena died from injuries.
Put in three caldarena, one of which was

partially eaten.

The Bionomics of South African Insects. 309

March 8. One other ca/darena completely eaten; the
third died, its head having been partly
eaten,

‘ 9, Put in one female ha/a/i and one male, and
one female caldarena. The former was
soon caught, but after a few bites was
rejected with evident disgust.

» LO. Male caldarena eaten completely, female
partially.

» 12. Put in male natalica, one male and one
female caldarena; the two former partially
eaten. The latter was caught three times
in quick succession, but promptly rejected
on each occasion after the first bite.

13. Put ina male halali, which the Mantis took

at once, throwing it away after eating
about half the thorax. Then gave her a
male caldarena, which was completely
eaten, so put in a second, which she
promptly caught, but threw it down after
the first bite at the thorax. She caught
it again about a minute afterwards and
started eating the apex of abdomen, but
two bites were sufficient. A third attempt
ended similarly.

, 14 Saw Mantis seize and reject the same
caldarena twice; removed it in the evening.

, 16. Putina male caldarena, which wascompletely
eaten, but a second which I gave her
immedately afterwards remained un-
touched. The Mantis began to show
distinct signs of weakness, and I observed
an opaque blackish spot in her left eye
to-day for the first time.

, 7. The male caldarena was killed to-day by a
bite on the head. Mantis began to nibble
off the end of one of her front tarsl, a
sign that her end is not far off.

, 18. Gave her a female caldarena, which was
caught several times but not eaten. She
continued to nibble at her tarsi.

» 19. Mantis oviposited during the night, but the
egg cocoon was only half-as large as usual

5
in this species [eggs proved to be infertile].

310 Mr. G. A. K. Marshall on

Gave her one male halali and two male
caldarena. ‘They were all caught in suc-
cession, but she only ate a very small
piece out of each, At times she seemed
very frightened of them, and in running
away she twice fell upon her back, when
she had some difficulty in righting herself
owing to weakness.

March 20. Put in three male caldarena, one of which
was caught and the whole of one fore-
wing and part of the thorax eaten.

21. Remaining two butterflies untouched.
Mantis had by now eaten off the ends
of al] her tarsi except the anterior and
intermediate on one side.

22. In the afternoon I found the Mantis dead
on her back.

X. SECOND EXPERIMENT WITH MANTIS. SALISBURY.
1898.

March 19. Caught a large green female Mantis of same

species as previous one [Sphodromantis
lincola, Burm.],and gave her one Junonia
cebrene and three Terias senegalensis, all
of which she ate. She also ate the
following :—
, 20. One TZ. senegalensis, one T. brigitta, one
Belenois severina.

, 21. One J. cebrenc, one Catochrysops ostris.

22. Two Alxna amazoula, without showing any

signs of distaste.

23. - One Spindasis natalensis, two J. cebrene.

, 24. One 7’ senegalensis, two Myrina ficedula.

,» 25. The Mantis escaped from the box this
morning, and [ did not find her till
5.30 p.m., when she was busy ovipositing
on the side of a book. She had then
laid about a third of her eggs, and did
not stop laying till 8.30 p.m:

Mantis ate one Parosmodes icteria, and two
Hesperia spio. She seemed very hungry,
following the butterflies about instead of
waiting for them to come within striking
distance.

bo
SD

d)

The bionomics of South African Insects. 311

March 27. She ate one Catopsilia florella ; 1 then put

”

XI. THIRD

1898.
April 3.

=e

in a Lelenois mesentina, and she be-
came much excited, running about after
it, and making several futile snatches at it
on the wing. At last she gave a vigorous
stroke, and. missing the butterfly caught
the gauze with which the box was
covered. Imagining apparently that she
had caught her prey, she began trying to
eat the gauze, in spite of my attempts to

drive her away, for fully two or three
minutes. At last she desisted and soon
caught and devoured the butterfly,
eating a b. severina and Axiocerces harpax
immediately afterwards.

One Precis sesanvus and one B. mesentina.

One Hamainumida dedalus and one B.
severina.

One Pyrameis cardui and one B. mesen-
tind.

One J. cebrene and two B. severina.

One J. cebrene and one C. florella.

Two C. florella and one P. cardut.

Mantis escaped. She was fully as vigorous
and healthy on the last day as when first
caught.

EXPERIMENT WITH MANTIS. SALISBURY.

Captured a female Mantis closely allied to
those of preceding experiments, being of
same size and colouring, but having a
much broader thorax and the mouth
pink. This insect I submitted to a
purely distasteful diet, combined with
periods of starvation, as follows :—

Three Acrea caldarena eaten.

Two s * i
Two AXING )
One ad caldarcna ,,
Two Limnas chrysippus ,,
One ” ” ”
One ” ” » +

One ” ” ”

April 16.
20.

?

June 4.
Db

>

Mr. G. A. K. Marshall on

One Limnas chrysippus eaten.

One x eS . ; then left
her a month entirely without food, which
however did not seem to affect the
health or vitality in any way.

One Acrvxa axina eaten.

One L. chrysippus ,,

One ,, .

Two ,, P , this insect never
once exhibited the least signs of distaste
for any of the butterflies, and devoured
them all with avidity, showing a marked
contrast to the Mantis first experimented
with (IX.), which throughout exhibited
an intense dislike to the Acreeas, and
evidently ate them from sheer hunger.

The close proximity of these two kinds of
Mantis suggests the idea that they might
possibly be seasonal forms of one species
in which the winter form has adapted
itself to an Acrwa diet, owing to the com-
parative scarcity of other butterflies at
that season. In spite of its diet and long
fast, this Mantis was still fully as vigorous
and healthy as when first captured.

Ate one L. chrysippus.

Gave it two Acrva axina ; it tasted both of
them several times, but in every case at
once discarded them with evident disgust.

Gave ittwo more A. axina with precisely the
same result.

Ate one L. chrysippus.

> » 3) ”)
” two ” )

Put in one ZL. chrysippus. The Mantis
showed its normal eagerness, and followed
it about for some time, finally attempting
to seize it, but failed. The vigorous
fluttermgs of the butterfly seemed to
frighten the, Mantis, which ran away
from it and made no further attempt to
catch it.

Put in two more chrysippus, but Mantis

The Bionomics of South African Insects. 313

was still scared and would not go near
them.

Sept. 8. Mantis died. There were no signs of ill-
health as in former experiments, and the
characteristic blotch in the eye was
absent. Death was probably due either
to hunger or natural causes. I only
wonder at its lasting so long, considering
its long fast and unhealthy food.

[The above experiments upon Mantide of four different

genera are summarized as follows :—

Mantis [L., in the Karkloof. Evident intense dislike,
after trial, of Acrwa horta.

Mantis IL., male, in the Karkloof. Evident dislike of
A. horta, although one specimen out of three was almost
entirely eaten. Two A.echeria were eaten, and one partially.
One L. chrysippus, one B. severina, and one P. demodocus
were eaten.

Mantis IIL, female, probably Polyspilota caffra, at
Malvern. Ate one A. petvwa, but rejected A. serena after
trial. Ate one male 1. imisippus, but after this refused all
butterflies, and exhibited signs of weakness.

Mantis IV., female, probably the same species, at
Malvern. Ate Papilios and Nymphaline freely, including
the probably aposematic genus Neptis, and the probably
aposematic Lyczenid genera Alena and Pentila, and Pierine
genus Mylothris. Ate LZ. chrysippus with hesitation, and
partially in one case, freely in two cases. Hence the Mantis
appeared to be a very general feeder on all butterflies
except the genus Acrwa, the species of which (cabira,
serena, encedon) were rarely eaten until after one or more
trials, and were sometimes finally refused. Natalica was
only offered once, and rejected after trial. Itis interesting
to note that immediately after trial of three different
Acreas, the Mantis refused species which she freely ate
at other times. The final weakness without power of
recovery was a probable result of the diet.

Mantis V., male, Pseudocreobotra wahlbergi, on the Lower
Umkomaas River. Ate Jf safitza, 7. omphale, and Neptis
agatha freely, the first-named on three occasions.
Acrexa cabira, refused twice ; encedon, refused twice after
trial and accepted once; serena, refused on five occasions,
eaten on four. It is probable that the weakness and loss
of sight was due to the Acrwxa diet.

314 Mr. G. A. K. Marshall on

Mantis VI., Phyllocrania insignis, on the Lower Umko-
maas River. The evidence that starvation for twelve and
fourteen days respectively does not produce the symptoms
observed in Experiments IIL, IV., V., VIL, and IX.

Mantis VII, male, in pupal stage, probably Poly-
spilota caffra, at Malvern. The Pierines 7’. achine and
B. severina freely eaten. Of the Acreeas, two encedon eaten
apparently freely, and one after an interval, others re-
fused : one serena eaten after atime: of four punctatissima
only one partly eaten. The Mantis then refused all food,
became weak, and one eye was affected. He was unable
to throw off the pupal skin properly. In Experiment VI.
two individuals of another species performed this change
of skin after ten and seven days of starvation.

Mantis VIII., female, Pseudocreobotra wahlbergi, at
Malvern. Only offered Acreeas. Nine punctatissima always
refused with or without trial ; encedon eaten freely several
times, refused once, and partly eaten once; two cabira eaten
freely ; serena eaten freely or after trial; neobule eaten
after two days’ interval.

In spite of this diet the Mantis remained apparently
healthy, September 26 to October 9, 1897, when the
experiment came to an end.

Mantis IX., female, Sphodromantis lincola, at Salisbury.
It was intended to offer this individual a purely Acrva
diet, but she ate her mate on the third day after their
capture im copuld. She was chiefly fed upon <Acrea
caldarena, which she ate sometimes freely, sometimes
after an interval and after trials: at other times she
refused it with or without trial ; one induna was eaten after
a day’s interval; one neobule was untouched; one natalica
was partly eaten; two dalali were rejected after trial, one
without. After a fortnight of this diet the Mantis became
weak, and her left eye was affected : a day later she began
to nibble off the end of one of her fore tarsi: two days
later she oviposited, but the egg cocoon was only half its
usual size (eggs infertile). After the first signs of weak-
ness the Mantis ate only a small part of three Acreeas out of
eight offered her during six days. She continued to nibble
at her tarsi, lost power over her movements, and died
after twenty days of captivity.

Mantis X., female, Sphodromantis lineola, Salisbury.
This individual was the subject of a control experiment,
being fed solely upon several species of the following groups

The Bionomics of South African Tnsects. ales

—Nymphaline, Prine, Lycenide, and Hesperide. The
only species with marked aposematic colouring and habits
was the Lycsenid Alena amazoula, and of this only two
specimens were offered, both being eaten freely. After
fifteen days of this diet the Mantis escaped : she was then
as healthy and vigorous as when first captured.

Mantis XI., female, species resembling Sphodromantis
lineola, Salisbury. Fed solely upon Acres caldarena and
A. axina, and Limnas chrysippus, with long periods of
starvation, two of them a month in duration. Seventeen
chrysippus, six caldarena,and three axina were eaten without
any signs of distaste, while four axzina were discarded after
tasting several times. The Mantis was captured on April
3, 1898, and refused food on September 4, dying on Sep-
tember 8 without signs of ill-health or blindness. Mr.
Marshall suggests that the species may be a winter form
(possibly of S. lineola) specially adapted to eat Acreeas
when other butterflies are scarce.—E. B. P.]

CONCLUSIONS FROM EXPERIMENTS ON MANTID.
(Ex: E)

Certain conclusions stand out very clearly, while others
are suggested as probable. These voracious insects did
not show any dislike of butterflies outside the Danaine
_ and Acrvine. The undoubtedly aposematic Pierine genus
Mylothris was freely eaten, and so were the following
genera with probable warning colours, movements, and
attitudes—WNeptis, Alena, Pentila, and the moth Lgybolis
vaillantina. Even the Danaine were generally eaten with-
out hesitation (II., IV. XL), and never rejected alto-
gether. In marked contrast was the behaviour of
Mantidz towards Acrxinew, which were constantly refused,
and often eaten only after one or more trials and long
intervals of time. When the Acraas were eaten freely
and without hesitation there is reason for suspecting
exceptional hunger. The summary of experiments

5
shows very clearly that “ Pardopsis appears to be consider-
ably more distasteful. . . . than the general run of

Acreas” (G. A. K. M., October 7, 1897, Malvern). There
were also less marked differences in the degree of dislike
shown towards other species; thus avina was less freely
eaten than caldarena (XI1.); caldarena appeared to be
eaten more freely than halali, neobule, indwna, and natalica

316 Mr. G. A. K. Marsball on

(IX.), although the number offered of these latter was
insufficient to warrant a certain conclusion; cabira was
rejected while a considerable proportion of the encedon
and serena were accepted (V.); horta evidently possesses
a high degree of unpalatability to Mantide (L., IL).

Mr. Marshall's evidence, by far the most important
collected in the case of the Mantidex, is in entire accord
with the few observations which had been previously
recorded. Thus the late Mr. de Nicéville found that
Acrea viele was the only butterfly refused by all the
species of Mantis with which he experimented in the
East (“Butterflies of India, Burmah, and Ceylon,” vol. 1,
pt. 1, p. 318). Colonel J. W. Yerbury informs me that he
watched the Mantis Gongylus gongyloides hanging from
the drooping lavender flowers of a species of Duranta at
Trinkomali (1890-91), and capturing the butterflies
which were attracted by the bloom. The insect hung by
its four posterior legs, with head thrown back and preda-
ceous legs held ready for striking. He saw it capture
and eat Delias ewcharis on several occasions, and also
Belenois mesentina and the Hesperid Hasora alexis (Fab.).
Colonel C. T. Bingham has also given me a male
specimen of the Harpagid Mantis, Creobotra urbana (Fab.),
found by him on a Lantana bush actually eating Delias
descombest (Boisd.). This observation was made in the
North Shan States, Upper Burma, on October 9, 1900.
The fact that two species of De/ias were thus freely eaten
compares 1n an interesting manner with the acceptance of
Mylothris by the African species of Mantis. We may safely
conclude that outside the Acrwinw, and doubtfully the
Danaine, Mantide devour butterflies very freely, the
species with warning colours as well as the others, and
that they are far more undiscriminating than the majority
of vertebrate insect-eaters. Thus Mr. F. Finn found Delias
eucharis to be one of the most distasteful of all butterflies
to many species of Indian birds (“Journ. Asiat. Soc. Beng.,”
vol, Ixvii, Pl. ii, No. 4, 1897, p. 667). Mr. Finn also
found in East Africa that a moth of the genus Hyybolis
(EZ. vaillantina) was refused by a Chameeleon and a Gecko
(‘Natural Science,” vol 1, No. 10, Dec. 1892, p. 747). It
is of deep interest to find such marked differences between
the preferences of the various groups of insect-eating
animals.

In addition to the observations recorded above, Dr.

The Bionomics of South African Insects. 317

David Sharp, F.R.S., quotes Mr, F. Muir concerning the
food of Ldolum diabolicum (Sauss.) at Mozambique :— Its
food seemed to consist of flies, Zimnas chrysippus being
rejected, even when hungry, and other butterflies only
taken for lack of other food” (Proc. Cambr. Phil. Soc.,
vol. x, pt. i, p. 175). Mr. Edward Barlow (Proce. Asiat.
Soc. Bengal, Dec. 1894) states that Hierodula bipapilla
(Serv.), kept in captivity at Calcutta, ate ordinary flies
(Musca sp.) with avidity, but attacked with great re-
luctance the common large green blowfly (Lucilia sp.),
only eating them when they could get nothing else. Two
bugs, Cyclopetia sp. and Physomerus sp., offered when the
Mantis was very hungry were never eaten, although often
killed. After tasting the former, the Mantis wiped its
mouth against its right fore-leg several times. This last
observation is the only record | have found of Hemiptera
offered as food to Mantidx.

The question arises as to whether the preferences
exhibited by MJantidxe in captivity are the same as those
which exist in the wild state. A Mantis is probably less
affected in this respect by confinement than a vertebrate
animal; but the same general criticism will probably hold
in both cases—that while the rejection of an insect by a
not over-fed insectivorous animal in captivity is evidence
of unpalatability or dislike, its acceptance is not sufficient
evidence of appreciation or that it constitutes an element
of the normal diet. An insect may be eaten readily in
captivity which would be rejected or only eaten under the
stress of hunger in the wild state; for it is generally quite
impossible to supply an animal under artificial conditions
with the variety and often the quantity of insects which
it would catch for itself. In this respect a large Mantis
can be kept in a more normal condition than an insecti-
vorous vertebrate, because of the much larger amount of
food required by the latter; although the young Mantis
would offer great difficulties to the breeder, because of the
vast numbers of very minute insects which it would
require. But Mr. Marshall’s experiments yielded plenty
of evidence of the positive refusal and acceptance, as it
were under protest, of Acrwinex, so that there can be no
doubt of their distastefulness to this class of enemy,
although acceptance might under the circumstances
have not been convincing proof of their palatability. It
is however in every way satisfactory to obtain evidence

318 Mr. G. A. K. Marshall on

from the behaviour of AMantide in the wild state, and
such as we do possess entirely confirms the conclusions to
be drawn from Mr. Marshall’s experiments. In the first
place we have the following observation of his own, made
in the Karkloof, Natal, in February 1897 :—

“Saw a Mantis catch a male horta on a flower in the
veldt. It began eating at the base of the abdomen, which
it consumed entirely, and then started on the thorax, of
which it only ate a very little, and then threw it away.”

This observation corresponds almost precisely with
many made upon the captive insects. Mr. Roland Trimen
also says that he never found the wings of Danais or Acrwa
among the fragments of butterflies which sprinkle the
eround below the feeding-place of a large Mantis, although
he is careful to add that he could not be sure that these
butterflies visited the exudations of Acacia sap, round
which the predaceous msects secure a plentiful supply of
food (Linn. Soc. Trans., vol. xxvi, 1870, p. 500). It has
already been pointed out that Colonel Yerbury’s and
Colonel Bingham’s observations upon Mantidx in the wild
state are entirely confirmatory of Mr. Marshall’s observa-
tions of them in captivity, as regards the food which
appears to be freely provided by certain Pierine genera
refused or disliked by other insect-eating animals.

Another question of deep interest raised by Mr.
Marshall’s experiments on Mantide is the inquiry how
far the species which they reject or eat only sparingly 1s
unwholesome or even poisonous to them. There is
strong a priori probability for the view that the
preferential appetite of such a form as a Mantis is merely
the strong instinctive tendency to eat the food which best
suits its organization and reject that which suits it least.
We should expect therefore that such marked disinclina-
tion to eat Acrweas as we observe in Jlantide indicates,
not distaste or unpalatability in an anthropomorphic sense,
but merely that Acreeas are unwholesome to Mantide,
The evidence requires to be sifted in detail.

In Experiment III. the signs of weakness seem to be a
too-excessive result of the single Acrva, and portion of
another, which were eaten. At the same time generic and
specific differences are almost certainly of great import-
ance, and it must be remembered that ITI., [V., and VII.
belonged to probably the same species, and all exhibited
weakness after an Acrva diet, resulting in the death of

The Bionomics of South African Insects. 319

IV., the deformity of VII., while III. was released.
Experiments V. and VIII. were also upon the same
species of Mantis. The first, a male, became weak and
probably blind after eating a few Acraeas; the second, a
female, remained apparently healthy after an exclusively
Acreeine diet for fourteen days. It is very unfortunate that
this latter experiment could not be continued. It is,
however, clear that in the case of this species and sex a
purely Acrzine diet for fourteen days is not necessarily
unwholesome. Experiments IX., X., and XI. were upon
species which were the same, or nearly the same, and all
females. The first died after an Acrva diet for twenty
days, the second was perfectly healthy after a mixed butter-
fly diet without Acrwiney and Danaine for fifteen days,
while the third lived healthily from April 3 to September 8
upon Acraas and Limnas chrysippus. The latter seems to
be an insuperable difficulty, but it must be remembered (1)
that chrysippus was given in especially large numbers, and
there is no evidence that Danaine are much rejected by
Mantidx, (2) that the Mantis may have recovered from
the effect of the Acreas during the long fasts, (3) that the
Acrexa chiefly made use of, A. caldarena, may be less
unwholesome than the majority of the group.

More experiments are greatly wanted, but Mr. Marshall’s
observations render it highly probable that Acraeas are
unwholesome to Mantide. The definiteness of the
symptoms exhibited, and especially the effect upon the
eye, constitute not unimportant evidence in support of
this conclusion. The appearance of an opaque blotch in
the left eye of three of the Mantises (V., VII., 1X.) suggests
further experiments in order to test whether we have to
do with mere coimcidence or a phenomenon of deeper
significance,

Mr. Marshall’s conclusions from his experiments were
written upon the results obtained with spiders as well as
Mantises, and will be found at the end of the section upon
the former (p. 322).

4. I.XPERIMENTS ON SPIDERS IN THE KARKLOOF.
(G. A. K. M.) Natal, February 1897.
[The Rev. O. Pickard-Cambridge, F.R.S., informs me
that the species made use of was the common and widely-

distributed Epeirid Nephilengys malabarensis, Walck.—
He Bs P.]

320 Mr. G. A. K. Marshall on

The spiders experimented on were all of one species
with very large females and minute males. Their webs
were all round the verandah, where they were strictly
preserved by Mr. Ball. I never saw the species in the
bush.

1. Gave a spider (A) a specimen of Acrea horta
(entire); she ran down and bit the thorax, then pulled it
out of the web and dropped it. At the same time gave
A. horta with its wings cut off to another spider (C) of the
same species, which ate it without hesitation.

2. Gave horta without wings to four spiders (A, B, D,
and EK), and also one in which half the wings had been
cut off to C. All were eaten readily.

3. Gave entire males of horta to A and B, and both
were at once rejected.

4. Wingless specimens ‘of L. chrysippus given to A and
C were at once thrown out of their webs.

5. The following wingless specimens were given to this
species :—Papilio brasidas to A, P. ophidicephalus to B,
P. euphranor to C, P. ophidicephalus to D, Eurytela hiarbas
to E, and P. lywus to F. All of them were promptly
eaten.

6. Caught a female horta, rubbed all the colour off its
wings, leaving them entire, and gave it to A, which after
careful examination wrapped it up and carried it off to eat.

7. Gave A a perfect male horta; she ran down and bit
it in the thorax and ejected it from the web. I then
rubbed all the colour off the wings and returned it. The
spider approached it carefully feeling round with her palpi,
and again cut the butterfly loose. I then gave it to B,
which also refused it. I then cut the wings off and gave
it to B again, with the same result. Finally I gave it to
A again, but she pulled it out of the web by the abdomen
and dropped it.

8. Gave wingless specimens of Papilio demodocus to B
and C. Both were eaten.

9. Gave a perfect female horta to D, which bit it several
times, being seemingly rather doubtful about it, but
eventually wrapped it up and carried it off to her chamber.
After a short time she threw it down, the butterfly being
still alive.

10. On two occasions saw dead specimens of A. horta in
spiders’ webs in the bush. They were both wrapped up,
but evidently had not been sucked.

The Bionomies of South African Insects. 321

11. Cut off the wings of three male horta and gave
them to A, B, and C, but they were all rejected. Gave
one of the same specimens to D, which cee it off to
eat, and was still sucking it when observed two and a
half hours afterwards.

12. Gave a wingless Amauris echeria to A, which came
down very cautiously and bit it in the thorax as usual.
Its taste was evidently unpleasant, as in extricating the
butterfly from the web it carefully abstained from biting
any part of the body. I then put the same specimen in
B’s web; she ran down at once and tackled it. After
giving it a few bites she paused as though in doubt, then,
as if thinking it was worth trying, she wrapped it up and
drew it up after her to her chamber. She was clearly still
doubtful, as she remained several minutes without attempt-
ing to touch it. She then sucked it for a few seconds, but
soon let it drop. Gave the same specimen to D, and it
was rejected. Gave another wingless specimen to GC,
which also was rejected.

13. Gave entire specimens of Zerias brigitta to B, C,
and D, and also female Nepheronia argia (agathina form)
to A. All were eaten readily. Subsequently gave entire
P. sesamus (natalensis form) to C, which was also eaten.

14, Gave a perfect male horta to D. She ran down, bit
it in the thorax as usual, wrapped it up and carried it off,
She then remained some minutes without attempting to
touch it, then after sucking it for a few seconds she threw
it sae: (Compare Experiments Grand alas)

15. Gave a wingless Acrva violarwm to spiders B, C,
and D, in succession. It was promptly ejected by each of
them.

16. Gave entire specimens of A. horta to spiders A, B,
C,and D. The two former ejected theirs at once; C cut
hers loose from the web, and was holding it in her jaws
preparatory to throwing it away, when she seemed sud-
denly to change her mind and ran up to her chamber with
it, without however enshrouding it with web. She
remained with it in her mouth for about half a minute,
and then threw it down, D took no notice whatever of
the insect in her web.

17. Gave male Acrwa buxtoni to A, and female Nephe-
ronia argu (agathina form) to C. Both were eaten.

18. Gave Pontia hellica to B, Papilio demodocus to C, and

TRANS. ENT. SOC. LOND. 1902.—PART IIL (NOV.) 22

on, Mr. G. A. K. Marshall on

Planema esebria to D. All of them were eaten, though
D seemed a little suspicious at first.

19. Gave Byblia goetzius-acheloia to A, which ate it
readily, although she was a long time before coming down
to see what it was.

20. Gave wingless specimens of A. horta to spiders
A, B, C, and D (six days since last were given—Experi-
ment 16). The first three promptly ejected them, but D
wrapped hers up and carried it off. She did not seem
very enthusiastic about it however, for she turned it over
and over a good many times, giving it a bite here and
there, and then left it alone for some time. This pro-
cedure she repeated several times, and then threw it away.

21. Gave entire males of Acrxa serena-buxtoni to
spiders A, C, and D, all of which were eaten. It should
be noted that experiments with this species are un-
satisfactory, owing to the fact that when captured it is able
voluntarily to exude from the thorax its bitter yellow
juice, and therefore when given to spiders it has lost much
of its nauseous quality, and would be less distasteful
than if caught by them direct.

5, RESULTS OF EXPERIMENTS ON SPIDERS AND THE
EARLIER EXPERIMENTS ON MANTIDA: ONE PROB-
ABLE MEANING OF TENACITY OF LIFE IN DISTASTEFUL
Inseets. (G. A. K MM.)

Malvern, Natal; February 21, 1897.—The danger of
arguing from insufficient materials was clearly shown me
in my first few experiments on spiders with A. horta
(Experiments 1, 2, 3, and 6). When I had got thus far I
felt sure I had got proofs of the appreciation of warning
colours by the spiders. For in these experiments they ate
every specimen without wings and refused all those with
them except one which had the colour rubbed off. Yet
subsequent experiments have convinced me that both
spiders and Mantises have no appreciation of warning
colours; and this fact has elucidated another which often
puzzled me, I mean the apparently constant correlation
between distastefulness and tenacity of life in Lepidoptera.
At first sight it would seem that tenacity of life or the
power to recover after severe injury would be useful to
any species in the struggle for existence. But a little
thought showed me that this power would be of no use to
edible species, as if once caught by insectivorous animals

The Bionomics of South African Insects. 323

they are not likely to be released. But in the case of
inedible species it is different. For if my surmise is true,
that insectivorous invertebrates are not capable of appreci-
ating warning colours but have to taste all their captives
before being able to tell whether they are edible or not
(which I think is clear from my experiments), then tenacity
of life (as a protective agency) will be as useful an
acquisition against invertebrates as warning coloration is
against vertebrates, and come into play when the latter is
useless. Of course tenacity is of use against the experi-
mental tasting of young birds, lizards ete., but this does
not seem to me to be a sufficiently cogent factor to
develop the power to such a high pitch. For if the
insects had only these enemies to contend against, even
supposing every specimen experimentally tasted died from
its Injuries, the protection afforded by the warning colours
would still be ample. Indeed I believe that the tough-
ness of inedible insects has been primarily developed to
counteract the injuries from invertebrate foes (which are
incapable of reasoning as to whether an insect is edible or
not), and that therein lies its chief utility, though it may
prove useful incidentally in other cases,

A. buxtoni appears to have more juice in proportion
than horta, and I regard it as a more highly-developed
species, from a distasteful point of view, in that it can
exude juice at will from its thorax, and thus show its
nauseous qualities without necessarily having to be injured
like horta. When squeezed the juice often oozes from
the ends of the antennze and all the nervures of the
fore-wings when they are cut. But, as I have pointed
out (vide Expt. 21), the results of experiments with it
are unreliable.

The treatment of A. horta by the spiders would almost
give some colour to your suggestion that the inedibility of
species may be due to unpleasant internal effects rather
than the mere taste, for B ate one specimen and A, C, and
D two each before they seemed to become aware that it
was not good to eat, from which I should conclude either
that the unpleasant effects are subsequent to eating or
that their sense of taste is not sufficiently acute to recog-
nize a nasty flavour at once. But the latter conclusion
appears to be invalidated by their prompt rejection of Z.
chrysippus and A. echeria. Anyway their selection seems
to show that there are grades of unpleasantness, and, as I

324 Mr. G. A. K. Marshall on

expected, those species in which the sexes are alike are
least edible. Thus I expect: to find that A. anemosa, A.
cabira, and P. aganice will prove more distasteful than
their allies in which the sexes differ markedly. However,
IT must admit that in the case of the Mantis this was not
so, and its persistent preference for echeria rather than
horta (on three occasions) is very curious and interesting.
Its dislike of the taste of horta was most marked, and yet
it did not appear to distinguish it by sight. In Experiment
Il. ¢(p. 298) the Mantis certainly avoided Horta after its first
taste, but it showed equal fear of echeria, which it after-
wards ate, and I presume could not distinguish between
them. But it is clear that it was unable to retain long
the impression which connected a butterfly with an un-
pleasant taste. The prompt acceptance of A. serena by
the spiders appears to support my view that the bright red
colour conveys no significance to them, although they find
the red horta distasteful. I was surprised at their unani-
mous refusal of the single specimen of A. violarwm I was
able to procure, as I had thought it would certainly be
more edible than horta, and I am almost inclined to believe
that it was rejected under a misapprehension.

I had an idea that perhaps Papilio demodocus was dis-
tasteful, which was suggested by its wide range and general
abundance; and that if this were so 1t would be probable
P. ophidicephalus, P. ewphranor, P. constantinus, ete., might
obtain protection from their strong resemblance to it on
the wing. But my experiments seem to negative the
idea as far as invertebrate foes are concerned.

Again, Terias has always puzzled me. They are so
widely distributed and always plentiful; moreover, their
flight is weak and their contrasting colours of black and
yellow are most conspicuous on the open veldt, which they
frequent ; indeed, far more so than the colours of the
females of A. violarwm and nohara. Yet they do not seem
to be protected, although some of the tropical Durbanias
and Teriomime appear to mimic them.

Malvern, Natal; Oct. 7, 1897—The experiments on
the effects of an Acreva diet, so far as they go, seem to
lend some measure of support to your view as to the un-
wholesome qualities of Acrva, though many more experi-
ments will be necessary to establish it. If I could only
get the material I should like to experiment contempora-
neously on a number of the same species, starving one,

The Bionomics of South African Insects, 325

giving one or two only edible butterflies, and confining
the remainder to a single species of Acrvwa or Amawris
each, But at present I find it not only difficult to get
hold of a Mantis, but it is even quite a job to catch sufficient
Acreas to continue the experiments. I have not seen a
single specimen of A. pefrwa for over six weeks, though
ee it should be swarming at this time of year.

[Mr. Marshall subsequently carried out a part of the
programme which he here suggests. See Experiments
IX., X., XI. on Mantide.]

When the experiments on spiders are compared with
those on Mantises the conclusion is suggested that Acre-
ine were distasteful to both, and only eaten under the
stress of hunger, while Danainw were far less distasteful
to the Mantises than to the spiders. To the latter they
appeared to be at least as distasteful as the Acraine.
Such differences in the susceptibilities of insect-eaters help
us to understand the puzzling case of Zerias, and the
Ethiopian ae genera which appear undoubtedly to
mimie it, and permit us still to look on Papilio demodocus as
a possible model. We see that the various insectivorous
groups have different tastes, and within each group we
must expect to find individual species adapted to feed
largely on insects which are as a rule rejected by the
other members of the same group.

In one respect spiders are extremely satisfactory for the
purpose of these experiments. They remain throughout
wild animals with their natural sources of food still avail-
able. The same may be the case with Mantides, as in
the Gongylus watched day after day by Col. Yerbury at
Trinkomali (see p. 316).

The late Thomas Belt (“ Naturalist in Nicaragua,’ Lon-
don, 1888, p. 317) states that a “spider that frequented
flowers seemed to be fond of” the Heliconide (including
Ithomiine), although a large species of Nephila used to
drop them out of its web when he put them into it.

Dr. A. G. Butler (Trans. Ent. Soc. Lond., 1869, p. 27)
long ago showed that the larvee of Abraxas grossular iata and
Halia wavaria were not eaten by the spiders he employed
—Hpeira diadema (the name given in the paper is Hreiba
diadema) and Lycosa species (2). In the former case they
were cut out of the web, and in the latter seized and
earried down the “dark silken funnel,’ but then relin-
quished apparently uninjured. Professor Platean (Mém.

326 Mr. G. A. K. Marshall on

de la Soc. Zool. de France, tome vii, 1894, p. 375)
gives reasons for doubting whether the latter spider
belonged to the genus named by Dr. Butler, and he
supposes from the described form of “funnel” that
the “ Zycosa,” which does not make a web, was in
reality Agelena labyrinthica. Professor Plateau’s fresh
experiments (lc. § 8) on spiders, however, entirely con-
firm the results obtained by Dr. Butler so far as the larva
of Abraxas is concerned. An Amaurobius ferox, which had
spun a characteristic web in captivity, paid no attention
during two days to three half-grown larvee moving about
and entangled in the web. From the Professor’s long
experience of spiders in captivity he is confident that it
was not afraid, and he thinks it probable that it refrained
from attack because the vibration of the web was not like
that caused by its habitual prey. It would have been
more satisfactory if this interpretation had been tested by
the offer of a few inconspicuous larvee. In the next ex-
periment four larvee were thrown into a large web spun by
a female Tegenaria domestica over the roof-light of a barn
in such a position that the observer could watch everything
without in the least disturbing the spider. The following
is a translation of Professor Plateau’s account :—“ At the
moment of the fall of the larvee into the web, the Tegenaria
.... rushes at one of them and bites it, or at least makes
two successive attempts to bite it. The spider then leaves
the first victim and attacks a second, which she also tries
to bite, but the skin of the larvze being too tough and the
caterpillars rolled up and pretending to be dead, she retires
slowly, a certain proof to those who know the habits of
these animals that she does not feel any fear.’ After-
wards, when the caterpillar began to move the web, the
spider, “having learnt the uselessness of her attempts to
bite, neglects them entirely and remains in her tube.”

Concerning this and the previous interpretation, it is
necessary to remark that no signs of fear were to be ex-
pected upon the hypothesis that the spider recognized
that the larva was uneatable; while the suggestion that
the skin was too tough to be penetrated seems to be very
improbable.

An experiment of the same kind was then made with a
female Tegenaria, which spun a web in confinement. After
keeping the spider for three days without food, two cater-
pillars were thrown into the web. The spider rushed

The Bionomics of South African Insects. 327

towards one of them but retreated again, “ recognizing that
it had been disturbed by a creature such as had never
before fallen into the web.” The same thing was again
repeated on four occasions, the spider never actually
attacking a caterpillar. Agelena labyrinthica was then
tried in the same way, but would not leave its retreat ;
when, however, an earwig was substituted for the larva, it
was instantly seized and devoured.

On the other hand, Professor Plateau has produced evi-
dence that the imago of the Abraxas is freely eaten by
Tegenaria domestica. Nearly every day for some weeks
he placed this moth, sometimes on one, sometimes on
another of three or four webs of this spider spun in a little
tool-house in his garden, They were always seized and
carried off.

Agelena labyrinthica, on the other hand, killed the moth
but abandoned it after some attempts to suck its juices.
Professor Plateau suggests that the spider found the prey
too large for it, an interpretation which might have been
advanced had the attempt to capture and kill been unsuc-
cessful; but, as the case stands, serves to show that the
author is willing to accept any explanation however im-
probable rather than the obvious one that there was
something in the taste or smell of the moth which pre-
vented the spider from devouring it.

A half-grown female Hpeira diadema devoured the moth
with avidity. It is to be hoped that this experiment will
be repeated many times, as in the case of the Tegenaria ;
and in all such researches comparison should be constantly
made with the behaviour of the spiders towards many other
kinds of insects.

It is quite probable from the experiments of Mr. Marshall
and Professor Plateau, and the observations of the late Mr.
T. Belt, that certain species of spiders, together with Man-
tides and other predaceous insects, will be found to be
among the chief, perhaps the chief, non-parasitic enemies of
aposematic insects.

Colonel J. W. Yerbury has kindly searched his notes for
any references to the attacks of spiders on butterflies. He
writes, “I can find very few references to the relations of
these two animals to each other in my old notes. The
following are two of them :—

““Aden, ? date-—A. large green flower-haunting spider
resting on a dried-up plant was preying on a female

328 Mr. G. A. K. Marshall on

Teracolus vi. The specimen was almost the first female
of the species which I obtained. The individuals of this
butterfly roosted regularly on the stalks of the plant in
question, their under-sides being of about the same tint
as the dried-up leaves and stems.

“<« RFutehpore Sikri, near Agra, May 1877.—Spiders lay
in wait for the Pierine Belenois mesentina, on the flowers
of a caper (Capparis aphylla). On this occasion the spiders
took a very heavy toll of the butterflies.’ ” *

[Mr. C. J. M. Gordon has sent to the Hope Department
a male specimen of Acrwa bonasia, which he found on
January 8, 1902, at Old Calabar, in the grasp of a flower-
haunting spider (Zhomisus, sp.). The falces of the arachnid
were fixed in the butterfly’s thorax, and the insect was
nearly dead.—k. B. P.]

6. THe ATTACKS OF PREDACEOUS INSECTS OTHER THAN
MANTIDA UPON CONSPICUOUS SPECIALLY-DEFENDED
LEPIDOPTERA, ETC, (EK. B. P.)

H. W. Bates, in the historic paper which contained the
first suggestion of the theory of Protective Muimicry
(Trans. Linn. Soc., vol. xxiii, 1862, p. 495), states concerning
the attacks of predaceous insects: “I never saw the
flocks of slow-flying /7eliconide [in the writings of Bates
and Belt upon Mimicry, the /eliconide always include
the Ithomiine or Neotropine, then called the Danaoid
Heliconidx| in the woods persecuted by birds or Dragon-
flies, to which they would have been an easy prey ; nor,
when at rest on leaves, did they appear to be molested by
Lizards or the predaceous Flies of the family Aszlide,
which were very often seen pouncing on Butterflies of
other families” (p. 510).

There is, however, good reason for believing that such
attacks are not rarely made, and that predaceous insects
are important enemies of aposematic butterflies.

In the following three sub-sections of this paper I have
brought together some slight evidence in support of this
conclusion. Far more requires to be done, and it is hoped
that the attention which is here directed to the inquiry

* Shortly after I had made the observation I came across a
reference to this habit of the spiders at the very same place, but I
cannot now recall the name of the publication.—J. W. Y.

The Bionomics of South African Insects. 329

may bring the subject to the notice of naturalists, especi-
ally in the parts of the world where the struggle for
existence is keenest.

A. Predaceous Hymenoptera and Newroptera.

The Neuroptera are included here and not under a
separate heading because I have as yet only re-
ceived a single record, and that one in association with
an observation on predaceous Hymenoptera. Accurate
observations on Odonata and J/antispide are greatly
needed, as well as on the predaceous Hemiptera, Large
Tenthredinide should also be observed, for I have seen
them devouring insects. Locustidx furthermore are con-
siderable and indiscriminate enemies of their class. I
have seen them eating Acridians, and there is a specimen
of one in the Hope Department together with its victim,
a moth.

The late Thomas Belt long ago recorded the capture of
Nicaraguan “ Heliconide” by a yellow and black banded
wasp for the purpose of storing its nest: “ Whenever one
of these came about, they would rise fluttering in the air,
where they were safe, as I never saw the wasp attack them
on the wing. It would hawk round the groups of shrubs,
trying to pounce on one unawares ; but their natural dread
of this foe made it rather difficult to do so, When it did
catch one, it would quietly bite off its wings, roll it up
into a ball, and fly off with it” (“ Naturalist in Nicaragua,”
Lond., 1888, p. 109). The following observation of Mr.
Marshall’s upon the chief unpalatable butterfly of the
Old World compares in an interesting manner with the
foregoing :—

“Tugea River, junction with Blaauwkraantz River, Dec.
14, 1896.—I have observed two enemies of Limnas chry-
sippus lately: one isa large wasp which I sawea rrying
off the larve, and the other was a very large red dragon-
fly which was devouring an imago.” It is quite likely
that the Odonata may not uncommonly attack such
conspicuous butterflies, but this is the only record I have
as yet received.

Experiments and observations on ants suggest an almost
boundless field of inquiry. ‘The following interesting
observation made by my friend Mr. C. J. M. Gordon, M.A.,
of Balliol College, clearly proves that certain ants neglect

330 Mr. G. A. K. Marshall on

specimens of Acrzeas when they can get other butterflies.
Mr. Gordon writes of two Acrzeas captured Jan. 13, 1902,
at Old Calabar: “So distasteful do these butterflies seem
to be that even the ants will not eat them. These speci-
mens are the only survivors of a set of about twenty. The
ants got in and ate all the rest, leaving these, as you see,
untouched.” It is interesting to note that the species
were very different, beng Acrva admatha and A. neobule.

5
The specimens are in the Hope Department, Oxford.

B. Predaceous Coleoptera.

A great deal of work remains to be done with the
predaceous Coleoptera. So far as I am aware Professor F.
Plateau is the only naturalist who has made any important
use of them, but there is reason to infer from his experi-
ments that they too are important enemies to aposematic in-
sects. One section of his paper (Mém. de la Soc. Zool. de
France, tome vii, p. 375, § 9) is devoted to experiments in
which Abraxas larvee were offered to Carabus auratus, Dytis-
cus marginatus [marginalis], and D. dimidiatus. Two of the
Carabi in confinement were starved for about eleven hours,
and then given one full-grown and two smaller larve of
Abraxas. One beetle fed upon the large larva continuously
for about an hour, only leaving the thoracic region. The
other Carabus, of which one antenna was mutilated, after
half-an-hour attacked one of the smaller caterpillars and
then abandoned it. When the observer returned after a few
hours both the smaller larvae were partially eaten. . Twelve
hours later the beetles were perfectly well. The experi-
ment was renewed with two fresh Carabi starved for
eighteen hours. The beetles began to devour the larger
of two larvee given to them, and even fought over it: in
an hour only the torn and empty skin remained. By the
next morning the second larva had been devoured, and
the beetles were quite healthy. Three imagines of Abraxas
were then offered to two freshly-caught Carali. After
three hours one moth was nearly devoured, after about six
hours the second, and by the following morning the third.
There only remained some fragments of the wings. The
beetles were as active as ever. Several larva were then
thrown into an aquarium containing the two above-
mentioned species of Dytiscus. The latter at once at-
tacked them, fighting over their prey, which seemed to
be entirely consumed,

The Bionomics of South African Insects. 331

These results are tolerably decisive; but it would have
been more satisfactory if the experiments had been con-
tinued for a much longer period and controlled by others
in which different forms of food were employed. In this
way a convincing test of the wholesome qualities of the
larvee would have been supplied. In other experiments,
again, it would have been desirable to offer a wide choice,
and ascertain if there are any marked preferences.

Mr. G. A. K. Marshall has also observed one of the
Histeride, Hister caffer (Krichs.), eating a far larger beetle
than itself, the Scarabeeid Onitis aleais. He has also
observed the same beetle devouring Aphodii.

On July 19, 1898, I observed a large Elaterid (Corym-
bites virens 9) eating the larvee of Vanessa wrtice on a
nettle beside the mountain road (6000 feet) from Leukerbad
to the Gemmi Pass, Valais. The specimens are now in the
Hope Department.

C. Predaceous Diptera.

It is convenient to bring together the numerous records
of the attacks of Aszlidew upon insects into a tabular
statement. In its preparation I have received the kindest
help and co-operation from Colonel Yerbury, Colonel
Bingham, Mr. G. A. K. Marshall, Mr. A. H. Hamm, and

Dr. Chapman.

Mr. G. A. K. Marshall on

332

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The Bionomics of South African Insects.

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336 Mr. G. A. K. Marshall on

Colonel Bingham has sent me the following notes of his
observations on Asilidx: “ With regard to flies of the
family Asilide and spiders attacking butterflies, I find
only a very few scattered notices of cases which I had
seen, but no details, [ am sorry to say. So far as I can
remember I have seen these flies once or twice actually
capture butterflies, and in one instance I find it noted in
my diary that I found an Asilus with a Junonia hierta
which was still shghtly quivering its wings. I have seen
the flies not once, but often swoop at butterflies, dragon-
flies, and bees. I cannot, however, find any particular
note of that case of the Asilus with a dragon-fly the
specimens of which are in the British Museum. What
note I made was written on the paper envelope in which
the specimens had been put away. With regard to the
Asilus which I found attacking the dammar bees (J/eli-
pon), I find that I have noted that they, the flies, persist-
ently hover round the nest-mouth of the dammar bees, and
catch the latter on the wing as they issue from the nest.
The flies, so far as I have noticed, never eat their prey on
the wing, but retire | to a bush holding their prey with
their long hairy legs.

A study of the table at once shows that the Asilide are
most indiscriminate in their attacks. The stings of the
Aculeates, the distasteful qualities of Danaine and Acre-
ine and of the odoriferous Zagria, the hard chitinous
covering of Coleoptera, the aggressive powers of Odonata,
are alike insufficient protection against these active and
voracious flies. The only tendency towards specialization
in the direction of any particular group of prey appears to
be manifested in the preference of the slender Asilids of
the genus Dioctria for Ichneumonids.* The far greater
frequency with which the female Asilid has been observed
with prey is sufficiently accounted for by the larger size of
this sex and the more important part borne by it in
reproduction.

Looking at the table as a whole, and the large propor-
tion of attacks made upon specially-defended insects, the

* Since this sentence was written I have captured (July 1902)
many specimens of Dasypogon diadema with prey at La Granja,
Sierra Guadarrama, Spain. The great majority of these were
sucking Aculeates, especially the hive-bee. Another species of
Asilid, on Peialara, also exhibited an apparent preference for
Coleoptera. —KE. Bb. P.

The Bionomics of South African Insects. 337

conclusion is suggested that Bates was mistaken in sup-
posing that Asilid flies play no part as the enemies of
Heliconine and Ithomiine.

7. LEPIDOPTERA WITH WARNING COLOURS SPECIALLY
LIABLE TO THE ATTACKS OF PARASITIC INSECcTs.
(Gi ASK: 'M.)

[The late Erich Haase in his work on mimicry (English
translation “ Researches on Mimicry,” ete., Pt. II, Stutt-
gart, 1896) continually made the assumption that the
immunity of Danainx, Acrevingw, and other specially-pro-
tected groups is absolute, and extends to the attacks of
parasitic Hymenoptera and Diptera as well as to those of
insect-eating vertebrates. A little reflection upon the rate
of multiplication of animals, and especially of insects,
makes it clear that any such absolute immunity is an
impossibility. A high degree of protection from the
attacks of the generality of insect-eating animals will
always be found to be compensated by the attacks of
special enemies, and probably very largely by that of
insect parasites. I brought forward this argument in
1890 (“Colours of Animals,” London, p. 181) ; and Haase,
without attempting to meet it, made the crude assump-
tions which will now be dismissed, once and for all, by
the numerous observations recorded below.—E. B. P.]

Estcourt, Oct. 15, 1896.—We brought seventy-five
larvee of Acrwa anacreon home with us from Ulundi to
Estcourt, and no less than twenty of them were killed by
a Dipterous parasite, so that, although it may be protected
in the imago stage, the percentage of larval deaths must
be very high.

Malvern, Feb. 21, 1897.—I certainly cannot understand
Haase’s attitude with regard to protection from parasites.
There are such patent examples to disprove it among
European “whites.” Out of eight pup of Acrea horta
that I bred this season no less than five were killed by a
Dipterous parasite.

[I have also received from Mr, Marshall two cocoons
and two imagines of an ichneumon bred from <Acreva
cabira at Malvern. They bear the date April 1897.—
HB: Ee]

Umkomaas Mouth, Natal ; Sept. 3, 1897.—I think it is
highly probable that Byblia ilithyia will prove to be dis-
tasteful as you suggest ; but so far as my experience goes

TRANS. ENT. SOC. LOND. 1902.—PART III, (NOV.) 23

338 Mr. G. A. K. Marshall on

the larve are very free from parasites, which in my
opinion tells somewhat against that view.

Salisbury, March 6-10, 1898.—Of four larve of Z.
chrysippus I have taken this season two were killed by a
parasitic fly [probably a Zachina] which attacks many
different butterfly larvee.

In his “ Rhopalocera Malayana” (p. 407) Mr. Distant
writes : “ Mr. W. F. Kirby has kindly drawn my attention
to the fact that several species of Chalcis have been reared
from East Indian Danaids.” *

[Colonel J. W. Yerbury at Aden “lost a great number
of chrysippus larvee from the attacks of a large dipterous
parasite, one of the TZachinine” (Journ. Bomb. Nat.
Hist. Soc., 1892, p. 209).

Professor Félix Plateau, in his interesting paper on
Abraxas grossulariata, L. (Mém. de la Soc. Zool. de France,
tome vii, 1894, p. 375), also referred to on pp. 325-7, states
that he found twenty-two caterpillars out of fifty-one, 43 per
cent., attacked by insect parasites, viz. of Hymenoptera, two
species of Microgaster and one of Ichnewmon; of Diptera,
the Tachinid Lxorista vulgaris (Fallen). The caterpillar,
pace Professor Plateau, is most conspicuous, and, as the
Professor admits in the above-quoted paper, is refused
by European insect-eating vertebrates with wonderful
unanimity.

In the autumn of 1888 I found the conspicuous gregar-
ious larvee of Pieris brassice suffered to an enormous extent
from the attacks of Jchnewmonide. No less than 424 mature
larvee out of 631 died from this cause (Trans. Ent. Soc.
Lond., 1892, p. 489). I have also observed an excessively
high rate of mortality from the same cause among the
conspicuous specially-defended larvee of Porthesia auriflua.
Dr. F. A. Dixey informs me that he has found the larve
of Huchelia jacobex much infested by ichneumons.—
HB.

8. EXPERIMENTS ON LIZARDS AND Frogs. (G. A. K. M.)

[Experiments with lizards and frogs were few and the
results negative. A large number of the 8S. African species
are no doubt specialized to eat only certain kinds of food,
and these would be useless for experiment if their natural

* The names of Chaleis cuplea, Hope, and 0. albicrus, Klug, are
specially mentioned,

The Bionomics of South African Insects. 339

prey did not include members of the insect. Order which it
was desired to test. European lizards freely eat insects of
all Orders, and have been found very satisfactory for such
experiments. The African species which Mr. Marshall has
actually seen hunting butterflies (see p. 435) should afford
very valuable testimony as to the relative palatability of
various Lepidopterous species and groups. Even if in-
tolerant of captivity, they would probably thrive in an
enclosure out of doors, with plenty of air, sun, and space.
—KE. B. P.]

Estcourt, Oct. 15, 1896.—I have just got a lizard to try
experiments on with regard to the edibility of insects, but
though it eats Diptera and some Orthoptera readily, it
will not look at any of the butterflies I have given it, viz.
Terias brigitta, Pontia hellica, and Zeritis taikosama. I
had previously tried an Agama lizard with the same result,
as I found that its natural food consisted almost entirely
of ants.

Malvern, Feb. 21, 1897.—I have found Pontia hellica
to be distasteful to a lizard.

Salisbury, March 6-10, 1898.—I kept a lizard for about
a week, but it refused to eat any of the butterflies I gave
it, but as it also refused termites the results were unre-
liable, and I let it go. In experiments of this kind the
natural habits of the predatory animal must be taken into
account, and the fact that a lizard prefers a fly to a butter-
fly may in some cases have no greater significance, so far
as mimetic problems are concerned, than that a frugivorous
bird prefers a berry to a butterfly.

Salisbury, June 5, 1898.—I have made several attempts
at experiments with butterflies and lizards, but with no
satisfactory results, as the latter seem to be unwilling to
eat anything at all in captivity, remaining in a sort of
listless condition.

Malvern, May 14, 189'7.—Caught three frogs and put
them in a box, and put in a wingless specimen of Acrwa
natalica, After a short interval the largest frog swallowed
it. He made no sign to show that it was distasteful, nor
did he disgorge it afterwards. For several days I put in
specimens of various species, both edible and otherwise,
but as none of the frogs paid the slightest attention to
them I ceased to experiment,

340 Mr. G. A. K. Marshall on

9. EXPERIMENTS ON CAPTIVE KESTRELS (Cerchneis rupi-
coloides and C. nawmanni). (G. A. K. M.)

1899. Salisbury, Mashonaland.

January 2. Offered a young kestrel (Cerchneis rupi-
coloules) an Acrva caldarena ; he took it
in his beak, held it for a few seconds and
then threw it away with a sharp shake
of the head. Then gave him a Byblia
ilithyia, which he accepted, but after
making one or two pecks at it let it drop
and would not touch it when it was again
offered. I then tried him with an example
of A. nohara-halali and A. doubledayi-
axina, but he would have nothing to do
with them.

7. Gave the kestrel a large Buprestid beetle
(Psiloptera valens, Pér, 7.1.); he seized it in
his beak with a cry of evident pleasure,
then holding it in his foot tried to eat it,
but after a peck or two it slipped from
his grasp; I gave tt back to him several
times, but always with the same result—
the beetle was too slippery. Psiloptere
are all eaten readily by baboons, and it
would appear from this that their hard,
shiny integuments combined with their
torpedo-like shape form a very efficient
protection (apart from their procryptic
colouring) from all birds which are not
sufficiently large to swallow them entire.

8. Offered kestrel two Dlepisanis haroldi, a
small Longicorn with Lycoid markings,
but he would not touch them; then a
Lycus rostratus, at which he pecked but
was evidently displeased with the taste,
and neglected it.

12. Gave kestrel a Precis pelasgis, Junonia
cebrene, Precis sesamus @) and Atella
phalantha. He ate them all with evident
relish, though he seemed to experience a
little difficulty in managing them at first,
as he could not get a good hold with his
claw in order to pull them to pieces, So

”

”

The Bionomies of South African Insects. 341

the first two were practically swallowed
whole, but the others were eaten piece-
meal. I then offered a larva of Limnas
chrysippus, which he accepted and held
for some moments in his claw as though
in doubt, but finally let it drop after a
half-hearted peck. On _ re-presentation
he would not touch it, so in order to
restore confidence I gave him a grass-
hopper, on which insects he is usually
fed. The species offered happened to
have some rather light-green, yellow, and
black markings; he took it with evident
distrust and soon dropped it, although it
was a species he had often eaten before.
Thinking that the refusal was due to
his experience with the brightly-marked
larva of ZL. chrysippus, I offered some
dully-coloured green and brown grass-
hoppers, which were readily eaten, and
after them he also ate the one previously
refused. I then offered a larva of Acrva
rahira ; he evinced a decided interest in
it, but, although it crawled about over his
feet, absolutely refused to touch it. This
however may have been due to a general-
ized impression that all caterpillars were
distasteful.

January 13. Cerchneis vupicoloides ate one Catopsilia

florella. A young bird of another species
(C. nawmannt) accepted a dead Papilio
demodocus; for a few moments he seemed
undecided where to attack it: then
noticing the eye-spots in the hind-wing
he promptly pecked them out, afterwards
eating the rest.

16. C. nawmanni ate three Terias brigitta ;

both this species and C. rupicoloides
refused the Lycoid Prionocerus dimidiatus
with unmistakable signs of dislike.

C. naumanni ate one Precis pelasgis, one
Axiocerces harpax, one Terias brigitta,
one Catopsilia florella, and one Precis
sesamus (natalensis form) @), all with

January 22.

25.

Mr. G. A. K. Marshall on

evident appreciation. C. rupicoloides
appeared to have become tired of butter-
flies, refusing all, even those it ate before.
Swynnerton found that it continually
refused the brightly-coloured grasshopper
mentioned above, but always ate it with
relish when it had been dipped in meal
to obscure its colours. C. nawmanni on
the other hand never refused the insect.

C. naumanni accepted a Papilio corinneus,
but seemed in some doubt as to its
edibility ; he finally ate the thorax and
threw away the abdomen. <Acrwa halali
was then offered, tasted, and rejected.
Acrxa anemosa and Limnas chrysippus
were likewise refused, and shortly after-
wards one byblia ilithyia and one Terias
brigitta were eaten.

C. rupicoloides escaped, and all the follow-
ing notes refer only to C nawmanne.
Kestrel ate one Dichtha inflata and one
Amblysterna vittipennis. A Lagria, sp.,
was tasted and rejected.

When very hungry the bird ate part of an
Acrexa caldarena, throwing away the rest.
Subsequently he ate four Belenois severina,
one Junonia cebrenc, and one Precis sesamus
(natalensis form).

Two Byblia ilithyia eaten by kestrel. A
Clythra wahlbergi, with strong Coccinellid
odour, was eaten by the kestrel after
some hesitation. Kestrel refused the
Coccinellid Epilachna dreget.

A. Longicorn (Ceroplesis fallax, Pér.) offered
to kestrel, which had been kept without
food for some time. He was evidently
nervous and much impressed by the
stridulation of the insect. I therefore
pulled the head off the beetle, and the
hawk then ate it, but very slowly and in
such a way as to lead me to suppose that
it was not altogether palatable. Gave
a Piezia selowst, head first, to kestrel, which
ate it readily though evidently noticing

The Bionomics of South African Insects. 345

the acidity of the abdomen. A Poly-
hirma enigma was at once eaten by
kestrel. I then offered him Graphipterus
lineolatus, tail first; he pecked at it and
received a small discharge of acid in the
mouth, whereupon he shook his head
and began wiping his beak vigorously on
the perch, as though to get rid of the
taste. Upon the beetle being presented
head first, he took it with caution and ate
it. The same results were obtained with
Graphipterus wahlbergi, G. bilineatus, and
G. lineolatus, they being refused when
presented tail first and eaten when
reversed. It should be noted that these
beetles all discharge their secretions
violently when captured, and therefore
the kestrel would probably receive a
comparatively small dose of the acid.
February 1. Gave kestrel an Afella phalantha ; he
seemed a good deal doubtful about it at
first, but finally ate it without any signs
of distaste. He then ate a Junonia
cebrene and a Byblia withyia with manifest
enjoyment. I then offered Z. chrysippus ;
he accepted it readily, pulled off the
head which he discarded, pecked a
little at the tough thorax and wings, and
then let it drop; on offering it again he
took it, gave it a few pecks and jerked it
away with his beak. He then ate a
Hamanumida dxdalus and accepted an
A, caldarena, of which he ate a small part
of the abdomen and threw away the rest.
After this he ate with pleasure a P. sesainus
(natalensis form), J. cebrene, and B. ilithyia.
Several Onitis alexis were then given
to the kestrel, which ate them readily.
Anomalipus plebeius was too hard for him,
and after five minutes’ hard pecking he
had only succeeded in pulling off the
head; I therefore broke it up for hin,
and it was promptly eaten. He then
refused Clinteria infuscata, Mylabris holo-

344,

Mr. G. A. K. Marshall on

sericea, Olerus sp. (entirely scarlet, with
strong verbena-like smell), and Prionocerus
dimidiatus,

February 3. Kestrel ate several dull-coloured Cureulio-

?

»”

”)

nide (Oosomus, sp., and LHremnus, spp.),
refusing several Onthophagus gazella which
were offered, also Lycus ampliatus, L.
rostratus, and L. constrictus, all of which
were tasted and were very evidently
unpleasant.

6. Kestrel refused Zonitis, sp., Hletica rufa,

Mylabris palliata, and Diacantha conifera,
after tasting each.

The kestrel had been starved for twenty-
four hours, and was very hungry. He ate
the following insects, in the order given,
with great avidity: two Teracolus achine,
one B. ilithyia, two Atella phalantha, one
Junonia cebrene, and one Papilio corinneus.
I then gave him an Aereva caldarena, of
which he first ate the head and swallowed
the rest whole, one A. rahira was also
swallowed whole; Z. chrysippus was then
offered ; the bird ate the head, which
seemed to raise suspicions in his mind,
for he sat considering for some moments,
and then began pecking at the thorax
and wings and finally dropped it; on
re-presentation he seized the butterfly,
gave a few pecks at it, and jerked it away. ©
Then one Acrva axina and two A. halali
were swallowed whole, but A. caldarena
which followed was only partially eaten,
fully half being discarded. Another
L. chrysippus was offered, and the entire
head and thorax was eaten before it was
thrown down. Later, the bird swallowed
whole another A. rahira subsequent to
eating several grasshoppers.

28. Gave kestrel a B. ilithyia, which was eaten

rapidly, and a second as well. He then
ate a Precis pelasgis and another aithyia,
but an Acrva halali was pecked at once
or twice and thrown away, and a fourth

The Bionomies of South African Insects. 345

ilithyia was treated in the same way,
being apparently mistaken for an Acrwxa.
I then offered a female Anoplocnemis
curvipes; the bird ate the head, but
evidently in some doubt; it continued
with the thorax however, but showed its
dislike by repeated sharp shaking of the
head, and finally dropped the abdomen.
I put a male of the same bug on its
perch, but though it examined it carefully
it would not touch it; yet this species is
eaten greedily by the baboons.
[There are several very significant results from the
above-recorded experiments on kestrels. The rejection of
Byblia, after trial and rejection of an Acrwa, may have
been due to the superficial resemblance. On the other
hand, this bird (@. rupicoloides) was apparently not fond of
butterflies, for after eating (Jan. 12) Precis, Junonia, and
Atella and (Jan. 13) Catopsilia, he refused all Rhopalocera.
The refusal of an Acridian marked with bright green,
yellow, and black, and its acceptance when the colours
were hidden was almost certainly the result of unpleasant
experiences with conspicuously-marked insects, of which a
particular instance was afforded when the larva of JZ.
chrysippus was offered. Such association of impressions
brought about by very imperfect resemblances are of great
importance in helping us to understand the origin of
mimicry, both Batesian and Miillerian, in slight accidental
resemblances of a very rough and imperfect kind. It also
warns us not to regard as far-fetched or absurd those
imperfect likenesses which may well be the early stages
of incipient mimicry. The refusal of the Lycoid Longicorn
Blepisanis may be similarly due to a previous experience
of Lycidex, or it may be truly distasteful and synaposematic.
The latter interpretation is certainly true of the Lycoid
Melyrid Prionocerus also refused by the kestrel “with
unmistakable signs of dislike.”

The other species of kestrel, C. nawmanni, was much
fonder of butterflies and of insects generally, eating the
brightly-coloured grasshopper on all occasions. The fact
that it took special notice of and pecked at the eye-spots
on the hind-wing of P. demodocus is of much interest, and
recalls an observation of my own quoted-on pp. 440, 441.
Such observations strongly confirm the interpretation of

346 Mr. G. A. K. Marshall on

eye-spots, especially upon the under-side of the wings, as
directive marks leading an enemy to attack a non-vital
part, and they tend to refute Portschinski’s explanation
of them as the representation in colour of drops of some
specially-protective fluid (see p. 398).

Butterflies of different groups, Hesperide, Pierine,
Nymphaline, were freely eaten, but the rejection of the
abdomen of Papilio corinneus by the captive bird which
afterwards ate Byblia and Terias, can only be explained
on the supposition of unpalatability, and the same was
evidently true in a more marked degree of Acreas and
L. chrysippus, although parts of these would sometimes be
eaten, while on Feb. 15, after starvation for twenty-four
hours, many Acreeas were swallowed whole. The behaviour
on this occasion renders it certain that, as in the case of
Bucorax caffer, L. chrysippus was far more distasteful to
the kestrel than the Acrieas.

The rejection, after trial, of the evil-smelling Coreid
bug 4. curvipes, greedily eaten by baboons, is a good
example of the difference in value of the same defence
with different enemies.

The treatment of Coleoptera almost invariably supported
the theories which explain the meaning of insect colouring
as cryptic, warning, ete.

The following beetles were eaten by the kestrel:
Curculionide, with cryptic colouring (Oosomus, sp., and
Lremnus, spp.) ; the large, slow-moving, conspicuous, black,
earthy Heteromeron Anomalipus plebeius, when the chitin
was broken; the smallish Buprestid Amblysterna vitti-
pennis (dark metallic green or coppery with white stripe
on each elytron); the Heteromeron Dichtha inflata, dark
brown with reddish stripes, conspicuous and slow-moving
like Anomalipus ; the medium-sized Scarabeid Onitis alexis
with elytra and legs brown, and thorax iridescent green.

It is probable that most of the defensive fluid had been
already discharged in the case of the Carabidxe of the
genera Piezia, Poly olyhirma, and Graphipterus, of which the
acid secretion was seen to be a very positive protection
when there was opportunity for its operation on a normal
scale. The Longicorn Ceroplesis fallax with a Cantharid
type of colouring may be synaposematic, as it was only
eaten very slowly although the bird had been kept without
food. The impression produced by the stridulation is of
much interest (see p. 403).

The Bionomics of South African Insects. 347

The following beetles were refused, usually after
tasting :—

CANTHARIDE :—Mylabris palliata, M. holosericea, Eletica

rufa, Zonitis sp. (all most conspicuous).

COcCINELLID& :—Lpilachia dreget (characteristic col-

ouring).

CLERIDE :—“ Clerus” sp. (scarlet).

PHYTOPHAGA :—Diacantha conifera (Lycoid).

MELYRIDE :—Prionocerus dimidiatus (Lycoid).

Lycipa#:—Three characteristically coloured species of

Lycus.

CETONUDH :—Clinteria infuscata (orange thorax with

two black spots, brown elytra, sometimes black).

HETEROMERA :—Lagria, sp. Probably distasteful, con-

spicuous and synaposematic with Phytophaga.

SCARABEIDA :—Onthophagus gazella, smallish Scarabzeid

with brown elytra and iridescent dark green thorax
and head.

With the possible exception of the last named, all these
species possess distinct aposematic colouring, and nearly
all belong to groups which are much mimicked, or fall
into important synaposematic combinations.

Mr. Marshall specially points out that the Kestrel, C.
naumanni, was young, and it is probable that it had

never before had experience of many of these species——
HB. P|

10. EXPERIMENTS ON A TAME GROUND HORN-BILL
(Bucorax caffer). (G. A. K. M.) .
Malvern, Natal, May 14, 1897.
March 14. Gave a tame ground horn-bill (Bucorax
caffer), belonging to Col, J. H. Bowker,
the following butterflies: two male A.
serena, one P. lyxus, one male H. misip-
pus, one male A. serena, all of which he
ate readily, taking them in the end of his
beak, crushing the thorax and throwing
them down his throat. I then gave him
L. chrysippus. He took it, crushed the
thorax and dropped it at once. A second
specimen given a short time afterwards
was treated in the same manner.
, 24. Gave the following butterflies to ground
horn-bill: three A. encedon, one A. petrea,

348. Mr. G. A. K. Marshall on

one P. aganice, two J. clelia, three male
H. misippus, one P. tropicalis, two P.
brasidas, two P. demodocus, one P. lyxus,
and two &. forestan. He ate every one
without the least hesitation, and evidently
appreciated them, as he would follow me
about, waiting for more.

April 1. Gave ground horn-bill one A. petrxa, two
A. cabiva, one P. brasidas, one male H.
misippus, and one P. esebria, all of which
he ate readily.

[It has already been pointed out that the acceptance of
insects by insectivorous animals in captivity is no proof
of their normal likes or dislikes in a wild state. Such
acceptance only proves what their action would be when
they had been, from some exceptional cause, kept without
their normal food in its usual quantity and variety. Hence
the fact that the Acrzas were devoured is no evidence
that they are normally eaten except in a time of unusual
hunger. On the other hand, the rejection of two L. chry-
sippus, after three Acrzeas had been readily eaten, indicates
that the former butterfly is decidedly distasteful to this
species of bird. It must be remembered that five Acraeas
were freely eaten on the next occasion. A comparison
with the experiments on Mantides is interesting.—K. B. P.]

11. Tue INSECT-FOOD OF WILD SoutH AFRICAN BIRDS.
(G, A. K.M.)

[Even more important than the results of experiments
are the observations made and collected by Mr. Marshall
upon the contents of the stomachs of birds, and the record
of actual attacks made by birds upon insects, which have
been witnessed in the field. The contents of birds are
clearly shown in the two following tables, A and B, which
are printed just as I received them from Mr. Marshall,
except that I have added a brief description of the general
appearance of those insects which seemed to require it.
Mr. Marshall had only supplied such a description in three
or four cases. In future records of this kind it will be
advisable for the observer on the spot to supply such
notes, together with a brief account of the habits, in-
asmuch as conspicuousness or concealment depend upon
these quite as much as upon colour and pattern —KE. B. P.]

The Bionomics of South African Insects.

349

TaBLeE A.—Contents of birds, probably 1898, unless otherwise
stated, and Salisbury when no other locality is mentioned.

BIRDS.

INSECTS, ETC,

GENERAL APPEARANCE OF INSECTS.

Macronyx capensis.

CoLEOPTERA, Clconus sp.,
Eremnus sp;
puncticollis,
Sympiexiorrhynchus sp.

Syagrus |
Lefev. ,

Syagrus, a shining black,
medium-sized Phytophagous
beetle (Humolpide). All
others are weevils, and pro-
bably all with cryptic
colouring.

Trrisor erythrorrhyn-
chus.

Homoprera, Pyrops sp.
CoLEoPTERA, Platypria
mashuna, Pér., Anthaxia
sp.

| Pyrops, at rest are red-brown
or greyish insects. Platypria,
tawny with brown spots,
very spinous (/Hispid).
Anthaxia, small green or
coppery Buprestid.

Coracias garrula. CoLEoPTEeRA, Gymnopleurus| Large, sooty-black, smooth
Sastiditus, Har. Scarabeid.
C. caudata. CRABS. Agrilus, small Buprestids,

CoLEOPTERA, Agrilus sp.,
Anthia pachyoma (!).

colour varies. Anthia, huge
black Carabid with very
powerful mandibles.

C. spatulata.

Drerera, fly-maggots from
carrion.

C. olivaceiceps.

OrTHOPTERA, Phymatews
morbillosus, L. (a large
evil-smelling bright-green
locust with purple and
crimson hind-wings).

Very conspicuous, with red
thorax and head; legs red
and yellow.

M elittophagus
pusillus.

Merops natalensis.

|

| COLEOPTERA, Onthophagus,
| sp.

Genus of Scarabxide, varying
much in size: metallic or
black.

COLEOPTERA, Mylabris
oculata, Thunb. (1).

Cuculus gularis.

| COLEOPTERA, Jyassinius
lugubris, Sphenoptera
disjuncta, Hoplonyx sp.

Characteristic Cantharid, apose-
matic, orange and black
colours.

Nyassinius, small red-brown,
rough Cetoniid, probably
cryptic. Sphenoptera, dark
metallic, coppery, moderate-
sized Buprestid. Hoplonyx,
medium-sized, black often
polished, Heteromera (Tene-
brionide).

Asturinula
monogrammica.

Falco subbuteo.

OrTHOPTERA, Clonia wall-
bergi. CENTIPEDES.

Clonia is a fine Locustid living
among the leaves of trees.
It is certainly procryptic.

CoLEorTERA, Pentodon
nireus, Burm., Onitis
alexis, Anomala sp.

Pentodon, large black Dynastid.
Onitis, medium-sized Scar-
abeid,. elytra and _ legs,
brown; thorax iridescent
green. Anomala, pale yellow
brown, or metallic Rutelide ;
moderate-sized beetles,

350 Mr. G. A. K. Marshall on

TABLE A.—(continued.)

BIRDS. INSECTS, ETC. GENERAL APPEARANCE OF INSECTS.
SON eet i, [ene $$ —_—___]—
Cerchneis CENTIPEDES.
rupicoloides.

C. nawmanni. CoLEopTERA, Heteronychus | Heteronychus, smallish black,
licas,Klug.,CENTIPEDES.| rather shining Dynastid.
ARACHNIDS, Solpuga mar- Solpuga is red-brown, distal
shalli, Poe. part of 4th leg black, broad

median black band on
abdomen which is clothed at
sides with yellowish-white
| hairs. It runs very swiftly
and its habits are strongly

procr yptic.

C. amurensis. COLEOPTERA, Hydaticus, bude sized, oval, polished
sp. dark-brown Di ytiscidee.

Bubo maculosus. COLEOPTERA, Hespero- | A large red-brown Cerambycid :
phanus amicus, white. procryptic.

Herodias lucidus. Dragon-flies and aquatic
Hemiptera.

Ciconia abdimii. CoLEoPTERA, Psammodes | Psammodes, both species are
ventricosus, Fahr., P. large dull- brown Tenebrionid
scabratus, Gerst., Poly-| Heteromera. Polyhirma,
hirma semisuturata, | moderate-sized, black Car-
Chd., Piezia marshalli, abid with white markings.
Pér., Scarites, sp. Piezia, a Carabid super-

HyMENOPTERA, Ants of | ficially similar to last.

the genus Carebara. Scarites, shining black Car-

| abids of variable size ; large
| mandibles.

Numida coronata. | COLBOPTERA, 2 Peiloptera | Psiloptera, largish iridescent
Shot by C. F. M. | chalcophoroides, eral bright-green Buprestids.
Swynnerton at. Hipporhinus bohemanii,| Macrocoma, a small golden-

| Mazoe, Mashona- Fihr., 1 Phantasis| green iridescent Phytopha-

land, 4000 ft. | gigantea, Gueér., 1 Macro- gous beetle. Hipporhinus,
| | coma awreov illosa, Mash. | a large brown rough eryptic
| These five specimens were | weevil. Phantasis, a large
| taken from the crop,,| Longicorn generally similar
| December 25, 1898, by | to the above, and probably
| tGiny: BALK: Marshall. mimetic of certain very hard
| They are now in the Curculionide.

|

Hope Collection, Oxford, |
| _ and are extraordinarily
| perfect, retaining the legs
| and in some cases the |

| antenne. |
Merops natalensis, ‘LEPIDOPTERA Hrrerocera, The moths both conspicuous,
September 1901. the Saturniide Psewd-| slow, day-flying, and pro-
aphelia apollinaris and bably distasteful species.
Cirina similis. Pseudaphelia, is large semi-

transparent, whitish with
black markings; Cirina is
still larger and dull pink.

eS

The Bionomics

of South African Insects. 351

TABLE B.—Jnsects, etc., in stomachs of birds (probably 1898),

COLEOPTERA.

Salisbury.

BIRDS.

| Polycleis decorus (largish weevil,
varying much in colour, but
always with more or less of
a pattern).

Oriolus notatus, Coracias caudata, Haleyon
pallidiventris.

Trochalus sp. (small rounded
often polished black-brown
or greenish Lamellicorns ;

pist.

probably mimics of Galeru-
cide, Chrysomelide, and

Coceinellidz),

Cassida (Aspidomorpha) punc-
tata, F. (medium _ size,
abundantly black-spotted,
colour probably reddish-
brown when fresh).

Alcides hxemopterus, Boh.
(smallish weevil with red-
brown white-spotted elytra
and black thorax and head).

B. mariquensis, Irrisor
Campothera bennetti.

erythrorrhynchus,

Prionops talacoma.

Oosomus sp. (an entirely black,

qraucalus pectoralis, Upupa africana, Irrisor
erythrorrhynchus, Campothera bennettt.

arboreal or — subcortical
weevil).
| Zophosis sp. (black quick-

running Heteromeran).

Onthophagus gazella (smallish
Scarabeeid with brown elytra |
and iridescent dark-green
thorax and head).

Geocichla litsitsirupa.

Cuprimulgus rufigena, Falco subbuteo (in |
large numbers).

OTHER INSECTS, ETC.

Pentatomid bugs,

| Geocichla litsitsirupa, Laniarius guttatus,
Trrisor erythrorrhynchus, Cerchneis amur-
ensis, Coccystes glandarius.

Reduviid bugs.
|

| Ant-lion larvee.

Ants.

Macronyx capensis, Rhinopomastus eyano- |
melas.

Thamunolxa cinnamomeiventris.

Bradyornis mariquensis, Pratincola torquata, |
Monticola angolensis, Saxicola pileata, |
Buchanga assimilis, Thamnolea cinna-
mometventris, Crateropus kirkii, Lopho-
ceros leucomelas, Campothera bennetti,
Crecopsis egregia.

S02 Mr. G. A. K. Marshall on

i

TABLE B.—(continaed.)

OTHER INSECTS, ETC, BIRDS.
| Other Hymenoptera. Rhinopomastus cyanomelas, and all four
| species of bee-eaters.
Hairy caterpillars. Oriolus larvatus, and all the cuckoos.
| Millepedes. | Turdus libonyanus.
| = es : a es =
Scorpions. | Coracias olivaceiceps, Cerchneis rupicoloides,

Asturinula monogrammica (apparently
favourite food with this species).

VERTEBRATES,

| Lizards. Melierax polyzonus, Astur polyzonoides,

|

Aquila wahlbergi, Circetus pectoralis,
Cerchneis rwpicoloides, and Glaucidiwm
perlatum.

| Snakes. Astur polyzonoides, Buteo jakal, Circetus

pectoralis, Bubo maculosus.

RESULTS OF TABLES A AND B.

[The almost complete absence of the members of apo-
sematic Coleopterous groups is very marked. In fact, the
whole of the numerous beetles are probably cryptic, with
the following exceptions. The species of the EKumolpid
genus Syagrus is probably distasteful; for it freely exposes
itself on leaves, where its shining black appearance renders
it conspicuous. It is worthy of note that the only bird in
which it was found, Macronyx capensis, also ate Reduviid
bugs. The Phytophagous Macrocoma auwreovillosa belongs
to a probably distasteful group, but it is itself green in
colour; it was only eaten by one species. The Hispid
Platypria is probably distasteful, and here too the only
species of bird which ate it, Jivisor erythrorrhynchus, also ate
the conspicuous Cassida (Aspidomorpha) punctata and
Pentatomid bugs. ‘The above-named Cassid-was also found
in two other species of bird. The most remarkable exception
is however the typically-coloured Cantharid, Mylabris
oculata, only detected in Merops natalensis. Here we
find the interesting proof that under certain circumstances,
and with certain enemies, the most marked distasteful

The Bionomies of South African Insects. 353

or unwholesome qualities accompanied by the most con-
spicuous orange and black aposematic coloration may
afford no protection. Furthermore, it is of great interest
to observe that the same species of bird was the only
one in which two conspicuous and almost certainly dis-
tasteful Saturniid moths were found. The Carabide of
the genera Anthia, Polyhirma, Piezia, and Scarites are not
so remarkable. Scarites is probably nocturnal and entirely
procryptic, while the defensive secretions of the three
other genera may be discharged and lost as the result of
the attacks of an experienced enemy.

Outside the Coleoptera, the number of birds which ate
Pentatomid bugs is remarkable (five species), and it
would be interesting if it were possible to obtain the
remains and make out the species of these Hemiptera.
The specialization of enemies to feed upon forms which
have become excessively abundant through specialization
in their modes of defence is seen in the two species which
contained ants, and the three which had eaten scorpions.
The hairy caterpillars eaten by cuckoos are a similar case ;
this group of birds being specialized to feed on insects
which are specially defended against the majority of insect-
eaters. The fact that Phymatews morbillosus, a large,
conspicuous, and strong-smelling locust, had been eaten, is
also of interest. Solpuga marshalli, in spite of its formid-
able appearance, is quite harmless, with procryptic appear-
ance and habits. The Tables as a whole afford wonderfully
strong support to the existing theories which explain
cryptic colouring and instinct as the defence of forms
which are eagerly sought for as food by numerous enemies,
and an aposematic appearance and mode of life as the
defence of specially-protected forms only attacked under
the stress of hunger or by comparatively few specially-
adapted foes.—E. B. P.]

12. RECORDS OF ATTACKS ON LEPIDOPTERA, ESPECIALLY
BUTTERFLIES, BY WILD SouTH AFRICAN BIRDS.
(Gea Ke Ms)

[The stimulus which induced Mr. Marshall to collect
observations on the attacks of birds upon butterflies was
provided chiefly by the account of the discussion which
followed Dr. F. A. Dixey’s paper on ‘‘ Mimetic Attraction”
(Trans. Ent. Soc. London, 1897, p. 317; Discussion in

TRANS. ENT. SOC, LOND. 1902.—PaART 11. (NOvV.) 24

354 Mr. G. A. K. Marshall on

Proc. 1897, pp. xx—xxxil, xxxiv—xlvii), The following
extracts from a letter indicate the line Mr. Marshall would
have taken had he been in England at the time.—E. B. P.]

Malvern, Natal ; Oct. 7, 1897—I am much struck with
the large amount of adverse criticism levelled against the
theory of even Batesian mimicry. The theory of converg-
ence (Miillerian mimicry) might perhaps be considered as
debatable, but how any one who has paid any attention
to the subject can doubt the reality of Batesian mimicry,
I cannot understand, and the attempt to explain it away
by climatic causes seems to me weak in the extreme. If
the view, advocated by many, that birds cannot be reckoned
among the principal enemies of butterflies in their imago
state, be true, then I consider that we may practically
abandon the whole theory of mimicry as at present applied
to the Acreine and Danaine of South Africa at all
events, for from what I have observed of these insects I
am convinced that their warning coloration cannot have
reference to either mantises, Asi/ida, or lizards, which are
practically the only other enemies that can be taken into
account, Moreover, the swift flight of the majority of
edible species can only have been developed to enable
them to escape from winged enemies, and that this de-
velopment is due to Asilidw or dragon-flies is more than
I can believe. Certainly the paucity of records of birds
eating butterflies is somewhat disconcerting, but this is
doubtless due to the fact that not sufficient attention has
been paid to the subject, which would entail long and
patient observation of the birds themselves, an occupation
that the average entomologist is not likely to indulge in
when out collecting. Personally I do not suppose I have
seen such an occurrence more than perhaps half-a-dozen
times; the birds being the Paradise flycatcher (Zerpsi-
phone perspicillata), the bee-eater (Merops apiaster), and
two rollers (Coracias spatulata and Lurystomus afer); but
then I admit that I have paid little or no attention to
the matter until quite recently.

The habits of the Zeracoli, especially in their winter
forms, have always seemed to me strongly suggestive of
their being frequently attacked by birds. With hardly an
exception they are fairly swift fliers (especially the.
“ purple-tips ”), keeping comparatively close to the ground
and dodging well. If struck at gently as they fly by,
they dodge and hurry onwards but still continue their

The Bionomics of South African Insects. 355

flight; if however they be thoroughly frightened by
continued strokes of the net, they will dart rapidly on for
a short distance, then vanish—or, in other words, they settle
with extreme suddenness, and their under-side colouring
harmonizes so well with the sandy soil they love that they
are very difficult to detect. It seems to me that such a
habit can only have been developed for the purpose of
escaping from birds, and must be very effectual in most
cases. I have noticed that the summer forms, which have
not the sandy-coloured under-side do not adopt these
tactics, but rely on their flight alone—probably because
food is more plentiful for insectivorous birds at that
season,

[After this, Mr. Marshall kept a careful record of obser-
vations. His results, including one observation made at
an earlier date, are shown on pp. 357-9 in the form of
a diary. The two following letters bear on the same
subject.—E. B. P.]

Salisbury, March 6, 1898.—I was much interested in your
arguments * for Common Warning Colours in butterflies
and your remarks on their probable enemies, but I must
candidly confess that I am not altogether convinced. The
difference in our views lies in your fundamental proposi-
tion that butterflies are an easy prey for birds to capture
from a general point of view. If this proposition be
correct, then I quite agree that your theory offers the most
natural and probable explanation of the predominance
of bright colours among butterflies. But from what I
have seen of the South African species I could not truth-
fully say that I consider that they would be likely to fall
an easy prey to birds, indeed I should say that the average
insectivorous bird would not have a chance against most
of the swift-flying species when on the wing, and would
only be able to catch them under exceptionally favourable
circumstances when the insects were off their guard. If
this supposition be correct it would go a good way to
explain how so many butterflies have been able to acquire
such brilliant colours, and particularly in the case of
those species which have protectively-coloured under-sides,
which is the rule rather than the exception. Birds would
soon learn the futility of attempting to pursue such species,
and would only capture them by stealth,and in a more or less

* Some of the arguments here referred to are set forth on pages
: ° 2 = te)
500 to 502 of the present memoir.—E, B. P.

356 Mr. G. A. K. Marshall on

unobtrusive manner, which might account for the paucity
of records. The fact that birds have been seen to capture
moths more frequently than butterflies need not neces-
sarily imply a preference for the former insects, but might
be explained on the supposition that they are aware that
they can be captured more or less easily on the wing, and
therefore that when a moth does happen to get well up
into the air in open country it is promptly pursued,
whereas under similar conditions a butterfly would be
allowed to pass unmolested. While on the subject of swift
flight I might mention that I was much struck during my
visit home with the slow flight of English butterflies as
compared with the generality of South African species.
I am inclined to agree with Trimen in his Presidential
Address to the Entomological Society, that birds are
among the chief enemies of butterflies. That they have
been the chief, if not the only, agents in the production
of mimicry, whether Batesian or Miillerian, I have little
doubt. It is highly significant that mimicry in its fullest
development is only to be found in forest-clad regions
where insectivorous birds are most abundant. Moreover,
T am not aware of a single instance of true mimicry
among species which habitually settle on the ground.
Salisbury, March 10, 1898.—-It would seem that mere
unpleasantness of taste or smell would hardly be sufficient
to give so great an immunity from attack from birds as is
apparently enjoyed by the Danainw and Acreine, unless
accompanied by poisonous or unwholesome qualities—at
least, if we may judge by other orders of insects. <A large
number of Rhynchota, for instance, possess a very un-
pleasant smell, and yet their colouring is procryptic stead
of aposematic. In the crop of the great spotted cuckoo
I have found a large green Pentatomid, which in the
strength and unpleasantness of its smell is only beaten
by Petascelis remipes, our largest Hemipteron. Again, in
the crop of the racquet-tailed roller (Coracias caudate,
Trim.) I have found a full-grown specimen of a large
Phymateus locust, which is a most evil-smelling beast.
This insect appears to combine procryptic and aposematic
colours; for when settled its general green colour is
eminently protective, but during its laboured flight it is
most conspicuous owing to its brilliant crimson and purple
hind-wings. If annoyed when settled on the ground they
often raise their wings over their backs (clearly to exhibit

The Bionomics of South African Insects. 357

the bright colours), exuding at the same time an odoriferous
- frothy liquid from the thorax.

1897.

March 28.

1898.

While out collecting at Malvern, Durban,

Natal, I saw a Paradise flycatcher
(Terpsiphone perspicillata) catch a speci-
men of Eronia cleodora. The butterfly
was hovering over a flower when the bird
swooped down, seized it with its feet,
and carried it off.

Feb. 27. Saw a Marico wood-shrike (bradyornis

March

mariquensis) dart down from a tree and
catch a Sarangesa eliminata (Holl.),
which was sitting with outspread wings
on a small plant.

6. Saw a flycatcher (Pachyprora molitor)

make several futile attempts to catch a
Tarucus plinius which was cireclmg round
the bush on which it sat.

Noy. 23. Saw a bush kingfisher (Haleyon chelicuten-

Dee.

PP)

1899.
Jan.

il.

ile

sis) catch and eat two butterflies, viz.
Junonia cebrene and Catopsilia florella,
both of which were captured when
feeding.

F. M. Swynnerton saw a drongo
(Buchanga assimilis) fly past him with a
white butterfly in its beak, probably (.
flovella.

15. Remains of Papilio demodocus found in the

stomach of a cuckoo (Coccystes caffe).

While watching an Atella phalantha hover-

ing over a bush of its food-plant, a
Paradise flycatcher (Terpsiphone perspici-
lata) darted past, and with a loud snap of
its beak tried to catch the butterfly im its
swoop. The latter escaped, however, and
on following it up I found that the tip
of one hind-wing had been cut clean off;
unfortunately I had no net and failed to

capture the insect.

Swynnerton shot a hobby (Falco subbuteo),

which had in its stomach an almost

358

April 26.

1900.

May 13.

1901.

Dec. 17

Mr. G. A. K. Marshall on

complete Zerias. The thorax and abdo-
men were quite uninjured, but the tips
of the fore-wings were gone.

I was watching a drongo hawking insects

from the top of a dead tree; there were
many Pierine about, chiefly Zeracolus
and Belencis, but the bird paid not
the least attention to them. At last
a Belenois came by which had its wings
very much shattered, so that its flight
was weak and erratic; the drongo
observed it at once, and swooped down on
it, but I saw the butterfly drop into the
long grass. Whether it was injured by
the bird I could not say, as I was unable
to find it, and I did not see it rise again.
This episode would point to the conclu-
sion that the fact that birds refrain from
pursuing butterflies may be due rather to
the difficulty in catching them, than to
any widespread distastefulness on the
part of these insects.

C. F. M. Swynnerton wrote from Gazaland :

“Tn March [1900] I saw a Pratincola
torquata [South African stonechat] in
chase of Turucus plinius. Had it not
been frightened off by coming face to
face with me, if would undoubtedly have
caught it. I think I told you long ago
of having found the wings of a lot of
butterflies, chiefly P. corinneus, below the
branch of a tree on which some swallows
were constantly settling.”

Salisbury. Saw a drongo (Buchanga assi-
milis) swoop from a rae and cateb, what
I took to be an injured Belenois, which it
dropped almost at once. I marked the
insect down, and found it to be a common
white moth of the distasteful genus Dia-
crisia (D. maculosa).

Melsetter, 5500 feet, Gazaland. <A speci-

men of the large, conspicuous, Hypsid

The Bionomics of South African Insects. 359

moth Callioratis bellatrix was seized and
rejected by a drongo, undoubtedly a
young bird, judging by its plumage.
[The moth, which is now in the Hope
Department, has lost most of the head,
but is otherwise uninjured.—K. B. P.]

13. Recorps oF ATTACKS ON BUTTERFLIES BY WILD
BrrRDs IN INDIA AND CEYLON, BY CoLONEL J. W
YERBURY, R.A.

1884.

1885.

[Colonel Yerbury has kindly extracted from

his notes all the observations he has made
bearing on this interesting question.—

E. B. Pl

“About the year 1884 a discussion arose in

the Bombay papers as to whether birds
preyed on butterflies, and the general
opinion expressed was that it was com-
paratively rare for them to do so. In
common with some other members of the
Bombay Natural History Society, I deter-
mined to watch and note the results. My
records taken from old diaries are as
follows :—

Neighbourhood of Poona and Aden, None.

Sept. 23. Aden,Campbellpore,and Murree Hills. Road

1886.
Sept. 2.

up Thundiani, near the Kala Pani
Bungalow. Saw a young king-crow,
Dicrwrus ater, stoop at a big blue
Papilio, either P. polyctor or P. arcturus,
and miss it. ‘The bird did not repeat the
attempt.

Campbellpore, Thundiani, ete. Road up

Thundiani, near top of the hill. Saw a
young king-crow stoop at a specimen of
Vanessa kaschmirensis, and after missing
it once take it at the second attempt.
Did not notice whether the insect was
eaten,

360 Mr. G. A. K. Marshall on

Rawul Pindi and home, vid Japan and
America. None.

1888_9.

At home.
1890.
June, Ceylon, Trinkomali. No record.
1891.

Noy. 14. On the Kandy Road between Trinkomali and
Kanthalai; butterflies in great numbers
sitting on the wet mud by the roadside ;
chiefly Pierine (Catophaga), but a few P
nomius with them. These butterflies rose
in clouds as one drove past. A bee-eater,
Merops philippinus, kept flying in front of
my carriage and taking specimens of these
butterflies as they rose. The bird seemed
to select the yellow females, which are
rare, the white females being to them
probably in the proportion of 100 to 1.
These flocks of butterflies often unite and
form what are known as snowstorms in
Ceylon; they then migrate right across
the island.

“These bee-eaters were often seen catching Pierine ; in
fact, it seems to have occurred so often that I ceased to
record the fact, for I can only find this one reference.
Probably the attacks were always witnessed at the begin-
ning of the N.E. monsoons during the time of the heavy
rains, 2.¢. September to December.

“T am not certain as to the date on which I saw the
Ashy swallow-shrike (Artamus fuscus) catching speci-
mens of the Euploea Crastia core. The fact is associated
in my mind with a particular place, and with the capture
of Charaxes psaphon 2 there. This is recorded for April
12th, 1891, so this may be the correct date on which I
watched the bird. At least six specimens of the Crastia
were captured by the shrike, all of which it carried away
to a branch high up in a big tree, but I could not see
whether they were eaten.

“ As regards my experience of birds catching butterflies,
it appears to have occurred more frequently in damp than
in dry districts ; e.g. it was frequent in Ceylon, rare in

The Bionomics of South African Insects. 361

places with moderate or small rainfall, such as Campbell-
pore, Poona, and Aden.

“Tn my opinion an all-sufficient reason for the rarity of
the occurrence exists in the fact that in butterflies the
edible matter is a minimum, while the inedible wings,
etc., are a maximum.”

[See Proc. Zool. Soc. 1887, p. 210, where Lepidoptera
and especially butterflies are spoken of in almost exactly
these terms, as a suggested explanation of the fact that
lizards, although they eat them, greatly prefer flies or
cryptic larvee.—K. B. P.]

14. Recorps oF ATTACKS ON BUTTERFLIES, ETC., BY
WILD BURMESE BIRDS, BY COLONEL C. T. BINGHAM.

[Colonel Bingham has kindly sent me the following
extracts from his 1878 diaries, for incorporation in the
present memoir.—E. B. P.]

“ April 23.—Marched from Kawkaraik to Thinganyina-
ung, fourteen miles. Started about 7.45, rather late as
there was some difficulty in collecting the elephants this
morning. ... The road, a mere jungle path, followed
the course of the Akya Chaung, a feeder of the Haundraw
River, and crossed the little stream some twenty or more
times in the first six or seven miles before turning up the
hill to the Taungyah Pass in the Dawnat Range. From
the outskirts of Kawkaraik right up to Thinganyinaung
on the other side of the pass, the road goes through dense
evergreen forest, and consequently the collecting is very
good on this road, both for insects and birds. To-day, the
day being hot, butterflies, bees, and dragon-flies swarmed,
and at every opening of the Chaung I found crowds seated
on the damp sand apparently sucking up the moisture.
Collecting as I went, it was past 11 o'clock before I got to
the foot of the Pass. I was hot and a bit tired, so I sat
down on a fallen tree to rest, just before crossing the Akya
Chaung for the last time. I had not been seated many
minutes looking at the swarms of butterflies, bees, and
dragon-flies, which were flitting about or sitting on the
sands, when my attention was attracted by a bird, a bee-
eater (Merops swinhoet), which swooping down from a tree
overhead caught a butterfly, a Cyrestis, within a few paces
of me. The bee-eater seemed to catch the butterfly with
ease, and I distinctly heard the snap of its bill. Then
holding the butterfly crossways the bird flew back to the

362 Mr. G. A. K. Marshall on

tree, and sat still for a minute or so, then came a little
jerk of the head, and the wings of the butterfly came
fluttering to the ground, while the body was gulped. On
the same branch some four or five more bee-eaters of the
same species were seated, and as I sat very still, one after
another these birds swooped close to me, sometimes after
a butterfly, sometimes at a bee or a dragon-fly. More
than once I saw a bird miss a butterfly, when the latter
would dodge and try to get away among the bushes of the
dense undergrowth around, but only very seldom was this
successful, for the bird would hover and twist and turn in
hot pursuit, and generally managed to catch the insect.
I was greatly interested, for though I had seen both bee-
eaters and king-crows (Dicrurus) go for butterflies and
moths, this was the first time I had witnessed a continuous
hawking of butterflies on the part of birds. I sat for
nearly half-an-hour watching. The birds seemed to
swoop only for the insects flying about, never at those on
the ground. A drove of pack bullocks with their shouting

5
Shan drivers coming down the road frightened the bee-

fo)

eaters, and they flew off. I got up and prepared to start
up-hill, when it struck me that it would be interesting to
see what species of butterfly had been taken by the bee-
eaters, so I set to work and collected all the loose wings I
could find. I did not get many, for the undergrowth was
very dense, and the wings dropped in it were difficult to
find. Also the place swarmed with ants, I could see them
on all sides carrying off whole wings, or portions bitten
out of them. Again I was pressed for time, so that I
managed to get together only nineteen wings, most of them
odd ones luckily. . . . I have just sorted out and put away
my collections of the day. The butterflies hawked and
eaten by the bee-eaters belong to the following species—
Papilio erithonius, P. sarpedon, Charaxes athamas, Cyrestis
thyodamus, and Terias hecabe. A meagre list, for I am
certain I saw the bee-eaters swoop for and catch Prioneris,
Hebomoia, Junonia, and Precis. I also particularly noticed
that the birds never went for a Danais or Huplea, or for
Papilio macareus, and P. xenocles, which are mimics of
Danais, though two or three species of Danais, four or five
of Huplea, and the two above-mentioned mimicking
Papilios simply swarmed along the whole road.” *

*T did not then realize the importance of my find, or I should
have spared more time for the collection of the fallen wings of the
butterflies, and taken more care of them,—C., T. B,

The Bionomics of South African Insects, 363

Looking through my diaries I find more scattered notes
of my having witnessed birds swoop for and catch butter-
flies and moths, but these were solitary incidents, and
only slight mention is made of them in the diaries with
one exception, which is given below—

“Camp Wabosakhan, December 3. . . . Going
through some fairly open jungle close to the main road
I put up a Melanitis zitenius, which fluttered across the
road and was swooped at by a king-crow (Dicrurus)
but missed; the butterfly dodged, got to the other side
of the road and dropped to the ground among the
herbage and fallen leaves, as is the habit of Melanitis.
The king-crow hovered for a minute not three feet from
the ground over the exact spot where I had noticed the
butterfly drop, failed to see it, flew off, but returned and
again hovered over the spot, but was again unsuccessful,
and flew up to a tree. I went forward very cautiously,
and having carefully noted the spot where the butterfly
had dropped, was enabled to make it out, but not. till
after fully ten minutes of patient and very cautious look-
ing. The Melanitis was there among dead leaves, its
wings folded and looking for all the world a dead dry leaf
itself. With regard to Melanitis, I have not seen it
recorded anywhere that the species of this genus when
disturbed fly a little way, drop suddenly into the under-
growth with closed wings and invariably lie a little askew
and slanting, which still more increases their likeness to a
dead leaf casually fallen to the ground.

“Only once again did I see the systematic hawking of
butterflies by birds. This second occurrence was also by
bee-eaters ; this time it was the large Merops philippinus.
I had been up in the Salween forests beyond the great
rapids, and had managed to get a bad bout of fever which
necessitated my returning to Moulmein, my head-quarters.
It was a hot steamy day in October, and I was lying with
the hot fever fit on me in the boat on the Salween below
Shwegon, when I noticed clouds of butterflies, chiefly
Catopsilia, migrating, crossing the Salween from east to
west in a continuous stream. These were being persist-
ently hawked by the Merops, mixed with which were
some king-crows.”

With regard to Microhierax cerulescens catching butter-
flies, I find the following note :— :

“March 20,1881... . Passing through a taungyah on
my way back to camp [ noticed a number of butterflies,

364 Mr. G. A. K. Marshall on

some seated, some hovering round a spot where some
Karens had been eating their food, and had left some rice
and gnapi scattered on the ground. I was approaching
the butterflies cautiously to see what species were there,
when a small black-and-white bird came down from a tree
close by and perched on the ground close to one little mob
of butterflies busy feeding away on the gnapi. I recog-
nized the bird at once as the pigmy hawk (Microhicracx
cerulescens). His coming flop down close to the butterflies
disturbed some, but not all. A few were too intent on
their meal. The hawk sat for fully two minutes looking
at the butterflies, then he crouched as birds do when they
are about to rise, and next moment with a quick snatch
he had taken a butterfly in his claws and was flying to
the nearest tree. Though I was watching intently J] am
quite unable to say whether he took one of the sitting
butterflies or one that was flying about. I watched him
eat the insect, which he held with his claw against the
branch on which he was seated, and he tore at it just as
the larger hawks do with their prey. I wanted a specimen
of the bird, so shot it, and afterwards picked up the wings
of the butterfly he had eaten; it was a Papilio sarpedon.”

N.B.—That same specimen of Microhierax is now, I
believe, in a small case by itself in the bird gallery of the
British Museum.

[Colonel C. T. Bingham has also made some interesting
observations on the use of insects’ wings as a pad at the
bottom of a hole in a tree, forming the nest of this same
species of bird, the falconet JMicrohierax caerulescens,
Linn. (JZ. eutolmus, Hodgs.). The following account is
quoted from “Stray Feathers” (vol.v, No. 2, June 1877,
pp. 79-81). . The observations were made in the “ Govern-
ment Teak Reserve on the Sinzaway Chaung, a feeder of
the Yoonzaleen River, which it enters about two days’
march below our frontier station of Pahpoon in Tenasserim.”
The nest was found on April 14, “in a hole on the under-
side of a decayed bough of a mighty Pymma tree (Lager-
stremia Flos Reginx).” The four eggs were found to be
“stained by resting on the broken leaves, wings of dragon-
flies, and bits of wood which composed the nest.” The
editor appends to this account a note of Davidson's which
had been in his possession for years. On March 25 the
nest of Microhierax fringillarius, Drap., was examined.
It had been made in a hole in adry tree in an old taungyah
(clearing) “near Bankasoon at the extreme south of Tenas-

The Bionomics of South African Insects. 365

serim.” “At the bottom of the hole, which was about
eighteen inches deep, was a soft pad composed of flies and
butterflies’ wings, mixed with small pieces of rotten wood.”

In March 1878, Col. Bingham found a second nest of
the same species (JZ. cwrulescens) which he sent to the late
Mr. de Nicéville in order to ascertain the species of insects
which had been made use of. Mr. de Nicéville wrote as
follows :—

“The fragments of butterfly wings you send are as
follows :—

No. 1. Portion of fore-wing of Papilio caunus.

, 2. Fore- and hind-wing of J/ycalesis perseus.

, 9. Hind-wing of Papilio erithonius.

» 9. Portion of fore-wing of Junonia orithyia.

» 4, 6, 7, 8, 9, too fragmentary to make out, but
seem to belong to some species of the Lycwnide.

, 10. Half of fore-wing of Chavaxes sp. ?.

, Ll. Portion of hind-wing of Symphedra dirtea &.

, 12 to 17 are the wings of dragon-flies.” *

A passage from another letter of Mr. de Nicéville to
Colonel Bingham indicates in a different manner the
severity of the nearly unseen struggle for existence which
butterflies of certain genera pass through. The wings
sent by Col. Bingham were found by him in 1888, Mr.
de Nicéville wrote concerning them :—

“See p. 275 of vol. 11 of my ‘Butterflies.’ Ferguson found
a single wing of Charaxes schreibert in Travancore on the
ground. It is curious that the only record so far of the

* In the Zoologist (4th Series, vol. v, 1901, pp. 224, 225) Colonel
Bingham states that he found, on April 23, 1899, a nest of the same
species of pigmy falcon in a hole on the under-side of a branch of
a dead tree in a deserted taungyah alongside the high-road leading
from Thabeitkyin, on the banks of the Irrawaddy above Mandalay,
to Mogok, the site of the famous Ruby Mines of Upper Burma. The .
hole had evidently been made by a Barbet. It was 15 inches long,
and at the end was slightly enlarged into an oval chamber contain-
ing ‘‘a fairly firm pad of chips of wood, a few leaves, with an upper
stratum quite two inches thick, composed almost entirely of the
wings of cicadas, with a few butterfly and moth wings interspersed
therein.” There were no eggs or nestlings. “ Further south, in
Tenasserim,” Colonel Bingham continues (l.c. p. 225), “I found the
eggs of this falcon in a precisely similar situation early in April,
as well as I can remember. That nest was composed almost entirely
of butterfly wings.” Colonel Bingham informs me that the last-
named nest was the one, described above in the text, which was found
in March 1878, and furnished the wings named by de Nicéville.

366 Mr. G. A. K. Marshall on ,

same species from Burma should be the three wings you
send me, which you say you found on the ground.”—

E. B.P]

15. Guy A. K. MARSHALL’s INDIRECT EVIDENCE OF THE
ATTACKS UPON BUTTERFLIES. (EH. B. P.)

At the meeting of the Entomological Society held on
August 1, 1883, Professor Meldola communicated some
observations made by Dr. Fritz Miiller in Brazil (Proc.
Ent. Soc. Lond., p. xxiii), together with specimens of dis-
tasteful conspicuous butterflies with wings notched or
otherwise injured apparently by birds. Dr. Fritz Miiller’s
well-known theory, which accounts for synaposematic
resemblances, implies that even distasteful butterflies
are experimentally attacked by young enemies. That
such attacks are made had been doubted, and Professor
Meldola therefore wrote to Dr. Miiller asking him to
collect observations upon the point. A specimen of Heli-
conius eucrate sent by him to Professor Meldola was
described (Ann. Mag. Nat. Hist., Dec. 1882, p. 419) as
having a symmetrical, jagged notch on both fore-wings,
and on Aug. 1, 1883, Professor Meldola exhibited examples
of thirty-six notched and shorn specimens of Acrea [Acti-
note| thalia, obtained in one week by the great German
naturalist. These examples and the Heliconius have been
presented by Professor Meldola to the Hope Depart-
ment, where they may be seen beside numerous similar
specimens from very different parts of the world, includ-
ing those figured on the accompanying Plates IX, X,
and XI. Similar observations upon Bornean butterflies,
including four Danaine, have been published by 8. B. J.
Skertchley (Ann. Mag. Nat. Hist. (6) 11, 1889, pp. 477—
485), while W. L. Distant has described unsymmetrical
injuries, apparently caused by a bird, in the wings of
LIimnas chrysippus (“ Naturalist in Transvaal, ” 1889, p. 65).
I noticed the same thing (1888) in many specimens of
Colias edusa captured in Madeira (“Colours of Animals,”
London, 1890, p. 206; see also Roland Trimen’s Presidential
Address to the Entomological Society of London, Jan. 19,
1898, where many of these and other records are collected
and commented upon).

It seemed of importance to obtain this kind of evidence
from as many parts of the world as possible and on a large
I therefore asked Mr. Marshall if he would kindly

scale.

The Bionomics of South African Insects. 367

look out for specimens of butterflies bearing injuries which
were probably caused by birds or other enemies. The
results, as in every other instance in which I have asked
for his help, far exceeded my most sanguine hopes. He
sent me the fine series of injured specimens represented
on Plates IX, X, and XI.

Looking at the species represented in this collection one
is at once struck with the repetition of the very forms
which have been seen to be attacked by birds (see pp. 357
to 359). Thus Atel/a phalantha, once seen to be mutilated
by a bird (p. 357), is represented by no less than five injured
specimens (Plate IX, figs. 9 and 12; Plate X, figs. 2, 4,
and 5). And nearly every other species observed to be
attacked or found in the stomach of a bird is also repre-
sented, often by two or more examples, in the three
accompanying plates.

The presence of specially-protected forms, Danaine
and Acrxinx, is as conspicuous as in the observations
made in other parts of the world; but new and interesting
light is thrown upon the problem by the exammation of
these specimens and comparison with those of other more
palatable groups. A large proportion of the former (Plate
IX, figs. 1, 5, 7, 10, 11) are far more extensively mutilated
than any but exceptional instances among the latter, and
remarking the peculiar toughness, flexibility, and power of
recovery in the wings of Danainw and Acrvxine, we are
driven to the probable conclusion that the results are in
many cases those of experimental trials by young enemies
and heroic attempts on the part of extremely hungry
enemies, rather than unavailing efforts at the capture of
palatable prey. The futile attempts of hungry animals,
accompanied by extensive mutilation of unpalatable in-
sects, are well known in confinement (Proc. Zool. Soc.,
1887, p. 191), and Mr. Marshall has made observations of
the same kind upon insect enemies in the wild state
(see pp. 318, 358, 359).

The conclusion that butterflies may be pursued when
specially easy to catch, suggested by the observations on
April 26, 1899 (p. 358), is somewhat confirmed by the
curious fact that all the five examples of Limnas chrysip-
pus are females (Plate IX, figs. 1, 5, 10, and 11; Plate X,
fig. 1). :

Of the conspicuous wet phases of the seasonally dimorphic
Precis only a single example is present (Plate IX, fig. 24),

368 Mr. G. A. K. Marshall on

whereas six examples of the cryptic dry phase are included
in the series (Plate IX, figs. 15,19, and 23; Plate XI, figs.
1, 2, and 4). These facts may possibly lend some support
to the suggested interpretation of these remarkable changes
(see pp. 431 to 442).

Some naturalists may be inclined to interpret the in-
juries represented on Plates LX, X, and XI as the ordinary
results of age and wear, or the accidental contact with
thorns or twigs. Such an explanation is not consistent
with the fact that the great majority of the specimens are
in other respects fresh and unworn, and the margins of the
wings not frayed as they become in individuals which have
been long upon the wing. Again, the very high propor-
tion of the injuries inflicted at the anal angle and along the
hind margin of the hind-wing is inconsistent with any such
interpretation. The part of the wing surface which is
certain to come most in contact with foreign objects is the
apical angle of the fore-wing, next, the costal and hind
margins of the fore-wing, last of all the border of the hind-
wing which is behind,and,as the insect finds its way through
an interlacing meshwork of twigs and leaves, is defended
by the greater width and powerful costa of the fore-wings.
It is true that the apex of one or both fore-wings is not
uncommonly snipped off, several examples being repre-
sented on Plate IX, and in the four lowest figures on Plate
XI, but the great majority of the specimens captured by
Mr. Marshall will be found to be injured in the hind-wing,
And of those snipped or notched in the fore-wing, some
exhibit symmetrical injuries which clearly suggest that the
insect was seized with the wings together, probably at rest.
Fizs. 12 and 17 on Plate [IX are good examples. Equally
symmetrical injuries are also common on the hind-wings,
either taking the form of a snip which suggests the very
shape of a bird’s bill (e.g. Figs. 4, 30, 31, 33 on Plate
X), or one in which both anal angles or even a large part
of both hind-wings are shorn completely off (¢. g. Figs. 2 and
28 on Plate X; Figs. 8, 9, 18 and 20 on Plate XI).

In one very interesting example of Vanessa atalanta
from N. Devon, presented by Dr. F. A. Dixey to the Hope
Department, there is only one possible position in which
the injury could have been inflicted, viz. the position
shown in Fig. 31, Plate X, for in that position alone can
the snip in all four wings be made to coincide. Further-
more, the position is that of complete repose, when the

The Bionomis of South African Insects. 369

white patch on the costal border of the under-side of one
hind-wing, wrapping round the front of the costa of
the fore-wing, meets the corresponding patch on the oppo-
site side, and is distinctly seen from the front. ‘The speci-
men captured by Mr. A. H. Hamm, represented in the
adjacent Fig. 33, was probably seized soon after it had
alighted, when the wings were held in the manner indicated
in the figure, and before they were lowered between the
hind-wings in the attitude of repose. Or it is possible
that this specimen was seized during flight at the moment
when the wings came together.

The theory of probability prevents the interpretation of
any but very rare symmetrical notches, except on the
supposition that the wings were together at the time of
the injury, and when the condition of the specimen is
fresh and the notch possesses a definite and similar shape,
fitting that upon the opposite side, there can be no hesita-
tion in inferring the attack of an enemy.

Turning to unilateral injuries, of which many examples
will be found in Plates IX, X, and XI, Mr. Marshall is of
opinion that they are the strongest evidence of the attacks
of birds because they were almost certainly inflicted while
the insect was upon the wing. Perfectly fresh specimens
with such injuries of a very pronounced type are shown on
Plate IX, figs. 15,19, and 23; Plate X, figs. 1, 3, 5, 19, 25,
29, etc.; Plate XI, figs. 4,6, 7,11, etc. It is true that a
butterfly settled upon a flower with outspread wings
might be seized by one side; but insects in that position
are on the alert, and many butterflies when slightly dis-
turbed will shut their wings with a snap when they do
not take flight.

Looking at the injuries as a whole it is seen that the
great majority are inflicted at the anal angle and adjacent
hind margin of the hind-wing,a considerable number at or
near the apical angle of the fore-wing, and comparatively
few between these points, at or near the inner angles of
the wings. I was at first greatly struck by the compara-
tive rarity of injuries in the last position, but in a later
consignment Mr. Marshall forwarded many excellent
examples, referred to in the following paragraph :—

“Salisbury, Sept. 27, 1901.—It was curious that just
after getting a letter from you, pointing out the greater
rarity of mutilation at the inner angles, I came across quite
a succession of excellent examples of this form. The

TRANS. ENT. SOC. LOND. 1902.—PART III. (NOV.) 25

370 Mr. G. A. K. Marshall on

Teracolus omphale is of special interest, as I think the
attack can only have been made by a bird. The same
applies to the Nyctemera, for this sect invariably conceals
its hind-wings when settled, dropping immediately into
this position as it alights. I have occasionally observed
that it holds its wings over its back for a very short time
before closing them. I think the damage to the hind-
wings must be the result of two separate snaps from a bird
while the moth was on the wing.”

The specimen of Teracolus omphale was accidentally
omitted from the illustrations, but the Vyctemera is shown
on Plate X, fig. 8, and three of the best examples of injury
at an inner angle on Plates IX, figs. 15, 19, and 23.

If it be granted that the injuries shown on Plates IX, X,
and XI are chiefly if not entirely due to enemies, the
question as to the kind of enemy remains to be settled.
The only probable foes are birds, reptiles, especially lizards,
and mantides. It is therefore of importance to show that
injuries entirely similar in character to those upon Mr.
Marshall’s South African captures, are also found on but-
terflies from parts of the Holarctic Belt where mantides do
not exist and the attacks of lizards amount to so little that
they may be safely neglected.

I have therefore included (on Plate X, figs. 26 and 28 to
33) the representation of a few butterflies with snipped
wings from the Northern United States, Switzerland, and
England. These are only a selection from a much larger
amount of material of this kind in the Hope Department,
but sufficient to show that the character of the injuries in
the northern land belt is the same as that of those far
south of the Equator, and in a country where lizards and
mantides are very important foes.

Much however may be determined by the character of
the injury and the habits of the butterflies. Such an
injury as that shown on Plate X, fig. 4, for example, is
hardly likely to have been caused by anything but the
beak of a bird. When a mantis seizes a butterfly with its
raptorial legs the wings are instantly crumpled and at the
same time torn and scratched with the thorn-like spines.
Only two or three specimens out of the 82 here figured
bear any such traces, viz. Plate I, figs. 7, 11, and Plate XI,
fig. 5, and in these cases the interpretation is very far
from certain. With regard to lizards, butterflies which
settle on low flowers, and especially those which alight

The Bionomics of South African Insects. 371

on the ground and rocks, are very liable to be attacked,
but in South Africa at least, species which haunt bushes
and trees and fly high are not likely to fall a prey to
lizards, and birds are the only probable enemies when no
traces have been left by a mantis. In the description of
Plates IX, X, and XI, a brief account of the habits of each
-South African species is given by Mr. Marshall together
with the conclusion which appears to be justified.

A very interesting general conclusion emerges after this
consideration and comparison of all the specimens here
figured, viz. the bionomic meaning of important elements
in pattern, and important structural developments of the
wings of Lepidoptera. On Plate IX we see evidence that
injury at the apical angle of one or both fore-wings is
fairly common. Now this angle is very remote from the
vital parts, and no great harm to the butterfly is done by
such injury. And this is a part of the wing which is
constantly rendered specially conspicuous below as well as
above by apical and sub-apical white spots and bars, black
tips, patches of bright colour, and by eye-spots (Plate IX,
figs. 1, 3, 15, 16, 20, 21, 22, 23, 25; Plate X, figs. 3, 8, 19,
25, 28, 32, 33; Plate XI, figs. 4, 21, 22, 23, 24). In the
four lowest figures on Plate XI the conspicuous apical
marking has been injured and, in three cases out of four,
partially or entirely shorn off on one side. This interpre-
tation of the meaning of the apical colour-patches was
suggested by Mr. Marshall in sending these very speci-
mens, and he alluded to two out of the three butterflies
figured on Plate XI, figs. 21-23,in the following passage :—

“Salisbury, June 20, 1899.—I would suggest that these
bright patches of colour [in the orange- and purple-tipped
Teracoli|, which were doubtless first developed by sexual
selection, have been of further use in diverting attack
from the vital parts, and this may perhaps explain their
almost universal transmission to the female sex. I have
sent you two good examples supporting this view, in that
the orange tip of one wing has been snipped off, presumably
by a bird. It should however be noted that the purple
tips are very inconspicuous in flight, and perhaps this might
account for the markedly-swifter flight of those species
which possess them, as they will have thus lost a useful
protection through the action of sexual selection, and have
compensated it by increased swiftness.” *

* Dr. F. A. Dixey points out to me that it is in favour of this

372 Mr. G. A. K. Marshall on

Notches close to and sometimes involving the same
kind of markings are to be seen on Plate IX, figs. 1, 3, 5
10, 16, 19, and 21.

We can thus understand the conspicuous apical mark-
ings of the fore-wings of butterflies, together with the
common prolongation, of the apex of the wing, as directive
marks which tend to divert the attention of an enemy
from more vital parts.

The comparison of Figs. 31 and 33 on Plate X will show
a common method in the use of this marking on the under-
side. It is exposed for a few seconds after the butterfly
alights (Fig. 33), and then hidden by lowering the fore-
wings between the hind (Fig. 31). The meaning is no
doubt that which is suggested on pp. 440, 441, where it is
however applied to the case of those sub- ‘apical eye-spots
on the under-side (Plate X, figs. 28 and 32) which are
exposed and then hidden in a similar manner.

Since the above sentence was written I have consulted
my assistants, Mr. W. Holland and Mr. A. H. Hamm, who
have had great experience in the ways of British Lepi-
doptera, and they both agree with me that our species
of Satyrine with special sub-aptcal eye-spots on the
under-side of the fore-wing are apt to expose these marks
for a few seconds after alighting, and then swiftly cover
them by lowering the fore-wings between the hind. The
“ Grayling,” Satyr us semele, 1s particularly noticeable im
this respect, as all three of us have often observed ;
but the movement is well seen in our other species with
similarly-placed eye-spots.* This characteristic movement
considered in relation to Mr. Marshall’s injured specimens,
and to the experiment with a lizard mentioned on pp. 440-1,
and a kestrel on p. 341, places the hypothesis advanced
on the former pages in a satisfactory position.

But the interpretation of markings and structures at or

interpretation that the females of the species with purple-tipped
males are themselves often orange-tipped.

* Dr. Dixey has specially observed this movement in semele. He
states that Epinephele janira, on the other hand, usually shows the
eye-spot when resting by day, although it quite conceals it when
settling down in the. evening: while a Q EH. tithonus, settled on a
pbramble-leaf in sunshine, exposed the ocellus, but concealed it when
a cloud came over the sun, again uncovering it when the cloud
passed. Dr. Dixey’s notes were made at the time of the observations
at Morthoe, North Devon, in 1897-8.

The Bionomics of South African Insects. 373

near the anal angle of the hind-wing is even more con-
vincing, inasmuch as both markings and structures are far
more specialized and examples of their injury much
commoner. Plate X is entirely occupied with the repre-
sentation of such injuries in species which are without
special directive marks and structures at this region of the
hind-wing, while Plate XI, with the exception of the four
lowest figures (2b to 24) and figure 4, is devoted to the
illustration of injuries received at the very spot which has
been rendered specially conspicuous. In fact, on Plate IX
we have evidence that the attacks of enemies are common
at the apical angle of the fore-wing, and on this Plate as
well as in the four lowest figures of Plate XI the special
directive marks developed at this specially-exposed area
are seen to be shorn off or in some way injured; while on
Plate X we have the same kind of evidence of still more
frequent attack at the anal angle of the hind-wing, together
with, on Plate XI, the evidence that this general tendency
on the part of the enemy is encouraged by the develop-
ment of directive features of all kinds, which are shown to
be successful in that they have been attacked. We see on
Plate XI the prolonged “tails” of the hind-wings of Precis
shorn off (Fig. 1), together with the large eye-spot
marking the same region in Papilio demodocus (Figs. 8
and 9), the two “tails” of Charaxes (Figs. 5, 10, 18, 14, 15,
20), the slender single “tail” with its accompanying single
or double eye-spot of some Lycxnide (Figs. 6, 11, 16, 17),
the conspicuous lobes combined with one or two “ tails’
and bright spots, sometimes in the form of eye-spots, of
other Lycenide (Figs. 3, 7, 12, 18, 19).

Many beautifully “tailed” forms occur among the
Nymphaline of tropical America, such as Protogouius,
Anza, ete., and the commonly de sveloped “tails” of Papilios

J

are probably to be explained in the same manner. When
“tail” is produced at the anal angle of the hind-wing

in relation to a dead-leaf-like under- side, the mid-rib-lke
stripe is developed in relation to the apparent leaf stalk,
as is seen in Plate XIII, figs. 4a, 4b, 6, 7, and 8. On the
other hand, Fig. 1 on Plate XI shows well that such “ tails”
may also act as advantageous directive structures.

The resemblance of the marks and structures at the
anal angle of the hind-wing under-side in many Lycenida
to a head with antennz and eyes has been independently
noticed by many observers. The movements of the hind-

374 Mr. G. A. K. Marshall on

wings by which the “tails,” the apparent antenne, are
made continually to pass and repass each other, add very
greatly to this resemblance. The head-like appearance,
first observed by Dr. Arnold in Thecla iarbas and con-
firmed in other species by Dr. Forsstrona, is quoted by
Kirby and Spence (People’s Edition, 1867, p. 423): it was
mdependently observed by Mr. R. C. L. Perkins (“ Colours
of Animals,” London, 1890, p. 208) in Thecla W-album, and
this keen naturalist obtained confirmator y evidence in the
case of the English Zhecla, similar to that shown upon
Plate XI. My friend Dr. Richard Evans of the Museum
at Georgetown, British Guiana, independently observed
the same thing in Siam, when taking part in the Skeat
Expedition. My friend Professor Wyndham R. Dunstan,
F.R.S., sending mea pair of Deudoriv antalus bred from
larvee which are destructive to the pods of “ Jnga dulcis”
at Manashi, near Cairo, wrote (July 4, 1900) that his
friend Mr. E. A. Floyer who sent the insects “remarks
that the butterfly has markings on its tail which resemble
the head. He considers this protective, as a bird is
uncertain which is the head and which is the tail, and the
insect often escapes by going off in the unexpected
direction.” My friend Mr. Champion B. Russell, who
presented to the Hope Department the beautifully
mutilated specimen of Spindasis natalensis represented on
Plate XI, fig. 3, also independently recognized the same
resemblance (1900), and thought that the lobes with their
two tails passing and repassing each other looked like
jaws opening and shutting. I asked Mr, Marshall’s opinion
on this subject and received the following reply :-—

“ Salisbury, June 11, 1901.—Mzr. Russell’s observations
on the tails of Lycenide are, as you say, of considerable
value as coming from an entirely independent source, but
I must confess that I am not inclined to believe that the
anal appendages in the wings of butterflies have been
modified in imitation of particular organs, for I fail to see
how this could be effected by ordinary selection, And I
think a valid argument against such an idea is the great
diversity of form shown by these appendages, not only
among the Lycenide but other families as well. It seems

safer to regard these curious lobes and tails as having been
developed by natural selection for the purpose of attract-
ing attention to that part, and that the particular form
they take is due to congenital variations which we cannot

The Bionomics of South African Insects. a)

at present explain. Asa matter of fact there seem to be
really very few “blues” in which the tails bear any real
resemblance to antenne. Again, so far as the special
explanation of jaws is concerned it seems to me that this
would rather prompt a bird or lizard to attack the insect
at the other end, which would be fatal. In some cases
there appears to be a possibility of explaining the particular
shape of a tail; for example, in Charaves the general rule is
two thin tails on each wing, and we can understand that
this would not be suitable for the leaf-like under-side of
C. varanes, which has consequently developed a single
thick tail which is more in keeping with its style of
coloration, Again, the thick twisted tail of Wyrina greatly
enhances its general resemblance to a bit of shrivelled
fig-leaf, and so forth.”

I think, however, that it is probable that such resem-
blance as there is to a head, in certain species of Lycwnide
may be of value and may have been produced by direct
selective action, and I would specially draw attention to
Mr. Floyer’s suggestion (p. 374) that the butterfly may
dart off in a direction which the head-like appearance has
caused to be unexpected by an enemy. Many years ago
my friend Dr. A.C. Haddon, F.R.S., showed me a specimen
of a little yellow fish, about 14 inches long, which he had
observed and captured among coral, Aug. 11, 1888, at
Thursday Island, Torres Straits. The head was crossed
by a dark, white-bordered, vertical, somewhat curved band,
which included the eye and tended to conceal it, At the
root of the tail was a very conspicuous eye-like mark.
The fish had the habit of often swimming for a little
distance very slowly tail first, but if disturbed it would
dart off with great rapidity in the opposite direction, viz.
head first. That so similar an adaptation should be met
with in such a very different part of the animal kingdom
affords considerable indirect support to the interpretation
of these Lyczenid marks and structures, at which so many
naturalists have independently arrived. Dr. Haddon
kindly permits me to make use of his interesting observa-
tion, which has not been hitherto recorded. Mr, G. A.
Boulenger, F.R.S., informs me that the fish is Chetodon
plebeius,

376 Mr. G. A. K. Marshall on

16. EXPERIMENTS ON A CAPTIVE MUNGOOSE (Herpestes
galera) WitH INSECT-Foop. (G. A. K. M.)

1899.

May 27. Offered a Lhopalocampta forestan to a very
young female Mungoose (//erpestes galera) ;
she rushed at it, but on touching it with
her nose drew back sharply (her eyesight
was still but feeble); so I partially cut off
its wings and let it flutter on the floor,
whereupon she ran at it several times but
did not attempt to eat it. I then gave her
four 7. senegalensis (without wings) which
she ate greedily, and on being offered a
Mylothris agathina she promptly seized it,
but immediately jumped back so violently
that she rolled head over heels. The
way she shook her head clearly demon-
strated the distastefulness of the butterfly,
and she would not let me bring it any-
where near her. I then offered a Lelenois
mesentina, but with the same result; she
refused to touch it every time. Thinking
this might be due to her experience with
the Mylothris, I put the elenois aside
and offered it ten minutes later, when it
was eaten with undoubted relish. An
Acreva caldarena and A. axina were then
refused, but without being tasted, the
smell being apparently quite sufficient.
More than an hour afterwards I again
tried her with R&. forestan, but she would
not touch it, though whether this was
due to fear of its size and violent fluttering,
or to some unpleasant smell, I could not
well decide; any way she did not attempt
to bite it.

June 8. Gave mungoose two Terias brigitta, two
Teracolus omphale, and two Belenois seve-
vind, All these had their wings cut off
and were thrown on the ground, when
they were promptly seized and eaten. A
Mylothris agathina was then offered in
the same manner, and even this was eaten.

The Bionomies of South African Insects. 377

June 4. Mungoose ate three Terias brigitta, three
T, senegalensis, two Precis sesamus, three
Byblia withyia, one Pyrameis cardui, and
two Mylothris agathina.

7. Mungoose ate three Acrva axvina and actually
one Limnas chrysippus. Whether this
eating of evidently-unpalatable species
is due to the voracious appetite of the
animal or toa youthful lack of discrimina-
tion it is difficult to say, but probably
the former is the truer explanation. I
could not observe any marked signs of
its having found the insects unpleasant.

11. Gave mungoose an Acrva caldarena, which
was promptly eaten. An L, chrysippus
was then thrown down; she seized it at
once, but quickly ejected it with unmis-
takable signs of distaste. An A. axina
was treated in a precisely similar manner,
so that she seemed to have learned
wisdom. Later on she was offered a
Phymateus morbillosus ; she made several
attempts to eat it, but its very unpleasant
smell deterred her each time.

[These results are interesting and in some respects re-
markable. It is probable that some of the apparently-
inconsistent results were due to the fact that a voracious
insect-eater in extreme youth was gaining its first experi-
ence of certain species. Thus the apparent fear of the
large Hesperid Rhopalocampta was probably, as Mr. Marshall
suggests, the inherent timidity of a young animal in the
presence of a strange sound and a method of wing-vibra-
tion very different from anything which it had witnessed
before. The treatment of J/ylothris suggests that the
animal was startled at first by something unusual in taste
or smell, but that when it became accustomed to the
experience the Pierme was no better defended against
the mungoose than against mantides, On the other hand,
the behaviour towards Acreas and chrysippus seems to
indicate a progressive recognition of distastefulness or
unwholesome qualities. It is unfortunate that the experi-
ments were not greatly extended and prolonged.—H. B. P.]

?

zy)

378 Mr. G. A. K. Marshall on

17. EXPERIMENTS ON A CAPTIVE MUNGOOSE WITH Brrp-
FOOD. WARNING CHARACTERS AND DISTASTEFUL
QUALITIES IN SouTH AFRICAN Birps. (G. A. K. M.)

[Although this section is strictly speaking outside the
scope of the present memoir, it is so closely associated
that I have ventured to include it.

Mr. Frank Finn noticed that his mungoose, Crossarchus
fasciatus, “appeared to be unwilling to attack birds, though
it did not seem to find them unpalatable” (“Natural
Science,” vol. i, No. 10, December 1892, p. 746).—E. B. P.]

Salisbury, Jan. 24, 1900.—I have been recently giving
my mungoose some wild birds, and I was much interested
to note the result. It ate a dove (Turtur capicola),
standard wing night-jar (Cosmetorius vexillarius), dwart
goose (Nettapus auritus), moorhen (Gallinula chloropus),
and wheatear (Saxicola pileata); while it emphatically
refused an owl (Asio capensis), kestrel (Cerchnets rupi-
coloides), buff-backed egret (Herodias lucidus), hobby (Falco
subbuteo), and drongo (Buchanga assimilis). Its dislike of
the smell of the common and conspicuous blue-and-black
drongo was very marked, especially as it was hungry at
the time, and I had plucked the bird clean; it made one
or two attempts to eat the meat, but finally gave it up.
In the case of this bird and the egret we would therefore
seem to have acase of true warning coloration. This is
also probably the case with the wood-hoopoes (Jrrisor
and Rhinopomastus), which are very conspicuous both in
voice and colour—the latter being in both genera black
shot with metallic dark-blue or green, with a large white
speculum in the wing, and a long tail.

Salisbury, June 26, 1900.—As to distastefulness in
birds I must further mention our wood-hoopoes, Lrrisor
viridis and Rhinopomastus cyanomelas, both of which emit
astrong unpleasant smell. They are both metallic greenish-
blue birds with long fan-like tails and a conspicuous white
bar on the wings, differing principally in the shape and
colouring of the beak. They are also both very noisy,
frequently uttering their harsh, chattering cries, and
especially when alarmed. Another bird which has well-
known distasteful qualities is the ground horn-bill (Bucoraxz
caffer). Indeed the Zulus use it on that account for rain-
making ; they will kill one and throw it into a river, for
they say its smell makes the river sick, so that it calls

The Bionomics of South African Insects. 379

down the rain to enable it to wash the body away. This
bird is black all over with only the primaries white, and
is so weak on the wing that after three or four flights a
good runner can run it down.

18. EXPERIMENTS ON Cercopithecus pygerythrus.
(G. A. K. M.)

The following experiments were all made at one time,
in February 1902, at Salisbury. The insects were offered
in the following order :—

Psiloptera chaleophorotdes : regarded with some suspicion ;
its head bitten off, and the remainder examined and eaten
cautiously.

Amblysterna vittipennis: was offered and eaten at once.

Praogena splendens: was smelt and at once thrown away.

Dichtha inflata: was cautiously smelt and refused.

Amblysterna vittipennis: was regarded with great
suspicion, carefully examined and then eaten slowly.

Precis sesamus (natalensis form): was received with
suspicion and very slowly eaten.

Precis pelasgis, P. sesamus (natalensis), Byblia acheloia, and
two Precis antilope were then eaten readily, but evidently
not so much appreciated as the beetles.

Acrva halali was then accepted without suspicion, but
when the monkey put it in his mouth, he at once took it
out again and looked at it with the utmost surprise for
some ‘seconds, and then threw it away. He would have
nothing to do with an A. caldarena which I then offered
him.

[Mr. Marshall was greatly struck with the caution and
hesitancy displayed by the monkey, and the evident effect
of distasteful forms in causing suspicion of palatable
species offered immediately afterwards. 1 have noticed
the same thing with the marmoset (‘‘ Colours of Animals,”
London, 1900, pp. 241, 242).

The refusal of the two Heteromerous Coleoptera, the
acceptance of the cryptic species, and the treatment of
the Lepidoptera, are in general correspondence with the
results obtained in other experiments.—E. B. P.]

380

19. EXPERIMENTS ON CAPTIVE BABOONS.

Lepidoptera

ACCEPTED.

P. cardui, R. forestan, J. cebrene,
B. severina, C. florella, T. achine
(apparently with doubt, the first
specimen being rejected at sight but
eaten immediately afterwards,
others eaten at once).

One P. corinneus (taken with
suspicion, pulled to pieces and
thrown away, then picked up,

smelt, and eaten).

Mr. G. A. K. Marshall on

(G. A. K. M.)

Rhopatocera.

REFUSED,

A. halali, A. axina (recognized
and refused untasted), LZ. chrysip-
pus (tasted and rejected), Protoparce
convolvuli (seized eagerly by the
female, but on smelling it she
regarded it with suspicion; she
rubbed it about on the ground and
then pulled it to pieces, smelling
and rejecting each piece).

A P. sesamus G was pulled to pieces and rejected by the
female, but the remains were eaten by the male; each

subsequently ate another specimen.

B. alithyia refused

by female but eaten by male; female ate the second one

offered.

rejected by female at sight,

C. florella eaten readily by both.

A. halali
tasted and rejected by male.

L. chrysippus tasted and rejected by male.

Coleoptera.

ACCEPTED.

Polyhirma xnigma (with doubt
at first) ; Peploptera anchoralis, Jac.
(rejected at first and rubbed on the
ground, but eaten immediately after-
wards) ; five Piezia marshalli (eaten
readily) ; Polycleis longicornis and
P. equestris; Adoretus flaveolus ;
Graphipterus mashunus ; Piezia
selousi; Polyhirma bowecardi and
PP. semisuturata ; Graphipterus
tibialis; Anomala, 0. sp. 3; one
Anonalipus plebeius; Brachycerus
brevicostatus ; Sternocera funebris ;
Psiloptera chalcophoroides ; Peplo-
ptera zambesiana; Micrantereus
carinatus.

REFUSED.

Gyimnopleurus smaragdinus ;
Onitis alexis; Epilachna dreget ;
Anthia thoracica (evidently recog-
nized at once; the baboon seized
and flung it on the ground, rubbing
it violently, with averted head, as
though to kill it); Clinteria infus-
cata ; Praogena festiva ; Chilomenes
lunata; Pachnoda flaviventris ;
Onthophagus gazella; Muylabris
palliata; Diacantha conifera ;
Protxtia amakosa ; Zonitis, sp. ;
Plagiodera thoracica ; Lycus amplia-
tus; L. subtrabeatus ; L. constrictus ;
L. rostratus.

A Mutilla offered twice but
evidently recognized and refused
(Polyhirma enigma eaten im-
mediately afterwards with scarcely
any hesitation). H/etica rufa (with
evident disgust); Malacosoma dis-
coidalis, Jae.; Prionocerus dimidi-
atus ; Decatoma lunata and AZ, tet-
tensis ; Onitis innuus ; Oniticellus
militaris; and Pachnoda rufa all
refused at sight.

The Bionomics of South African Insects. 351

Four Protetia amakosa and four Oxythyrea dysenterica
were refused by female at first but eaten rapidly by male.
The female eventually ate one of the latter, but evidently
in doubt.

The above experiments were conducted at Salisbury in
December 1898.

1899:

Jan. 7. Psilopterx are all eaten readily by baboons.

, 2. Byblia ilithyia eaten by young baboon
(Papioparcarius). A Clythra wahlbergiwas
offered to baboon, but immediately refused
on being smelt (it has a very decided
Coccinella-like odour). The Coccinellid
Epilachna dregei was refused.

, 29. Offered the Longicorn, Ceroplesis fallax, to
the baboon, which smelt it and shook his
head, showing evident alarm when the
insect stridulated; on my pulling off the
head he took the body and pulled it to
pieces, smelling each part, but would not
eat it. I then offered him a large brightly-
coloured Cetoniid, Cvlorrhina loricata; its
strong smell was clearly distasteful to
him, so I put the insect on his hand; he
was much frightened and tried to shake
it off, but could not, owing to its long
claws, and was thoroughly terrified by
the loud buzzing it made as it finally flew
off. Offered Piezia selousi to the baboon ;
I could tell by the way that he tried to
snatch sharply at it that he recognized it,
and when he did seize it he threw it
violently on the ground, rubbing it with
his hands (as is their custom with dis-
tasteful insects). I picked it up and gave
it to him again ; he then ate. the head,
took a bite out of the abdomen and threw
the rest away.

Feb. 1. Another young baboon ate two B, severina
and two B. mesentina with evident relish.
It then refused Onitis alexis at sight and
would not even touch it; An example of
Anomalipus plebeius was then taken, smelt

382

Mr. G. A K. Marshall on

and thrown away, although I can detect
no smell in this species, which, however,
stridulates strongly by rubbing the head
against the thorax.

Feb. 14. Baboon ate one Colias electra and one Byblia

ilithyia, It tasted and rejected first an
A. caldarena, then an L. chrysippus, then
rejected at sight several A. axina and A.
halal.

Gave female baboon two Teracolus achine,
which she ate with appreciation. A MMy-
lothris agathina was taken with some
doubt, the head and thorax were eaten
and a small part of the abdomen, the rest
being thrown away; it was clearly not
much appreciated. An LZ. chrysippus was
tasted and refused. Then an Acrea
anemosa was offered with under-side
exposed to show the bright red and black
markings; it was tasted and refused. A
Precis sesanvus GY was then offered in pre-
cisely the same manner ; the baboon took
it, held it in her hand for a few moments
and then let it fly away without attempt-
ing to smell or taste it. I then cut the
wings off another specimen of the same
insect, and this was promptly eaten with-
out any signs of distaste. This appears
very significant. Then two Limnas
chrysippus and one Acrwa anemosa were
refused at sight without trial; one Meptis
agatha was tasted and neglected, and
two Lelenois severina were eaten.

Baboons ate with great avidity two Ano-
plocnemis curvipes, a large strong-smelling
Hemipteron all brown in colour. They
would have nothing to do with a Mylabris
dicincta or a M. tettensis.

Baboons ate readily four Brachycerus
brevicostatus, one Psamimodes scabratus,
one Chlenius cylindricollis, one Tragi-
scoschema wahlbergi, and also an imago
and nymph of the very large and evil-
smelling Hemipteron Petascelis remipes.

The Bionomics of South African Insects. 383

A Dblepisanis haroldi which was offered to
one was taken, immediately thrown down
without even being smelt; on re-present-
ation it was smelt and tasted cautiously,
found to be palatable, and eaten; there
can be little doubt that it was mistaken
for one of the synaposematic members of
the powerful Lycoid group. A larva of
Precis sesamus was neglected by both
baboons, neither taking the slightest
interest in it.

Feb. 24. Baboons ate greedily two Anoplocnemis
curvipes. As I was taking an Anthia
massilicata out of a box the male baboon
made a snap at it and received a good
dose of its powerful acid in his mouth,
which made him start back in a great
fright, making most comical grimaces ; I
then offered the beetle piecemeal to the
other baboon, and it was eaten with
relish except the anal portion of the ab-
domen, which was rejected. This is of
interest in connection with the immediate
recognition and violent rejection of A.
thoracica by the female in an earlier ex-
periment. One Catopsilia florella and
three Terias brigitta were next eaten.
Then two byblia ilithyia were offered
with the upper-side exposed, and were
eaten with undoubted appreciation by
the female; I then showed her a large
brightly - coloured example of <Acreva
rahirva, also exposing the upper-side, but
she recognized it, merely taking it, pull-
ing its head off and throwing it on the
ground without even smelling it. She
then ate two more &. tithyia, and finally
I offered her an withyia and an Acrea
axina together, both having their wings
closed and being held close to one an-
other; she took and ate the former, but
quite neglected the latter. This experi-
ment tends to show that the general
resemblance which Byblia bears to an

384 Mr. G. A. K. Marshall on

Acrea is not sufficiently close to deceive
a baboon.

Gave baboon a &. ilithyia, which was
eaten without hesitation, then an Acreva
halali, which she rubbed in her hands
as though preparatory to eating, but
eventually threw down without even
smelling it; she then ate another B.
uuthywa.

March 20. Gave baboons three specimens of a
brightly-coloured black-and-yellow spider
(Gasteracantha ornata). They viewed
them with some distrust at first, but
eventually ate them with appreciation.

» 22. Baboons ate one Precis archesia and two
male Hypolimnas misippus, but rejected
a Mylothris agathina. They further ate
two Lb. ilithyia, three Atella phalantha,
and one Precis sesamus G). I then offered
an Acrexa natalica, which was seized,
smelt, and thrown away; another LP.
sesamvus ©) was offered immediately after-
wards and neglected.

,» 26. Gave baboon a Mylothris agathina; she
regarded it with some suspicion, but
eventually ate it very slowly and evi-
dently did not care about it. I then
gave her another specimen; she pulled
the abdomen off, smelt at the exuding
juices, and after tasting them gingerly
with her tongue, threw the insect away.
After this two Belenois severina were
eaten with relish. I then gave her a
Neptis agatha, and she seemed to hesitate
about taking it, and after tasting a portion
of the abdomen neglected it. Of a second
specimen she ate rather more before
throwing it down; the remains I offered
to the other baboon; he tasted it and
seemed in doubt for some time as to
whether it was fit to eat or not, but
finally rejected it.

April 9. Gave baboon a female Hypolimnas misippus,
var, inaria, of which I had cut off the

Feb. 2

Or

The Bionomics of South African Insects, 385

wings; she ate it readily and likewise a
P. sesamus.

April 22. The female baboon ate with relish a male

Hypolimnas misippus and a Precis sesa-

mus @. I then gave her a B. ilithyia,

which she pulled to pieces and threw
down, but then changed her mind, picked
up the thorax and ate it, and immediately
afterwards she ate three more examples

in quick succession without hesitation, A

single Bb. ilithyia given to the male was

pulled to pieces and neglected.

» 23, A Rhopalocampta forestan and R. pisistratas
were eaten with avidity by female baboon ;
she watched with great eagerness while I
opened another box, and was evidently
disappointed when I produced a Belenois
mesentina ; she ate it, however, and an-
other after it. She seemed doubtful
about a Herpenia eriphia, but while
she was tasting it the male made a grab
at it, and I could not decide whether the
insect was really appreciated.

May 1. The female baboon ate a Papilio corinneus
without any signs of distaste, and likewise
a Belenois mesentina. She then took a
H. eriphia, but without eagerness, and
pulled it to pieces, tasting it gingerly,
and finally rejected three-parts of it.
Another &. mesentina was then offered
and neglected.

» 21. Offered female baboon a larva of B. mesen-
tina. She was clearly afraid of it, snatching
it from my hand and throwing it down,
but she soon picked it up again and began
examining it very cautiously; finally she
decided to taste it, and after some con-
sideration came to the conclusion that it
was all right. She then eagerly devoured
over twenty of them, though it was
evident she did not like to feel them
wriggling in her hand. I noticed that
she almost always squeezed out the ex-
creta of the caterpillars before eating

TRANS. ENT. SOC. LOND. 1902.—PART Ill. (NOV.) 26

386 Mr. G. A. K. Marshall on

them. The male was much too fright-
ened to eat them in spite of the example
of the female. I then gave a BP. severina
to the female and a &. mesentina to the
male, both being eaten readily. The
female made a grab at my box, seized a
Mylothris agathina and a B. mesentina
together and began eating them at the
same time; she soon found something
was wrong and dropped them, then picked
up the IMylothris, tasted it and threw it
away, eating the elenois afterwards.
The male, who had meantime eaten an-
other B. mesentina, picked up the dis-
carded Mylothris, but on tasting it also
at once rejected it. Then four Tera-
colus omphale and three 7’. achine were
eaten with undoubted appreciation by
both baboons. On giving each a HH.
eriphia they were both tasted and re-
jected ; the female ate a 7. phlegyas and
neglected another H. eriphia.

There can be little doubt as to the
distastefulness of this latter insect to a
certain extent, but I should be inclined
to rank it rather lower than JMylothris

and about on a par with Neptis agatha.
Finally, six Zerias senegalensis were
eaten without any sign of distaste by
both baboons, though the insects are too
small to be much appreciated by them.
Salisbury, Sept. 21, 1900.—I could not yet say what
protective character is possessed by the Ichnewmonide,
but with the Bracons there can be no doubt that it con-
sists in their having a very strong and unpleasant smell,
somewhat resembling that of the Coccinellidx, only rather
more pungent, They do not attempt to defend them-
selves by stinging, and their flight is slow, rendering them
very conspicuous on the wing. I gave one of the common
red species to my monkey (Cercopithecus pygerythrus) the
other day. He put it straight into his mouth without
smelling it, but soon spat it out again. I then offered
the mangled remains to one of the baboons, but she merely
smelt and rejected them, It is interesting to note that

The Bionomies of South African Insects, 387

the baboons, which eat insects largely, are much more
cautious in receiving any food than is the Cercopithecus.

20. CONCLUSIONS FROM EXPERIMENTS ON CAPTIVE
Bapoons, MUNGOOSE, AND KEsTRELS. (G. A. K. M.)

[Mr, Marshall’s notes on these insectivorous animals were
in one series, which I have analyzed for the purpose of this
memoir. Hence the following references in his letters
deal with the experiments as a whole, except in those
instances in which a particular animal is named.—E. B. P.]

Salisbury, June 20, 1899.—In view of the above experi-
ments 1t seems to me impossible to regard such genera as
Terias, Teracolus, Belenois, Byblia, etc. as unpalatable. I
quite agree with your excellent suggestion that distasteful
characters probably first arose in the larval stage and were
transmitted to the imago, and this view lends further support
to the presumed palatability of Selenois, for my baboon
ate the larvee with much relish. Zeracolus likewise I must
still regard as a non-distasteful genus, at least so far as the
orange- and purple-tipped groups are concerned.

I was much pleased with the undoubted proof of mimicry
[in the experiments with baboons] in the case of Precis
sesamus GD), for Ido not recollect having seen an account of
direct experimental proof before.

With Lyblia I was not so successful as I had hoped, but
I am inclined to attribute the hesitation in accepting this
species, which was observed in some instances, to imperfect
mimicry. I fully recognize the difficulty in distinguishing
in such experiments between Batesian and _ slightly
Miillerian mimics, as either might be received with
hesitation at first, though if subsequent specimens were
eaten readily (as in the case of Lyblia) I should think they
must be included in the former category.

The eager acceptance of the malodorous Coreid bugs by
my baboons came as a very great surprise to me, and
several other results in these experiments point to the
great danger of generalizing on the unpalatability or other-
wise of insects, from the results of experiments on only
one kind of animal.

Salisbury, April 25, 1899.—I may mention that so far
as my experiments go I have no evidence for the unpalat-
ability of TZerias, Teracolus, Belenois, Byblia, Precis, or
Hypolimnas, whereas Mylothris and Neptis are certainly
distasteful to some extent.

388 Mr. G. A. K. Marshall on

21. CONCLUSIONS FROM EXPERIMENTS ON BABOONS.
(Ki. B: P.)

Some of the most interesting results were those which
show the actual working of the principle on which the
theory of mimicry depends ; especially the twice-repeated
experiment with the exposed under-side of the distasteful
Acrea anemosa, resulting in the escape of the Precis
sesamus GJ with a somewhat similar under-side, while the
same species deprived of its wings was readily eaten. At the
same time a P. sesamus EG) was on one occasion rejected by
the female without any preparatory display of an Acraa.
The suspicion of the Lycoid Longicorn, Blepisanis haroldi,
dispelled when it was tasted, points in the same direction.
On the other hand, the baboons were not imposed upon
by the resemblance of the Carabid Polyhirma xnigma to a
Mutilla. In the natural state the swiftly-running Carabid
would have a much better chance of this pseudaposematic
protection than under the conditions of an experiment with
captive insect-eaters. Byblia ilithyia was similarly dis-
tinguished from an Acrwxa ; but this by no means proves
that the resemblance is not beneficial under natural
conditions.

As regards Lepidoptera, the refusal of a Protoparce
convolvuli, after examination of every fragment of it, is
most remarkable. The insect at rest is apparently a
beautiful example of cryptic colouring. Further experi-
ments are very desirable.

The Acrv@ine (axina, caldarena, rahira, anemosa, halalc)
and LZ. chrysippus were never eaten, although sometimes
tasted ; on many occasions they were recognized at sight,
and refused. Mylothris agathina was sometimes partly
and once completely eaten, but it was usually rejected and
evidently unpalatable to them. Neptis agatha was also
generally tasted and neglected, and never entirely eaten.
The Pierine Herpenia eriphia was also evidently dis-
tasteful. Other butterflies which were usually eaten did
not appear to be a food which is much appreciated. Thus
B. ilithyia was sometimes neglected and sometimes only
partially eaten; Zeracolus achine was on one occasion eaten
“with doubt,” and the same was true of one Papilio corin-
neus. The two large Hesperids of the genus Khopalocampta
were, on the other hand, eaten with relish, and the baboon
showed evident disappoimtment on receiving Lelenois

The Bionomics of South African Insects. 389

mesentina after them. Apart from these examples of
hesitation, disappointment, and refusal in part or complete,
the baboons ate many species of Pierinw (Teracolus, Colias,
Terias, Belenois, Catopsilia) and Nymphaline (Junonia,
Precis, Pyrameis, Atella, Hypolimnas), but refused to touch
the larva of Precis sesamus. Considering what has been
already argued about insect-eating animals in confinement,
the acceptances (excluding the Hesperidw) probably do not
justify the conclusion that the Lepidoptera were palatable,
or that they would be sought for in the wild state except
under the stress of hunger. The treatment of the larva
of Belenois mesentina certainly seems to indicate palatability
to the baboon, and also that it was a first experience of
this caterpillar.

The experiments with Hymenoptera are particularly
valuable; for although M/utilide and Braconide are so
greatly mimicked, there is practically no direct evidence
that they are refused by insect-eaters.

The fact that the two large species of Hemiptera were
eaten, in one case “with great avidity,” is a further example
of the failure of a mode of defence which produces a great
impression upon man, viz. the emission of an odour offensive
to us. The suspicion created by the conspicuous spiders
was probably due to unpleasant experiences of insects
with a similar combination of colours.

The numerous experiments with Coleoptera are excep-
tionally interesting. I have below tabulated the results
under A, acceptances, and B, rejections. It is seen that
the species of the following groups were refused without
any exceptions :—Lycidx, Melyridx, Cantharidx, Cocei-
nellide, and Scarabeide, while those belonging to the
Rutelide, Buprestidx, and Curculionide were invariably
accepted. The uniform refusal of five species of Scarabeids
is somewhat surprising, but all of them possessed iridescent
colours. Of the six Cetoniide offered, four were always
refused, two being sometimes rejected and sometimes eaten.
The only Phytophaga eaten were two Clythridx of the
genus Peploptera, and one of these was refused at first.
The cryptic Heteromera were accepted, although one of
them was smelt and refused on one occasion ; the iridescent
species was rejected. Longicornia were eaten, except the
Cantharid-like Ceroplesis, which is probably synapose-
matic (compare p. 396). The Carubidx. were extremely
interesting. The acid secretion appears to be their

390 Mr. G. A. K. Marshall on

undoubted defence against baboons as it was against the
kestrels (pp. 342-3). Hence the small and medium-sized
species were eaten somewhat freely, although with evidence
from time to time that the acid was disliked, and, on the
other hand, a large species with a greater amount of
secretion was avoided, and evidently recognized at sight.
Another one being seized, discharged its secretion into the
baboon’s mouth with immense effect. When however it
was killed and offered piecemeal, all the parts except those
which contained the acid were eaten by the other baboon.
If we allow for the fact that Carabidx, attacked under
natural conditions, have a much larger supply of the acid,
we must admit that they possess a very powerful defence,
and that the meaning of the chief quality which under-
lies the aposematic appearance of the large species is
tolerably clear.

A.—Coleoptera accepted by Baboons.

GROUP. SPECIES. S1zE AND APPEARANCE,
CARABID&. Piexia marshalli. Medium; black with white
lines and patches.
59 seloust (some- Largish ; similar coloration to

times treated as if above.
distasteful and only
partly eaten).
Polyhirma enigma Smallish ; black, white-
(with doubt at first). spotted.
Polyhirma boucardi. Largish; black with white
dorsal thoracic line.
Polyhirma semisutura- Medium; black with white

ti. lines and patches.
Graphipterus mashu- Smallish; black with tawny
NUS. lines and spots.

Graphipterus tibialis. Small; grey-brown, with grey
linear markings.

Chleniuscylindricollis. Medium; green, with yellow
margin to elytra and yellow

legs
gs.
RUTELID&, Anomala, sp. nov. Medium ; testaceous.
Adoretus flaveolus. Smallish ; 5G

Burrestipm. All Psidopterx (includ- Cryptic colouring.
ing the largish P.
chalcophoroides).
Sternocera funebris. Large; shiny black, with
white hairs on thorax.
CuRCcULIONIDA. Brachycerus brevicosta- Large ; eryptic.
tus.
Polycleis longicornis. Largish; black, with yellow
spots on elytra.
= equestris, var. Largish ; iridescent blue-green
with red oblique band on
elytra.

The Bionomies of South African Insects.

GROUP.

HETEROMERA.

PHYTOPHAGA.

LONGICORNIA.

SPECIES.

Psanmodes scabratus.

Anomalipus plebeius
(once refused after
smelling).

Micrantereus carinatus.

Peploptera anchoralis
(rejected at first).

Peploptera zambesiant.

Tragischoschema wall-
bergi.

Blepisanis haroldi.

391

SizE AND APPEARANCE.

Large ; cryptic.
Large; black, often earthy,
and probably allocryptic.

Medium; probably earthy and
allocryptic female, shining
black male.

Medium; yellow with black
spots and bands (Clythri-
dx).

Similar to above.

Smallish ; black with con-
spicuous orange markings.

Small ; Lycoid.

B.— Coleoptera rejected by Baboons,

GROUP.
LycID#.

MELYRID.
CANTHARID.

CoccINELLID &.

PHYTOPHAGA.

CETONIID&.

ScARABHIDA.

SPECIES.

Four species.

Prionocerus dimidiatus.
Zonitis sp.

Eletica rufa.

Decatoma lunata.

>, tettensis.

», dicincta.

» palliata.
Chilomenes lunata.
Epilachna dregei.
Malacosoma discoida-

lis.

Diacantha conifera.
Plagiodera thoracica.
Clythra wahlbergi.

Clinteria infuscata.

Pachnoda flaviventris.
5 ruft.

Protxetia amakosa
(sometimes eaten).
Oxythyrea dysenterica
(sometimes eaten).

Coelorrhina loricata.

Onitis innwus.

Size AND APPEARANCE.

Warning coloration charac-
teristic of African Lycidex.

Lycoid coloration.

Characteristic Cantharid and
Lycoid (c.g. M. palliata,
etc.) coloration, orange and
black.

Characteristic Coccinellid col-
oration.

Medium ; nearly black with
narrow yellow band at apex,
and at base of elytra.

Lycoid coloration.

Largish ; red-brown thorax,

iridescent blue or green
elytra.
Medium; Cantharid type,

orange and black.

Medium ; brown elytra, some-
times black, orange thorax
with two black spots.

Large ; greenish-black with
orange markings.

Large ; thorax green, greenish-
orange elytra.

Smallish ; black with many
small white spots.

Smallish ; shining green or
blue elytra and red thorax.

Largish ; green thorax and
seutellum, yellow elytra
with 4 black spots; brick-
rea head and legs.

Largish ; iridescent
green.

dark-

392 Mr. G. A. K. Marshall on

GROUP. SPECIES. SIZE AND APPEARANCE.
SCARABEIDH. Onitis alexis. Medium ; brown elytra and
legs, iridescent green thorax
and head.

Onthophagus gazella. Smallish ; similar colouring to

above.
Gymnopleurus smarag- Small ; iridescent green, blue,

dinus. or red,

Oniticellus militaris. Small; probably when fresh
iridescent dark-green with
orange markings.

Hereromera. Praogena festiva. Medium ; almost black with
purplish iridescent reflec-
tions on elytra, bluish on
thorax.

CARABID&. Anthia thoracica. Large ; black, white spot on
thorax and white margin to
elytra. Huge mandibles.

ss massilicata Large ; black with pale mar-
(only eaten when gin to elytra. Huge man-
offered piecemeal). dibles.

LONGICORNIA. Ceroplesis fallax. Large ; black, yellow-banded,

Cantharid type:

92. THe CHIEF CONSPICUOUS SPECIALLY-DEFENDED
GROUPS IN THE COLEOPTERA INFERRED FROM
G. A. K. MARSHALL'S EXPERIMENTS. A COMPARI-
SON BETWEEN COLEOPTERA AND LEPIDOPTERA IN
THIS Respect. (&. B. P.)

It is of great interest to attempt to conclude from the
results of the experiments on the palatability of conspicu-
ous Coleoptera contained in this memoir, and from
previously recorded observations and experiments on the
same order of insects, what are the chief specially defended
groups which may be considered to stand in the same
position towards their allies as the [thomiinx, Danaine,
Feliconine and Acrxinx do to the rest of the Rhopalocera,
and the Agaristide, Syntomide, Zygxnide, etc., do to the
rest of the Heterocera.

The chief memoirs upon which the conclusions stated
below have been based are published in the Transactions
of the Entomological Society of London. They are the
papers by Mr. CG. J. Gahan (1891, p. 367), by Mr. H.
Donisthorpe (1901, p. 845), and the Presidential Address of
Canon W. W. Fowler, Jan. 15, 1902 (Proc. 1901, p. xxxiii).
I have also had the opportunity of reading the manu-
script of an important paper by Mr. R. Shelford on mimicry
in Bornean insects, now being published by the Zoological

The Bionomies of South African Insects. 393

Society of London, and of arranging an abstract which is
printed in the Report of the British Association at Bradford,
1900, p. 795.

But, above all, I wish to express my thanks to Mr. C. J.
Gahan and Mr. G. A. K. Marshall for their kind advice and
assistance in the attempt to arrive at sound conclusions.

The groups about which there seems to be no doubt at
all—conspicuous, constantly refused by insect-eaters, and
liable to be mimicked by other Coleoptera—are the
following :—

1. Eroryntip&. Very apt to enter into conspicuous
combinations which are doubtless synaposematic with
other beetles.

2. CoccINELLID&. Greatly mimicked by other beetles
and insects of other orders. Very commonly form synapo-
sematic assemblages (see p. 520).

3. MALACODERMID&, including the Lycinx, Lampyrine,
and Telephoring. Greatly mimicked by beetles of other
families, and also by insects of other Orders (see pp. 515—
518).

4. MeLyRID&. Some of the species convergent towards
Lycine and Telephorine ; others characterized by the
possession of thoracic glands, which are exserted when the
insect is irritated.

5, CANTHARIDa. Undoubtedly a distasteful group
with conspicuous warning coloration. Some of the species
are synaposematic with ‘other beetles, and with Aculeate
Hymenoptera (see pp. 516-518 and 525-527), while
others afford models for mimicry and synaposematic
approach (pp. 518, 519).

6. CHRYSOMELIDA. The sub-families, Galerweine and
Hispinx are especially largely mimicked by other beetles,
and fall into synaposematic combinations. The Chrysome-
line, Humolpine, ete., also enter into combinations which
are doubtless Miillerian (synaposematic). The Megalopine,
however, may be mimetic (pseudaposematic) rather than
synaposematic.

Concerning the last-named family, Mr. Gahan writes to
me, March 3, 1902 :—

“In reference to my previous paper on Diabrotica
[Trans. Ent. Soc. Lond., 1891, p. 367], there are a few
facts since published in a paper by F. M. Webster ‘ On the
probable origin, development, and diffusion of North
American species of Diabrotica,

394 Mr. G. A. K. Marshall on

“<«The fact that several species of this genus are literally
swarming over large areas of country, and their habits are
such as to expose them almost continually during the
adult stage to attacks of birds, while in all of the investi-
gations of the food of birds they rarely appear, has raised
the question of their being inedible.’

“ Webster gives also a quotation from Bates’ ‘ Naturalist
on River Amazon, which I had overlooked :—

“The Eumolpidee and Galerucidze were much more
numerous than the Chlamydes and Lamprosomas, although
being also leaf-eaters, and having neither the disguised
appearance of the one nor the hard integuments of the
other; but many of them secrete a foul liquor when
handled, which may perhaps serve the same purpose of
passive defence.’ ”

There are two other groups which may eventually be
placed beside the six families named above.

ENpomycHID&. Mr. Shelford’s experiments show that
several of the Bornean species are most distasteful. They
are abundant and extremely conspicuous; they form
synaposematic groups, especially with the Hvotylide, and
there is one beautiful example of mimetic resemblance to
an Endomychid model by a Bornean Longicorn. The style
of colouring in the family suggests that it contains
Miillerian groups (see also p. 522).

PyrocHroip&. The colouring and habits suggest that
these Coleoptera are highly distasteful; they may even
belong to the first rank in this respect.

We now pass to a Coleopterous family which may with
more probability be placed beside the Hypsidx or Chalco-
stinv, undoubtedly distasteful groups of moths which
nevertheless are exceedingly apt to display Miillerian
resemblances to other presumably still more stronely-
protected Lepidoptera. In such synaposematic combin-
ations they appear perhaps invariably to take the patterns
and colours of others, rather than impress the stamp of
their own likeness on the assemblage.

CLERIDa&. These beetles are, like the above-named
moths, most apt to take on the appearance of still more
distasteful allies, such as the Lycinw, the Cantharide, the
Galerucine, and, in the genus Allochotes, the Coceinellide.
They are great mimics of JZutillidx, and less commonly of
ants. Mr. Shelford has come across one beautiful example
of the mimicry of common Bornean Clerids of the genus

The Bionomics of South African Insects, 395

Lemidia, by a Longicorn of the genus Daphisia. The
common mimicry of Muéillide may be due to an original
body-form, size, and colouring, which rendered the resem-
blance to such models peculiarly easy and rapid of attain-
ment by selective means. Cleridx also occasionally
possess warning colours of their own, Examples are
found in the genus Zemidia mentioned above, and in an
abundant, bright-red, strongly-smelling South African
species of a genus which is probably new. The latter was
rejected by insect-eating animals (see p. 344).

We now come to four groups which the experiments
here recorded show to be at any rate partially distasteful.
They are often very conspicuous, sometimes from an
aposematic colouring peculiar to themselves, sometimes
from their sluggish movements and size, and the manner
in which they expose themselves or move freely in the
open. They are, however, not as a rule mimicked by
other beetles, and they do not largely enter into synapo-
sematic association with the most distasteful Coleoptera.

SCARABZIDH. The diurnal South African species are all
conspicuous, and freely expose themselves. Their colours,
black, green, or coppery, are all conspicuous against the
ground. on which they are always found. The largest
South African species (Scarabeus femoralis) adopts a
warning attitude when it is disturbed. Many of the
species were evidently distasteful to insect-eaters. It is
possible that their special defence is due to the nature of
their food.

Cetonupa&. The majority of South African species,
including all the larger species, are conspicuous on flowers
or exuding gum. They freely take wing in sunshine, but
are quite sluggish in cloudy weather. Their colours vary
greatly, but very conspicuous and contrasted tints are often
present. Many of the species were found to be unpalatable.

TENEBRIONIDA. Mostly dull browns and blacks, gener-
ally diurnal, terrestrial, and slow-moving. Several species
proved to be distasteful.

Lacriip&. The South African species are often iri-
descent green or purplish, many brown or black. They
are abundant and very sluggish; they freely expose them-
selves in conspicuous positions on leaves, and have a
strong smell, Although but few experiments were made,
it is probable that the whole group 1s distasteful.

The case of the Longicornia is peculiarly interesting.

396 Mr. G. A. K. Marshall on

They may be considered as parallel to the Nymphaline
among Rhopalocera. In both we have a great preponder-
ance of species with cryptic habits and colouring, while
genera mimetic of the most distasteful groups of their
respective Orders are also common. In some of these
examples the mimicry is almost certainly Miillerian, as in
the case of Neptis and Limenitis among the Nymphaline
and Cymatura and Ceroplesis among Longicorns. Further-
more, the Clytinw and Callichromine have been shown by
Mr. Shelford to be mimicked in Borneo on a large scale by
other Longicorns, although the former tend strongly to
mimic Aculeates in nearly all parts of the world. Simi-
larly, there is good reason for thinking that the genera
Neptis and Limenitis, which may be mimetic, are also
themselves mimicked by other Vymphalinz, ete.

There remain the remarkable cases of the Ctcindelide,
Carabidx,and Curculionide, which are probably without any
strict analogy in the Lepidoptera. The two former require
special mention, the last-named are treated separately on
pp. 522-525.

CICINDELIDH. These Coleoptera supply models for
mimicry by a Locustid in Borneo, and Mr. Shelford also
considers that one of them is mimicked by a fly. They
are also known to be mimicked by Longicorns. Many
South African Cicindelide are convergent towards, or
mimetic of, Carabide, especially those of the latter which
are themselves convergent towards Mutillide (see pp. 511-
515). Some of the species have a peculiar scent. On the
other hand, their swift movements and retiring habits are
inconsistent with a high form of special protection.

CARABID#. Mimicked in Borneo by a Locustid.
Probably more strongly defended by the possession of
anal glands than are the Cicindelide, and in the com-
binations between the two families it is seen that the
latter have approached the former, rather than vice versd.
Certain groups of Carabidx form pseud- or synaposematic
combinations with the Galerucine and also with the
Mutillide, The South African smaller and moderate-
sized diurnal species have habits very similar to the
Crcindelidx, but are not so swift. They commonly pos-
sess directive marks indicating the specially-protected
anal region. The largest South African species (Anthia)
have a very large charge of the defensive secretion and
extremely powerful mandibles. They freely expose them-

s

The Bionomics of South African Insects. 397

selves, and are most conspicuous, often possessing a
highly-characteristic warning pattern. ‘They adopt special
warning attitudes, and do not run away when they are
attacked (see p. 510).

These two families may be perhaps compared to the
powerful group of the hawks, which are mimicked by the
feeble cuckoos, and yet, when attacked, are themselves
swift in flight, but can render a good account of themselves
when active defence becomes necessary.

23. EXPERIMENTAL EVIDENCE OF THE VALUE OF THE
TERRIFYING MARKINGS IN Cherocampa LARV#.
(G. A. K. M.)

Salisbury, April 16, 1899.—I offered baboons a full-grown
larva (about seven inches long) of Cherocampa osiris. The
larva is remarkably snake-like, the general colouring some-
what recalling that of the common puft-adder (Lites arte-
tans). "The female baboon ran forward expecting a tit-bit,
but when she saw what I had brought she flicked it out of
my hand on to the ground, at the same time jumping
back suspicionsly ; she then approached it very cautiously,
and after peering carefully at it from the distance of
about a foot, she withdrew in alarm, being clearly much
impressed by the large blue eye-like markings. The male
baboon, which has a much more nervous temperament,
had meanwhile remained at a distance surveying the
proceedings, so I picked up the caterpillar and brought
it towards them, but they would not let me approach, and
kept running away round and round their pole, so I threw
the insect at them. Their fright was ludicrous to see;
with loud cries they jumped aside and clambered up the
pole as fast as they could go, into their box, where they
sat peering over the edge watching the uncanny object
below. After a while the female seemed inclined to descend
to investigate matters again, but owing to the manner in
which they had entangled their ropes she could not descend
without the male, and he very emphatically refused to
move. On concealing the larva I managed to coax them
down again, and then seizing the rope to which the male
was tied, I drew him slowly towards me holding up the
larva in the other hand; he simply screamed in abject
terror, so I let him go, and they retired to their box. The
whole performance was a most remarkable demonstration
of the high value of the terrifying colours in these larvee.

398 Mr. G. A. K. Marshall on

.. Their terror of the insect was most amusing, and
was an eloquent testimony to the great value of this form
of colouring to so bulky a larva. I do not think any one
could now argue that the theory of terrifying coloration is
far-fetched, as I have heard contended. The snake-like
appearance seems capable of deceiving more intelligent
animals than baboons, for it is not long since I received
a box containing a mutilated specimen of this caterpillar
accompanied by a note inquiring, “Is this a snake ?”

[This evidence recently obtained by Mr. Marshall,
added to that already published by Professor Weismann, Lady
Verney, and the present writer (“Colours of Animals,”
London, 1890, pp. 260, 261), leaves no doubt that the
conspicuous eye-spots of Chwrocampa and other large
larvee are really terrifying and do actually alarm their
enemies. The sesults observed are consistent with the
production of a feeling of terror rather than of distaste or
repugnance such as Portschinski supposes to result from
the sight of an ocellated spot. In his remarkable papers
on “Coloration marquante et Taches ocellées” (St. Peters-
burg), this acute and imaginative naturalist states his
belief that ocellated spots represent the appearance of a
drop of warning liquid. He develops this hypothesis
with the greatest ingenuity, and describes and illustrates a
large number of such spots in insects of many kinds. In
some ocellated spots he sees represented the reflection of
the sky in a drop of warning liquid; in others, the dis-
torting effect of gravity upon a drop resting on a
vertical surface: in the sounds made by certain irritated
Mantides, as they display the spots on their raptorial legs,
he believes he hears a representation of the rushing sound
of a warning liquid forced through a fine aperture. My
kind friend Professor W. R. Morfill has given me the
opportunity of learning the remarkable and highly-imagin-
ative views of the distinguished Russian naturalist. On
some future occasion I hope to be able to lay them before
English-speaking naturalists in much greater detail. For
the present I desire to point out that the results obtaimed
by experiment do not support his conclusions, but suggest
in the most convincing manner that terror, such as is
caused by the appearance of a serpent, is produced by the
display of eye-like marks on a large caterpillar, Terror
may be similarly caused by the display of large
ocellated spots on the wings of imagines, while in other

The Bionomics of South African Insects, 399

cases they probably act as directive marks, diverting the
attention of an enemy from the body of the insect (see
pp. 371-5 and 440-1). To the former category probably
belong the remarkable eye-like spots on the tegmina of
certain Harpagid Mantides; for Mr, Marshall writes (1902)
of a South African species: “The eye-like markings on the
wings of the Mantis, Pseudocreobotra wahlbergi, are, I
think, almost certainly of a terrifying character. When
the insect is irritated the wings are raised over its back in
such a manner that the tegmina stand side by side, and
the markings on them then present a very striking resemb-
lance to the great yellow eyes of a bird of prey, or some feline
animal, which might well deter an insectivorous enemy.
It is noticeable that the insect is always careful to keep
the wings directed towards the point of attack, and this is
often done without altering the position of the body.”—

E. B. P.]

24. EVIDENCE OF A SUPERSTITIOUS DREAD OF THE
LARVA OF Cherocampa elpenor, (EK. B. P.)

Professor Weismann and Lady Verney have shown that
the larva of C. elpenor terrifies birds, and I have found that
Lacerta viridis was at first much intimidated, but finally
overcame its fright and devoured the larva. An account
of these observations is given in “Colours of Animals,”
Poulton, 1890, pp. 260, 261. I have recently ascertained
that the larva is regarded with superstitious fear in certain
parts of Ireland. This I owe to the kindness of my friend
Mrs. Nuttall, the American anthropologist, who has drawn
my attention to an article by Mrs. Frances J. Battersby,
of Cromlyn, Westmeath, in “ Knowledge,” vol. 21, 1898,
p. 256, and reprinted in “Public Opinion,’ Nov. 11,
1898, p. 622. The writer quotes the following quaint and
amusing account of the larva, and the sympathetic magic
by which its supposed evil influence is cured, from “A
Chorographical Description of the County Westmeath,”
by Sir Henry Piers, of Insternaght (1682): “We have a
certain reptile found in our bogs called by the Trish the
‘Connagh worm. ‘This is an ugly worm, sometimes as
thick as a man’s thumb, about two or three inches long,
having, as all reptiles have, many short feet, a large head,
great goggle eyes and glaring, between which riseth or
Jutteth forth one thick bristle, in shape like a horn, which

400 Mr. G. A. K. Marshall on

is prominent and bendeth forward about three-quarters of
an inch. Whatever beast happeneth to feed where this
venemous worm hath crept (some say if he do but tread
there) is certainly poisoned, yet may be infallibly cured if
timely remedy be applied; the case is twofold, yet in effect
but one, both proceeding from the very worm itself. Some
there are that take this worm and, putting it into the hand
of a new-born child, close the hand about it, tying it up
with the worm closed in it till it be dead. This child ever
after, by stroking the beast affected recovers it, and so it
will if the water wherein the child washes be sprinkled on
the beast. I have known a man that thus would cure his
neighbours’ cattle though he never saw them. The other
method of cure, which I like much better, is by boring an
augur-hole in a well-grown willow-tree, and in it imprisoning
but not immediately killing the worm, so close by a wooden
peg that no air may get in, and therein leaving him to die
at leisure. The leaves and tender branches of this tree ever
after if bruised in water, and the affected beast therewith
be sprinkled, he is cured. The All-wise and Ever-gracious
God having thus in His Providence ordered it that not only
this venemous reptile, but divers others, and who knows if
not all, did we know the right method of using them,
should have in themselves their own antidotes, that so we
might have a remedy at hand as the poet sayeth :—

‘Una eademque manus vulnus opemque ferat.’”

The authoress also shows that a superstitious dread of the
larva still persists among the Irish folk. In all the deserip-
tions the terrifying eye-like marks have a prominent place.
A “clergyman’s daughter, walking near a ditch, ‘saw her
little dog barking and snapping at a most curious-looking
creature with staring goggle eyes.’” One of the country-
folk said that the creature “had a round head like a cat’s,
and gogele eyes.’ “ He was afraid to touch it, as its eyes
glared like a frog’s, and said it bit or stung cattle, when
their heads swelled up; and a man was once bit on the
leg, which swelled up, and he nearly died.” A labourer,
going to fetch a tin basin from a field, “found a Connagh
sitting in it, glaring at him.” A woman lost one of these
caterpillars which she was carrying on a stick, and was
reproved by her father “for not having killed the Connagh
by smashing it with a stone, ‘as now it would sting the
cattle”” The authoress, who is an experienced collector

The Bionomics of South African Insects. 401

of Lepidoptera, has also shown specimens of the larva
to the country-folk, and ascertained that it was what they
eall the “ Connagh,” so that we are not dependent for the
identification upon the loose descriptions of ignorant
and excited people. She states that there are two models
of the “Connagh ” in the Dublin Museum “studded with
coloured stones, and supposed to have been used as
charms.” * .

Miss Eleanor A. Ormerod in her Eleventh Report, for 1887
(p. 126), also shows that this larva is looked upon “at least
in one district in Ireland as the cause of murrain in cattle.”
Thus “In the course of last year Miss Fleming, writing
from Derry Lea, Monasterevan, Co. Kildare, Ireland, men-
tioned :—‘ There is a very large caterpillar sometimes
found here (I have seen it four inches long), which is said
by popular voice to give the disease called “murrain”
when licked or swallowed by a cow. The people call this
creeping thing a Murrain Worm.” On Aug. 7, 1887,
Miss Fleming sent a specimen which proved to be the
larva of C. elpenor. Another specimen was sent on Aug.
20, 1887, to Miss Ormerod, by Mr. N. Richardson, from the
Estate Office, Castle Comer, Co. Kilkenny.

In the autumn of 1898 (Twenty-second Report, for
1898, p. 72) Miss Ormerod received from Mr, Thomas
Wade, of Newcastle-West, Co. Limerick, an account of the
disease of a cow which “the farmers, not only here, but
all over Munster, seem convinced ... is caused by ‘a
worm.” Although Mr. Wade suggests that they refer to
“a lizard, or something akin to it,” it is almost certain that
we have here another case of the same superstition.

Mr. G. H. Carpenter, B.Sc., F.E.S., of the Science and
Art Museum, Dublin, informs me that in 1901 a police-
constable in Co. Mayo forwarded a larva of elpenor as
“a rare kind of reptile,’ and that a similar description
has been given to him by other country correspondents.
The evidence of alarm and superstitious dread is however
of greater significance than the employment of a word
which is so often used inaccurately.

* Mr. Carpenter informs me that the cylindrical form of these

models and the large size of the caudal horn on one of them suggest
Acherontia rather than Cherocampa.

TRANS, ENT, SOC. LOND. 1902.—PART III. (NOV.) 27

4.02 Mr. G. A. K. Marshall on

95. EXPERIMENTAL EVIDENCE OF TERROR CAUSED BY
THE SQUEAK OF Acherontia atropos. (G. A. K. M.)

[I had been told by Mr, Roland Tiimen that the South
African native races commonly have a superstitious dread
of this moth, and J was anxious to know whether this was
the case in Rhodesia. The observation recorded below
indicates very clearly that the sound and the attitude are
the cause of the fear; for it was inspired in a native who
had never before seen the moth. It is improbable that
the moth is distasteful, but its legs are very powerful, and
the spines on them sharp enough to cause an unpleasant
prick even to human fingers. The behaviour of the
Cercopithecus is strong evidence that the sound possesses a
terrifying significance.—E. B. P.]

Salisbury, Jan. 11, 1901.—1 was deeply interested in
your investigations into the sounds produced by A. atropos,
but I regret to say that I have never gone into the subject
at all. The larva is fairly common with us, feeding on
Solanum and foxglove, but the imago is not often seen.
I showed one to some Mashonas the other day, and asked
them if they knew it. ‘‘Oh yes!” they said, “it’s an
‘imvemvane’ [a general term for butterflies and moths]; it
flies up in the air—whr-r-r-r—and the black man doesn’t
know how to catch it; only the white man can catch it.”
When asked if it were noxious, they seemed surprised and
said, no, not at all. A Zulu replied in much the same
way, and seemed to have no particular ideas about it.
Some Zambesis said they did not know it, and when I
suggested that 1t might be a “schelm” [a Dutch word for
anything noxious or obnoxious], they said no, and one of
them stepped forward and touched it with his finger. But
when it arched its back and squeaked loudly, he jumped
back in a fright saying: “ Oh yes, boss, that’s a ‘skellem ’
right enough.” I finally gave the insect to my monkey
(Cercopithecus pygerythrus), making it squeak while doing
so. He was evidently struck by the sound, and after
watching a few moments grabbed it from my hand, bit off
its head, and threw it down violently. He then approached
cautiously, and began pulling it to pieces in a nervous
spasmodic way, evidently fearing to get stung or bitten
every moment; on tasting one of the bits he found it palat-
able and proceeded to eat it all, I should imagine that
the curious movements and squeak of the moth are of a

The Bionomies of South African Insects. 403

terrifying character, or it may be really aposematic for
certain mammals and birds, to which it may be distasteful.
I cannot find from other sources that the Kafirs here have
any superstitions with regard to it; the only insects they
take any interest in seem to be the various beetles and
larvee which they eat.

26. INSECT STRIDULATION AS A WARNING OR INTIMIDAT-
ING CHARACTER. (G. A. K. M.)

Salisbury, April 19,1901.—I have been thinking of
trying to get some material together to support the view
that stridulation in insects where occurring in both sexes
may be explained in a large number of cases as a warning
character, its value in this respect being especially well
brought out in a number of obscurely-coloured Hetero-
mera, etc., which are known to be distasteful, while it is
largely absent in brightly-coloured, distasteful groups, as
Cetonude, Mylabride, Lycidx, etc. I should also expect
to find it more prevalent among distasteful nocturnal
species, where warning colours are of little avail. One of
my chief difficulties lies in the larval stridulating organs
in Coleoptera referred to by Gahan in his interesting
paper (Trans. Ent. Soc. Lond., 1900, p. 438), and I
should be much interested if you could kindly tell me
whether these larvee really do stridulate, for I see that
Sharp (Camb. Nat. Hist. Ins., Vol. I, p. 198) throws much
doubt on the larval stridulation of Me ne and Scara-
beidxe suggested by Schiddte. Lucanus cervus seems to be
a well- Pernonientcd case, and it would be most interesting
to know whether the larva is distasteful. Darwin’s sug-
gestion as to the acquirement of stridulation by one sex
and its subsequent transference to the other has always
seemed to me unsatisfactory, and its possible warning value
occurred to me immediately I began experimenting with
Coleoptera. Of course in some cases it might be pseud-
aposematic, as in Hymenoptera-like Longicorns in which it
would suggest the shrill, angry buzz of a wasp. Pocock
has already suggested this explanation with reference to
scorpions and Mygale spiders, but I am not aware of any
one else having referred to it.

[For this interesting investigation a piece of apparatus
invented for me by my friend Mr. G. J. Burch, F.R.S., would
be extremely useful. It consists of an ordinary double

4.04 Mr. G. A. K. Marshall on

stethoscope (for both ears) with the usual form of end-piece
replaced by a cork traversed by a glass tube about one-
eighth of an inch in internal diameter, and with its terminal
lip very slightly expanded into a small funnel. If, while
the ear-pieces are inserted in both ears, the open end be
moved about near to a stridulating insect, an extraordinary
reinforcement of sound takes place as the source is
approached, so great indeed that I found no difficulty in
localizing it within a small area. Excessively minute
sounds become clearly audible by the use of this valuable
and simple piece of apparatus. If there be good reason to
suppose that the stridulation of any insect is inaudible to us,
viz. if the structure of its organ and the movements set up
asa result of irritation suggest stridulation, it would be
feasible, I believe, to transmit the vibrations to some re-
cording surface other than the tympanum of the human
ear, and thus to investigate them.

It is usually possible to distinguish readily between the
sounds which are emitted in courtship and those which are
produced on irritation and are probably of a warning or
terrifying significance, inasmuch as they arise from quite
different stimuli and tend to be accompanied by character-
istic movements or attitudes. This latter association is
exhibited by the imago of Acherontia atropos, and the snap-
ping sound made by its larva, due to the movements of the
mandibles, is also of terrifying significance. I have once heard
the epigamic sound of Halias prasinana, but only when
the male was pursuing the female and the whole mode of
flight was subordinated to the ends of courtship. Generally
speaking, any sound produced by both sexes on irritation
or attack, and accompanied by threatening attitudes or
movements (as of the mandibles), or merely violent strug-
gles, is to be interpreted, with a high degree of probability,
as a warning or intimidating character. The decision
between warning and intimidation can only be arrived at
after an experimental investigation into the qualities of
each separate species.

Pseudaposematic sounds are also well known in birds,
especially those which build in holes in trees and hiss like
a snake when disturbed. Many lizards also hiss when
extremely irritated. Professor J. W. Gregory, F.R.S.,
describes a grasshopper at Kurawa which hissed so that
he at first mistook it for a snake (“The Great Rift Valley,”

London, 1896, p. 273).—E. B. P.]

The Bionomics of South African Insects. 405

27. HUMAN EXPERIENCE OF THE TASTE AND SMELL OF
INSECTS AFFORDS UNTRUSTWORTHY EVIDENCE OF
THE EFFECT UPON THE SENSES OF INSECTIVOROUS
ANIMALS. (HE. B. P.)

The idea that human experience would supply a valuable
test as to the palatability or unpalatability of insects to
their natural foes has occurred to more than one naturalist.
Tn Section 7 of his paper (Mém. de la Soc. Zool. de France,
tome vii, 1894, p. 375, § 7), Professor Félix Plateau describes
what he calls the “real taste” (‘‘saveur réelle”) of the
larva, pupa, and imago of Abraxas grossulariata : meaning
by the words “real taste” the impression produced upon
his own palate. After some natural hesitation he tasted
a fine lively specimen of the larva, first cutting it trans-
versely into two pieces. After masticating it sufficiently
long to be sure of the impression produced, he affirms that
it is almost without taste, very slightly sweetish, with
nothing whatever unpleasant about it: “ni nauscabonde,
ni poivrée, ni acide, ni amere, sans arriere-gott, et J ajouteral
méme agréable, rappelant un peu celle des amandes douces
seches ou de la noix de coco.” ‘The skin is however some-
what tough, which Plateau suggests as a possible reason
for the rejection of the larva by certain vertebrate enemies.
The pupa was very similar, but even more trssteless than
the larva; while the abdomen of the moth produced
much the same effect when masticated: “la saveur m’a
paru faible, agréable et analogue a celle de la chenille.”

Plateau considers his experience to be entirely convincing
and decisive as to the real taste of the insect to all in-
sectivorous animals. But it is perfectly obvious that the
only point which has been proved is that a single individual
of an animal not habitually insectivorous has found this in-
sect to be rather pleasant in taste although decidedly insipid.
But this fact does not enable us to judge in any way of
the impressions produced upon the senses of a truly
insectivorous animal. That evidence must be judged upon
its own merits, and, as Plateau appears to consider that he
has shaken it, I give a somewhat detailed account of his
treatment of the subject, and especially of his own valuable
and interesting experiments, some of which were conducted
upon insect-enemies hitherto unobserved and untried from
this point of view. In view of the far-reaching character of
Plateau’s conclusion it becomes necessary to re-examine the

406 Mr. G. A. K. Marshall on

whole of the evidence for the unpalatability of Abraxas gross-
ulariata, especially the larva, which has been chiefly em-
ployed for the purposes of experiment. Professor Plateau
gives a fair and admirable résumé of the work of the earlier
experimenters on the larva, showing that they arrived at
the opinion that it was unpalatable, from the uniform
behaviour of many European insectivorous animals, viz.
many species of birds,* two species of spiders, the common
frog, the tree frog, and many species of lizards. In describ-
ing the last experiments, those of F’. E. Beddard, F.R.S., and
F. Finn, conducted almost exclusively upon non-European
animals (“Animal Coloration,” London, 1892, pp. 149, 164,
165), the conclusion is omitted : “these experiments show
that, with a few exceptions, the caterpillar of the magpie
moth is distasteful to animals.” Although a marmoset
“ate one up quite greedily,” the behaviour of two Cebus
monkeys and a Cercopithecus callitrichus, as described by
Mr. Beddard, is most suggestive of the errors likely to
arise from a too-exclusive study of insectivorous animals
able in confinement to eat but little, or nothing at all,
of a food they would obtain in the wild state in plenty
and variety. These monkeys “sucked at the caterpillar
and threw away the skin after the contents had been
entirely extracted; they paused every now and again to
sniff suspicrously at the caterpillar, but nevertheless they
steadily persevered in munching it.” Mr. Beddard’s

account of the behaviour of non- European birds entirely
confirms the conclusions derived from a study of European
insect-eaters. It is probably safe to conclude that the
species had never seen the larva before, and it is not
surprising that many of them should peck at it. But
although from Mr. Beddard’s statement (Joc. cid. p. 149) it 1s
clear that over a dozen species were experimented upon,
the only bird which certainly swallowed the larva was a
large ground cuckoo (Carpococcyx radiatus). The author

* T am now glad to be able to add the evidence that a sparrow in
the wild state rejected this moth after capturing it. The observation
was made by my friend Mr. G. C. Griffiths, F.E.S., of Clifton, Bristol, a
naturalist of great experience and powers of observation, Mr. Griffiths
writes as follows :—-“ 1 was standing on June 30, 1900, among the
trees on Clifton Down at mid-day, w hena specimen of A. gr ossulariata
flew out from a wych-elm and passed slowly across over a gravel
path. A sparrow darted after it and bit off all its wings, but held
its body searcely an instant, dropping it upon the path, where I
picked it up—a very satisfactory proof of its distastefulness.”

The Lionomics of South African Insects. 407
is also “inclined to think” that a white-eye (Zosterops)
ended by swallowing the insect after masticating it for a
long time. The behaviour of the cuckoo is really con-
firmatory of previous observations, for it has been long
known that our own cuckoo is a coarse feeder, and there
is even evidence that it eats this very species, as pointed
out by our President in the Annual Address for the present
year (Proc. 1901, p. xli). The experiments made by Beddard
and Finn on lizards support previous conclusions : “chewed
and refused by Z. viridis, and disregarded by Zowuwrus and
Amphibolurus ;” but they state that a toad ate the larva.
From their other observations on these Amphibia, it appears
certain that toads are indifferent to modes of defence
which are efficacious against the majority of insect-
eaters; but many more observations and experiments
under various conditions are needed before we can safely
conclude that Abraxas is palatable to these enemies. As
possessing some bearing on the question, it is to be noted
that a toad which had swallowed a caterpillar of Huchelia
jacobex, “ almost immediately threw it up” (p. 159).

We now come to the more recent experiments of
Professor Plateau. He found (§ 5) that eight Cistudo
europea, Which were certainly insectivorous, inasmuch as
they devoured beetles, paid no attention to the larva of
Abraxas, with the exception of one individual which seized
a larva and then abandoned it. Coluber xsculapii and
an Italian variety of Lacerta muralis paid no attention to
them, and if, when one of them opened its mouth, a larva
was inserted, it was immediately rejected. As regards
Amphibia (§ 6), Professor Plateau found that the common
frog acted exactly as Dr. A. G. Butler has described. He
further believes that the frog never again attacks the
caterpillar which it has once seized and rejected, although,
as the larva walks off quite uninjured, the frog sometimes
follows its movements for a few seconds. He then threw
some larve into an aquarium containing a number of
Triton punctatus, many of which seized and “ made efforts
to swallow them, giving, according to their custom, sharp
blows with the head to right and left. But the prey was
too big, or the skin too tough, for the Tritons abandoned
them after a few minutes.” The newt swallows its prey
whole, and has great powers of expansion to meet the
special difficulty of size, so it is far more probable that
these Amphibia were influenced in the same manner as

408 Mr. G. A. K. Marshall on

the frogs and the water tortoises experimented on by
Professor Plateau. He then offered the larve to the
larger Triton alpestris, which paid no attention to them.

Professor Plateau’s experiments on spiders are described
on pp. 325-7, where it will be seen that the larvee were either
neglected or treated as A.G, Butler described by Amauwro-
bius ferox, Tegenaria domestica, and Agelena labyrinthiea.
The imago on the other hand was constantly eaten by
Tegenaria, offered once to Lpeira diadema and eaten with
avidity, killed but rejected by Agelena.

His experiments on Carabus auratus and two species of
Dytiscus are described on p. 330: they indicate clearly that
these predaceous beetles freely attack and devour both
larva and imago of Abraxas.

The above is a résumé of the evidence as Plateau gives
it, although he also includes the attacks of insect-parasites.
These I have quoted on p.338; but the other observations
recorded in the same section of this memoir indicate that
insects with warning colours and distasteful to the (non-
parasitic) enemies of their class are specially lable to these
attacks, so that the results obtained by Plateau in this
section (S 10) of his paper tend, so far as they go, to support
the conclusions he seeks to destroy.

As regards the vertebrate enemies Plateau has supplied
a considerable amount of evidence in support of the con-
clusions of his predecessors. He shows that certain
spiders are probably, and some predaceous beetles almost
certainly, enemies of Abraxas in one or more stages. These
latter facts are most interesting and valuable, but they no
more controvert or throw doubt upon the behaviour of the
generality of insectivorous vertebrates than the admitted
exception of the cuckoo, and the pleasant impression pro-
duced upon the Professor himself. All the theory of
aposematic colouring requires is that the conspicuous form
should be unfit as food for the majority of the enemies
of its class; and this has been abundantly proved in the
case of Abraxas. It is unimportant whether our anthropo-
morphic terms unpalatability, distastefulness, etc., truly
express what an animal feels, if we admit the fact, as we
are bound to do after the experiments, that the majority
of insect-eaters after trial do not eat the insect, and after
one or more trials do not seek to eat it unless driven by
starvation. Exactly what impressions they feel we can
never know, and it is quite unnecessary for the theory

The Bionomics of South African Insects. 409

of warning or aposematic coloration that we should know.
The behaviour is sufficient, and affords convincing proof.
Plateau also brings two indirect lines of argument to
bear on the question. He points out (§ 3) that the larva
is not wholly conspicuous, but requires searching for if it
is to be obtained in large numbers. This is due to its
habit when young of resting along the serrated edges of
leaves, exposing only the reddish lateral band sprinkled with
black points. This appearance is at a little distance very
like the reddish edges of many leaves. When it is older
he observes that it stretches itself longitudinally along the
branches in the inner darker part of the bush at no great
height above the ground. When the bush is shaken it
falls or lets itself down by a thread; and as soon as it
reaches the ground rolls ina ring. In this position, which
it retains for a long time, it resembles the excrement of a
bird. Mr. Beddard too points out (Joc. cit., pp. 167, 168)
that these larvee “like other Geometers . . . do not begin
to feed until evening. I have a quantity of these cater-
pillars on some thick-leaved shrubs in my garden; during
the daytime none are visible, but in the evening they
commence to crawl about quite actively.” I have ‘nyseif
often observed the larva moving and freely exposed by
day on the sides and tops of hedges, but after the state-
ments of these two naturalists I am quite ready to believe
that my observations were exceptional. Indeed on general
grounds I must believe that this is so; for it would not be
an advantage but a great disadvantage to the Abraxas to
display its “aposematic colouring at too creat a distance.
It has certain enemies, such as the cuckoo, and it would
doubtless be dangerous if these were able to see it upon
the bush at a great distance. Its colours would be con-

5
spicuous enough to the majority of insect enemies hunting

the bush for food ; and the very rough resemblance to
leaf-edges, branches, and birds’ excrement would never
impose upon the eyes which enable tieir owners to find
abundant food in spite of the extraordinarily close likeness
to each one of these objects which is attained by so many
of the cryptic species which they hunt.

Plateau’s second indirect argument depends on the
undoubted fact that the pupa of the Abraxas resembles a
wasp. This he regards as an example of (Batesian)
mimicry, and believes moreover that it has never been
pointed out before. This is very far from being the case.

410 Mr. G. A. K. Marshall on

In my paper “The Experimental Proof of the Protective
Value of Colour and Markings in Insects in Reference to
their Vertebrate Enemies,” Proc. Zool. Soc., 1887, p. 191,
I published a Table (A) of the various combinations of
colours of conspicuous insects. Section III of Table A,
on pp. 232, 233, contained the combinations of black and
yellow and of yellow and black; under the former was
placed the pupa of Abraxas grossulariata and several
Hymenoptera Aculeata, ete, under the latter the larva
of Huchelia jacobex and other species. Another Table (B)
grouped conspicuous insects according to their patterns.
Section I of Table B, on p. 236, is as follows :—

I—Ring Pattern.

|

Pattern developed ( Pupa of 4. grossulariata.

on abdominal ~| Imago of Vespa, Nomada, | Alternating rings |
segments. | and Bombus. | of yellow and |
Developed on whole , Larva of /. jacobex. black. |

length of body.

The following sentences refer to this very section of
Table B:—*There is probably in some cases a certain
amount of true mimicry in the acquisition of patterns and
colours. Thus it is more than probable (as has been
previously suggested by other observers) that the species
rendered conspicuous by alternate rings of black and
yellow gain great advantages from the justly respected
appearance of hornets and wasps. It must not be for-
gotten, however, that the latter forms also probably gain
fo some extent by the greater pubheey which follows
from the resemblance ” (pp. 235, 238). Furthermore in
1890 the same conclusions were re-stated in almost the
same words, save that the species indicated in Table B,
Section I, are here introduced into the text :—*“ Thus it 1s
more than probable that the species marked by alternate
rings of black and yellow (including the chrysalis of
the Magpie Moth and the caterpillar of the Cinnabar
Moth) gain considerable advantages, ete.” (“Colours of
Animals,” London, p. 186).

It is hardly necessary to point out that the resemblance
to a wasp is no evidence of palatability, any more than the
resemblance, which is often strongly marked, in the same
district. between wasps of different genera and between

The Bionomis of South African Insects. 411

them and Hymenoptera Aculeata of other sections. The
present memoir contains splendid examples of Miillerian
or synaposematic associations between inedible forms such
as Lycide and stinging Aculeates (see p. 517), and the
resemblance between Abravas and a wasp is probably of
the same kind, as I suggested in 1887.

My friend Professor W. M. Wheeler of the University
of Texas has also tasted a Syrphid fly, Spilomyia fusca
(Loew), which mimics Vespa maculata as well as a smaller
wasp. The fly was “found to have an agreeable flavour,
the alimentary tract of the insect being full of honey.”
Hence in this case Professor Wheeler concludes that the
colours of the fly are “associated with the absence of dis-
agreeable smell and taste, as the generally accepted theory
of mimicry requires” (“Science,” N. §. vol. vi, No. 154,
p. 887, Dec. 10, 1897). Still later in “The Century
Magazine” for July 1901, p. 378, Professor Wheeler
describes another experiment as follows :—<The writer
while riding through the deserts of Wyoming some years
ago was impressed with the day-flying moths (Psewdohazis)
flitting leisurely along near the ground or resting fully
exposed on the glaucous spikes of the sage-brush. . . .
They had black-and-white wings and black-and-orange
bodies. So striking was this case of apparent warning
colour that the writer after much hesitation decided to
ascertain by means of the only available experiment
whether the imsect really possessed the ‘nauseous pro-
perties’ so generally assumed in such cases by writers on
the subject of animal coloration. He dismounted from
his horse and proceeded to masticate the body of one of
the moths. To his astonishment, the little flavour that it
contained was mild and pleasant—one might almost say,
nut-like.” The writer also records that lizards previously
fed on house-flies, aud therefore not very hungry, “ devoured
with evident signs of relish” several of the conspicuous
day-flying moths Alypia octomaculata. Professor Wheeler
concludes “that if every field-entomologist could only
bring himself to repeat the writer’s experiment on one of
many cases of ‘ flaunted nauseousness, and place his taste-
impressions on record, we should in the course of time
have a really valuable body of evidence, for we can hardly
assume that beasts, birds, and reptiles can find things
‘nauseous’ which are quite tasteless or even pleasant to
the human palate.”

412 Mr. G. A. K. Marshall on

I believe, on the contrary, that we are justified in the
opinion expressed in the last words quoted from Professor
Wheeler, and I have proved that we have very good
grounds for maintaining that a conspicuous insect pleasant
to the human taste is rejected with probable signs of
disapproval by many truly insectivorous animals. The
tasting of conspicuously coloured insects by entomologists
in general, recommended by Professor Wheeler, would I
believe be of very little value or more probably of no
value at all as evidence of the likes and dishkes of insect-
eating animals. Carefully conducted experiments upon
such animals, and still better observation of them and
their prey in the field, and the examination of the contents
of their digestive canal and the components of their feces,
are the only means by which trustworthy conclusions can
be reached. We have to deal with a heterogeneous group
of animals, alike in one respect, viz. the specialization of
digestive apparatus and sense-organs to an insect diet.
It is reasonable to suppose that, whatever we may find
in man and other forms not markedly insectivorous, in the
members of this particular group there will be a specially
acute sensitiveness to qualitative differences amongst the
innumerable species from which they select their food.
It is probable that especial keenness for certain species
indicates a high nutritive value, and that the sense-organs
of insectivorous animals enable them to detect and thus
to reject species which would have an injurious effect, or
at the least would be of comparatively low value as food.
There is no reason for the belief that an equal degree of
specialization exists in animals which are-not insectivorous.
Least of all is this likely to be the case in man, with his
comparatively minute olfactory lobes, the central organ by
which there is appreciation of every kind of flavour as
well as every variety of scent.

I have thought it desirable to argue the question at
some length, inasmuch as the two distinguished naturalists
I have quoted, as well as others, find significance in human
experience of the taste of insects. The opposite conclusion
has always seemed to me inevitable from the considerations
set forth above; but if any doubt remains it must be
dispelled by reading the following account of Mr. Marshall’s
experiments upon himself, and coinparing it with those
which he conducted upon insect-eating animals of many
kinds. It is interesting that he should detect bitterness,

The Bionomics of South African Insects. 413

acridity, and a Coccinellid-like smell in certain species, but
even in these the human experience is probably very
faint compared to that of a truly insectivorous animal.
Nor does it by any means follow that a scent which is
excessively unpleasant to man indicates unpalatability to
all or even the majority of natural foes, as in the obvious
case of Anoplocnemis curvipes, the evil- smelling Coreid
bug eaten greedily by baboons (sce p. 382), ‘although
rejected after tasting by a kestrel ae 345). Professor
Plateau and Professor Wheeler will probably reconsider
the significance of their own sense-impressions when they
read below that Mr. Marshall could detect nothing un-
pleasant in the much-mimicked Limnas chrysippus.

“ Estcourt, Oct. 15, 1896.—I have for some time intended
to make experiments on the taste of the Acreas... .
However, my tasting experiments have not helped me
much, for my sense of taste is evidently not as acute as
that of birds and lizards. However, I send you the few
notes I have made in case they may be of interest or use.

“In Acrva nohara, buxtoni, cabira, and Planema esebria, I
was unable to detect any trace of bitterness or acridity,
and they appeared to me quite insipid; indeed, as far as
mere taste is concerned, I doubt whether I could dis-
tinguish them from elenois severina, though their outer
integuments are much tougher. [Mr. Marshall subsequently
found that buatoni does possess a bitter juice, which
exudes freely from the wings when they are cut, especially
at the bases.] This alone is sufficient to prove the un-
reliability of the test, for cabira is most certainly an
inedible species, even if esebria is doubtful. A. encedon
and its var. /ycia exude a yellow juice from the thorax
which is slightly bitter, but not very markedly so. <Acrwa
anacreon and natalica both exude juice in the same
manner, but it seems distinctly more bitter than that of
encedon. A. aneimosa in addition to the bitter juice emits
a very strong smell when pinched, being the only Acrwa
in which I have noticed this, though possibly acara does
the same, as I regard these as the two best-protected
members of the genus. Planema aganice has no smell,
but emits a lot of acrid juice, not only from the thorax,
but also from the antennz and the nervules of the hind-
wings. With regard to the juice-exuding species I may
note that the bitter taste appears to ‘lie only in the
exudation and not to permeate the tissues of the body.

414 Mr. G. A. K. Marshall on

“Tn the few specimens of Amauris echeria that I tried
I found that no juice was emitted, but they had a nauseous
taste and a strong smell, which reminded me somewhat of
that emitted by many Coccinellide, But it was L. chry-
sippus which showed me the futility of trying to arrive at
any definite conclusions from this line of research, for it
emits neither juice nor smell, and I could detect no trace
of any taste, unpalatable or otherwise, but the tissues have
a somewhat soapy feel to the tongue, which I noticed in
A, echeria and some of the Acreas. The same may be
said of IMylothris agathina, though from its conspicuous
colouring, slow flight, and wide dispersal, I feel sure it is
an inedible species.

“ Malvern, Feb. 21, 1897.—Acrvxa horta exudes a bitter
yellow juice from the thorax when it is injured, and this
juice permeates the costa of the fore-wing. The head and
abdomen do not appear to me to have any unpleasant
taste. Trimen refers to their smell, but my smelling
powers are not sufficiently acute to detect it.

“Malvern, May 14, 1897.—Alxna amazoula feigns
death most persistently ; it has an unpleasant taste and
strong smell not unlike that of the Coccinellidx.”

28. Guy A. K. MARSHALL'S PROOF OF SEASONAL
CHANGES IN SoutH AFRICAN BUTTERFLIES OF THE
Genus Precis. (E. B. P.)

A. Introduction.

The attempt will be made in the following section of
this memoir to explain these astonishing changes as due
to the adaptation of a moderately distasteful and protected
genus in two directions—towards conspicuous warning
colours in the generations of the wet season, the time
when insect-food is abundant; towards procryptic conceal-
ment in the pressure and scarcity of the dry season.

Facts which require for their interpretation the hypothe-
sis of adaptation in the direction of conspicuousness will
be brought forward, much use being made of the conelu-
sive proof only recently obtained “by Mr. Marshall, by
breeding the one from the other, that Precis simia is the
wet phase of P. antilope.

The distinct habits and stations of the two phases, their
relation to other seasonal forms of butterflies, the observed

The Bionomies of South African Insects. 415

differences in the insect life of the two seasons, will all be
shown to be consistent with the above hypothesis.

The results of Mr. Marshall’s experiments as to the
nature of the stimulus by which the change is started in
any individual will be discussed, and further lines of
investigation suggested. The much greater size and
weight of the dry phases will be shown to have an import-
ant bearing upon the inquiry, indicating, as it does, that
the phase must be predetermined in the larval stage.

Finally, it will be argued that the facts proved by Mr.
Marshall, although most startling and indeed astounding,
are not subversive of any of the principles of the science
of systematics.

B. Historical.

In his great work on “South African Butterflies ”
(London, 1887, vol. 1), Mr. Roland Trimen describes several
intermediate varieties between Precis natalensis and P. sesa-
mus, and records Mr. F. N. Streatfeild’s capture of the two
butterflies in coitw. He concludes (doc. cit. pp. 231 and 233),
“Tt is only to such occasional unions, and to their fertility,
that the origin of the intermediate examples under notice
can be attributed.”

Mr. Trimen also makes a similar suggestion as to the
intermediate varieties between pelasgis and archesia, which
are also recorded as having been taken in coitw (loc. cit. p.
235).

Mr. Guy A. K. Marshall first published in 1896 the
suggestion that a group of South African butterflies
described and known as different species of the genus
Precis or Junonia were in reality the seasonal phases of a
comparatively limited number of species. He pointed
out, however, that octavia and aimestris (s. l.) had been
previously considered as two forms of a single species by
M. Charles Oberthiir of Rennes (Ann. Mus, Genov. XVI,
1883, p. 721), and also that Mr. C. N. Barker, the distin-
guished Natal naturalist, had been long convinced of the
existence of these seasonal phases, and especially of the
most remarkable case of all, P. sesamaus, and its wet-season
form, natalensis.

Mr. Marshall’s general description of the differences be-
tween the two phases is as follows: “The dry-season form
is smaller, and usually assumes a duller type of colouring
on the upper-side, sometimes of quite a different hue; the

416 Mr. G. A. K. Marshall on

under-side becomes of a general brownish tint, more or
less resembling a withered leaf, the likeness being
heightened by an oblique line running from the apex of
fore-wing to the anal angle of hind-wing, representing the
mid-rib ; also by the marked prolongation of the fore-wing,
so well known in the winter form of Melanitis leda.
Lastly, the ocelli on the under-side are much reduced or
obsolescent” (Trans. Ent. Soc. Lond., 1896, p. 557). I
am unable to understand the opening statement that
“the dry-season form is smaller,’ indeed, Mr. Marshall
criticizes a more general statement of the same kind made
by Mr. C. N. Barker (loc. cit. p. 551). In the very first
species described from this point of view by Mr. Marshall,
Precis tugela, he speaks of “the smaller summer form,”
and the only other Precis in which he speaks of a differ-
ence in size is P. artaxia, of which he uses almost the
same words (p. 561). I have since compared the two
phases of the following species in the Hope Collection, with
results shown below :—

Precis tugela, dry phase larger.
» ceryne, ,, ,, much larger.
acti, am 7

“5 ‘5 53
» pelasgis, ,, » rather ae
» sesamus,, ,, distinetly larger.*
i Lantiulope. 9. much) see
iy LATLALIG. gk . *

See also pages 451 and 456 for the proof by weighing of
the great difference between some of these phases.

Since the above paragraph was written Mr. Marshall
has informed me that the statement was certainly a slip
of the pen, which remained uncorrected, because he was
unfortunately unable to see the proofs of his paper.

Although Mr. Marshall anticipated the results of future
discovery in a truly remarkable manner in this memoir
(Trans. Ent. Soc. Lond., 1896, p. 557), and brought forward
evidence of a most convincing kind, yet the conclusion
which required to be proved was to most naturalists so
highly improbable, because of the extraordinary differences
between the supposed species, that nothing less than the
actual breeding of one form from another was sufficient.

In his second paper on the subject (Ann. and Mag. Nat.

* Difference much less marked in specimens from northern part
of range.

The Bionomies of South African Lnsects. 417

Hist., ser. 7, vol. u, July 1898, p. 30), which it is only
just to describe as eee “making i in the history of-seasonal
dimorphism, Mr. Marshall recorded this great discovery,
and published the fact that he had succeeded in breeding
sesamus from natalensis in three cases. The specimens
themselves Mr. Marshall presented in two cases to the
Hope Collection at Oxford, in the third to the British
Museum of Natural History. The Oxford specimens are
figured on Plate XII, figs. 1 and 2 the parents, figs. la, 10,
and 2a the offspring. The dates of the various stages are
given in the description of the plate.

These specimens must always have historic interest, and
I have therefore published in the plates accompanying
this memoir a representation of the whole of the evidence
obtained by Mr. Marshall in 1898, so far as it is at my
disposal. ‘The extracts from Mr. Marshall’s letters bearing
on the same subject have also historic interest, and are
therefore recorded in full below.

“Umkomaas Mouth, Natal; Sept. 3, 1897.—I am sorry
to say I have never yet bred natalensis through to sesamus.
At Karkloof, Natal, I managed to secure three egos 1
March (just the right month for the purpose of testing
the hypothesis that they are the same species), and one of
the resulting larvze was fully half-grown when I left there
for Malvern, near Durban. I brought them down with
me, as I knew that C. N. Barker had bred nata/ensis from
the larva, and so would know their local food-plant. The
Karkloof plant does not occur at Malvern, and the larvee
utterly refused the Malvern food-plant and everything
else I tried them with; so they pined away and died.

“T have not the least doubt of the specific identity of
these two forms; they are undoubtedly confined respec-
tively to the wet and dry seasons, they have been frequently
observed 72 coitu, and intermediate forms occur at the
change of seasons. The larvee are identical and feed on the
same plant; for out of twelve larvie taken by Hutchinson
off one plant, ten were natalensis and two sesamus. I
always think natalensis is an interesting species as showing
the brilliant colours which can be acquired by an unpro-
tected species without detriment. A newly- -emerged
natalensis is a glorious insect, and rivals the brightest
Acrieas in its colouring on both surfaces ; moreover, it 1s
a frequenter of open country, where its salmon-red wings

TRANS. ENT. SOC. LOND. 1902.—PART III. (NoV.) 28

418 Mr. G. A. K. Marshall on

are a conspicuous object as it sits sunning itself on plants
or stones. But it is very wary and difficult of approach,
being kept on the alert by its enemies, the lizards. I
have often watched these little reptiles stalking both
natalensis and pelasgis round the stones, and have seen
them capture and eat both species.”

“Salisbury, March 6, 1898—You will be pleased to
learn that within another few weeks I hope to have been
able to have solved the natalensis-sesanvus question.
Three weeks ago I obtained five eggs from a female of
typical natalensis ; two proved infertile, one young larva
I lost, but the remaining two are thriving and growing
splendidly. Later on I got three more eggs, which
have hatched successfully. To-day I took one more,
and also, which pleased me much, an egg of typical Precis
simia, which I am convinced is the wet-season form of
P. cuama (Hew.), in spite of Butler's remarks. The
natalensis question I am all the more anxious to settle, as
I have now strong collector's evidence against me, viz.
Distant, who records that he only took one natalensis at
Pretoria, whereas sesamus was abundant and occurred all
through the wet season.”

“Salisbury, June 5, 1898.—You will be glad to learn that
I have at last proved the identity of P. sesamus and natal-
ensis by breeding the former from eggs laid by the latter
in three instances, and I send you the parent and offspring
in two of the cases, the third I am sending to the British
Museum. You may imagine my delight on seeing the first
specimen emerge, for though I felt convinced that the result
would be as I anticipated, yet Distant’s remarks raised
a haunting fear that perhaps I had made a big mistake
after all. However, I am glad to say this was not so.”

C. The Demonstration by Guy A. K. MARSHALL that
Precis simia is the Wet Phase of P. antilope.

Only a few weeks ago Mr. Marshall obtained this further
proof of the soundness of the conclusions he reached, and
the validity of the evidence he adduced in 1896. A
female specimen of P. simia was tracked while she laid nine
egos, on Feb. 23, 1902, at Salisbury. The butterfly was
then captured, and is represented on Plate XII, fig. 3,
and the under-side on Plate XIII, fig. 4. Offspring of

The Bionomies of South African Insects. 419

the form antilope were successfully reared from two of
these eggs. In the case of the first, shown on Plate XII,
fig. 3a (under-side on Plate XIII, fig. 4a), the egg
hatched on March 1, the larva pupated on April 10, and
the imago, a female, emerged on April 27. In the case
of the second, shown on Plate XII, fig. 3b (under-side on
Plate XIII, fig. 40), the egg hatched on March 1, the
larva pupated on April 14, and the imago, a male,
emerged on April 29. All three specimens have been
presented by Mr. Marshall to the Hope Collection at
Oxford. The great difference between the under-sides of
the two offspring (compare Fig. 4a with 45 on Plate
XIID) is deeply interesting. Although so widely differ-
ent, both equally resemble dead leaves, recalling the
various distinct forms of dead leaf represented by the
under-sides of the individuals of the same species of Xal-
lima. The difference between the outline of the wings in
parent and offspring is seen to be far greater in this species
than in sesamus and natalensis, and archesia and pelasgis,
as will be at once seen when the figures on Plate XII or
Plate XIII are compared.

Mr. Marshall’s account of his success in obtaining the
material by which he proved the identity of antilope and
simia, was received in the following paragraph of one of
his letters.

“ Salisbury, Feb. 26, 1902.—I cannot even now agree with
Butler’s arrangement of Precis antilope and cuama. For
although their extreme forms appear to be very distinct,
yet all the chief distinctive characters are unstable and tend
to converge. Aurivillius agrees with me in regarding them
as conspecific, though he separates trimenit and simia. A
pair of these latter I took in copuld last season, which is
sufficient evidence as to their identity in my mind, for lam
very sceptical as to interbreeding in a case such as this.
However, I have determined to solve the problem this
season, and since my return most of my time out-of-doors
has been spent in trying to secure authenticated eggs of
either summer form. Last Sunday I succeeded at last! I
got nine eggs from a single simia, and they will probably
hatch to-morrow ; I hope I shall succeed in pulling most of
them through. All my larvee died in the first stage last
year, for owing to the erratic way in which the females
lay I could not ascertain the true food-plant, but I think
I have it all right this time. I expect to breed both

420 Mr. G. A. K. Marshall oz

antilope and cuama from these eggs, as these forms are
just beginning to appear.”
As explained above, both pupz emerged as antilope.*

D. The Habits of the two Seasonal Phases of the South
African Species of the Genus Precis, and the Stations
which they respectively occupy.

That these butterflies should exhibit a marked differ-
ence in habit andstation corresponding to a difference in
appearance at the wet, as compared with the dry season,
is of such paramount importance in the consideration of the
significance of these phenomena, that I quote at length
all available observations of naturalists—some of them
made before there was any suspicion that such forms as
sesamus and natalensis were the two phases of a single
species.

In “South African Butterflies,’ vol. 1, London, 1887, p.
230, Mr. Roland Trimen, F.R.S., speaks of the habits of
Precis octavia {natalensis, GE) wet phase], as he had seen it
“widely spread over Natal in the summer of 1867, It
frequents open, grassy hills, especially their summit ridges
or highest points, and is very conspicuous, whether flying
or settled.”

Of the habits of P.sesamus © he writes (p. 233): “ Though
constantly to be seen flitting about with its congeners,
octavia, archesia, and pelasgis, 1 have noticed that sesamus
has a greater liking than any of them for shady places,
preferring to settle under a bank or in some deep road-
cutting. ... The very dark bronzy-green under-side is
well adapted for concealment in such spots.”

Of Precis archesia (dry), he notes that it “delights to
bask or repose on rocks or large stones. Colonel Bowker
has noted that it sometimes congregates under rocks, and
is often met with in small rocky caverns in deep forest
kloofs.”. Mr. Trimen has noticed at Highlands, near
Grahamstown, “a habit in the ¢ pelasgis [wet] which I
have never witnessed in the case of archesia [dry], viz.
that of perching himself on the projecting twig of some

* [ Although Lam still of opinion that Dr, Butler is in error im
regarding antilope and cuama as distinct species, yet it may be
pointed out that the results of this experiment do not in any way
disprove his contentions.” —G, A. K. M.]

The Bionomics of South African Insects. 421

high bush at the edge of a wood, and thence giving chase
to other passing butterflies” (p, 237).

In suggesting the seasonal relationships in the genus
Precis which he afterwards proved to exist, Mr. Marshall
speaks at the beginning of his first paper on the subject
(Trans. Ent. Soe, Lond., 1896, p. 557) of the special
interest which attaches i the Sltcnaatione in this genus,
“not only on account of the great differences in the colour
ing of their two forms, but also because of their marked
change in habits.”

He points out that the species of Precis in which there
is but little seasonal change (P. tugela, P. elgiva, and 1.
natalica) are, when at rest, leaf-like in both phases, although
they are more leaf-like in the dry season. He states that
they are furthermore especially forest insects, being
confined to “the warmer, low-lying, or heavily-timbered
districts.” The other species of Precis in which the
seasonal forms are very different “occur more abund-
antly, or even exclusively, in the uplands and in open
country.” There are, however, differences of habit in
the species of the former group. Thus Mr. J. M.
Hutchinson is quoted by Mr. Marshall as stating that the
summer form of P. tugela “is a bolder insect, sailing
around and settling on trees at a fair height, almost after
the manner of Charazes, whereas the other is much more
retiring, keeping among the thick bush and settling low
down, or on the eround among dead leaves, etc.” (lov. Cis,
p. 558),

Turning to the second group of species in which the
seasonal changes are pronounced, Mr, Marshall describes
the habits of P. simia [antilope] (loc. cit., p. 560): “The
dry-season form only frequents the bush, settling on the
ground among the dead leaves, or very rarely on small
plants, the under-side colouring affording it excellent
protection. As the season advances the habits of the
insect change, and in October and November the later
form (c) may be found in company with the early form of
simia (b), frequenting open tops of kopjes, flying boldly
about within a limited area, and settling with expanded
wings on shrubs and bushes. This is the habit of all
summer forms of Precis... .’ With regard to P. sesamus ©,
Mr. Marshall remarks that it “differs from the normal
type of winter Precis in the absence of leaf-like colouring
below, and in the very slight faleation of fore-wings. This

422 Mr. G. A. K. Marshall on

is accounted for by its different habits, for instead of
frequenting dead leaves in the bush it prefers the dark
rocks on stony and wooded kopjes.” Evidence is brought
forward to show that P. artaxia only exists in the cryptic
dry phase in the warm timbered coast belt, although it
develops a much less well-concealed wet-season phase
(nachtigaliz) in open country (pp. 561, 562).

In his second paper on the subject (Ann. and Mag. Nat.
Hist., ser. 7, vol. u1, July 1898, p. 30) Mr. Marshall gives a
further account of these interesting differences in habits :
“Speaking broadly, the natalensis form frequents the
highest points in any neighbourhood, especially if they be
more or less open (for it is anything but a sylvan insect) ;
whereas the sesaimus form is more partial to shady spots,
and is to be found in ravines and spruits or rocky wooded
slopes, and shares with the Hesperid Sarangesa climinata
a marked affection for disused mine-shafts and cuttings.
This distinction must not be taken too strictly, for true
sesamus is occasionally found in company with the summer
form on open hill-tops, but principally at the change of
seasons; but natalensis, so far as my experience goes, is
never to be found in the more shady stations frequented
by sesamus. The latter, moreover, is distinctly warier and
more difficult of approach when not feeding, and if alarmed
flies off with a rapid, and often zigzag, flight, settling
abruptly among rocks or herbage, when its greenish-black
under-side colouring is equally protective. Sesamus 1s
more often observed in gardens, and not unfrequently
enters human habitations in search of a shady resting-
place.” Mr. Marshall also quotes Mr, J. M. Hutchinson’s
experience on his farm in Natal, lying in an open plain
between two ranges of hills: “He has found sesamus
resident on the farm, occurring fairly commonly along the
banks of the spruits during the winter, whereas natalensis
is very much scarcer and non-resident, the examples seen
having always been travelling from one range of hills to
the other, on both of which it is common.”

Since his return to England, in the present summer,
Mr. Marshall has summarized his experience of the habits
and stations of the seasonal phases of the species under
discussion as follows :—

“There are three types of stations in South Africa which
may be occupied by butterflies of this genus.

I. Forest country, with heavy timber affording deep

The Bionomies of South African Insects. 423

shade. Found on the coast belt and also in the interior,

but,

south of the Zambesi, only in patches,

Il. Woodland country, without timber. Trees small,
affording light shade.

III. Open country, without trees. Nothing higher than
small scrub,

I.
le

The following species are only found in the forest :—
P. tugela. It has been already explained that the
wet-season form exposes itself more freely (p. 421).
Both phases have leaf-like under-sides, but the dry
has a more hooked tip to fore-wing and more pro-
longed anal angle to hind-wing,

. P. elgiwa. No ditference in habits observed. The

slight differences in appearance are due to a more
hooked tip with a somewhat different direction, and a
more leaf-like under-side in the dry phase as com-
pared with the wet.

. P. natalica. No difference in habits observed, both

phases being low settlers. Ocelli and white spots on
the under-side tend to disappear in the dry phase.

II. The following species are found in the woodland
country :—
1, and 2, P. antilope and P. actia. In both these

species the dry-season phase and the female of the
wet are found in the more shady places, viz. the lower
slopes of kopjes. The ¢ of the wet phase is usually
found on the less-wooded higher slopes.

P. artaxia. The habits are like those of the two
preceding species. Towards the forest belt in Umtali
the dry phase encroaches on the wet, and in the low
country between Umtali and the sea Selous never saw
the wet form at all. The larger dry form has a far
more leaf-like under-side, with a mid-rib, and hooked
tip to the fore-wing, wanting in the wet. The dry
phase is also more wary.

I]. and HI. The following species are found in open as

well

as woodland country :—

1, and 2, P. sesamus and P. archesia have very similar

habits. The dry phase generally frequents the wooded
and the wet phase the open country. In entirely
open country the former would occur on the lower
slopes in whatever shade is to be found. In entirely
woodland country the dry phase would be found on

424 Mr. G. A. K. Marshall on

the lower, more shady slopes, the wet on the upper
slopes where there is less shade.
III. The following species only occurs (in Mashonaland)
in open country :—
1. P. ceryne. Both forms are found in open swampy
districts, without bush. Rare.”

HE. Evidence of Adaptation in the conspicuous under-sides of
the Wet-Season Phases almost equal to the proof of it in
the cryptic Dry Phases,

The evidence of adaptation in the cryptic under-sides of
the dry phase in the species under discussion is so clear
and so generally admitted that it is unnecessary to say
much about it. J will only point to the manner in which
the various distinctive elements of this phase are co-
ordinated to a common end, that of concealment. Thus
in the dead-leaf-like forms such as archesia and antilope
we find the prolonged anal angle of the hind-wing, the
produced and bent apex of the fore-wing, the angulated
outline between these points, the stripe representing a
mid-rib, the colours and patterns varying in different
individuals but always resembling some type of dead leaf
with discoloured blotches or eaten into holes (archesia).
Most important of all there is the co-ordination of all
these diverse elements with appropriate habits and the
choice of an appropriate station. In archesia, which com-
monly frequents rocks, the intensely variable mottled
appearance produces a strongly eryptic effect at a little
distance, while a close inspection only brings out the details
which produce a graphic representation of a dead leaf. In
sesamus the outline and under-side differ from those of the
dead-leaf-like dry phases of the other species, and differ in
a direction which is specially cryptic, because of the peculiar
habits of this phase of the species (see pp. 420, 422).

All this will be at once admitted by every naturalist
who studies the specimens, as it 1s proclaimed by all who
have had the advantage of observing the species in the
wild state. What is not admitted, but is I think almost
equally clear, is the fact that adaptation in the opposite
direction, viz. the direction of conspicuousness, is characteristic
of the under-sides of the wet phase. If the under-side
merely reproduced the conspicuous pattern of the upper-

The Bionomics of South African Insects. 425

side of the wet phase the case would be strong and con-
vincing for adaptation, and an interpretation based on the
principles of warning colours or mimicry, Batesian or
Miillerian. But the under-side does more than this; it
differs from the upper-side, and so far as it differs, it
becomes more conspicuous. The following details render
the case for adaptation in the direction of conspicuousness,
as it seems to me, overwhelming.

In comparing the upper- and under-side of the wet
phase of the species to which the chief attention of
naturalists has been directed, it 1s of special interest to turn
to the accurate descriptions of Mr. Roland Trimen, F.R.S.,
written long before Mr. Marshall’s discovery was thought
of, a time when natalensis and sesamus were not only
considered distinct but were even separated by ceryne. In
“Rhopalocera Africe Australis” (London, 1862-66) we read,
on pages 130, 131, of Junonia octavia [ Precis sesamus@)]:
“ UNDER-SIDE.—Much paler, more creamy in tint, with a
glistening pinkash tinge.’ Again, on the under-side of
the fore-wing the author speaks of “the row of spots
parallel to hind-margin commencing distinctly from costa,
the first two spots increasing the number’ to seven ;
double row of bluish lunules more conspicuous than on
upper-side, whiter.’ And the under-side of the hind-wing
is thus described: “Basal black containing four rather
large, very conspicuous spots of the ground-colour, and dusted
with blue scales, which form a transverse streak between
costal and subcostal’ nervures near extremity of black ;
whitish-bluish lunules, in hind marginal border, large and
very conspicuous.” The fact that the ground-colour of the
under-side 1s much paler than the salmon-red of the upper
and thus affords a far more effective contrast with the
black markings is seen when Fig. 1 on Plate XIII is com-
pared with Fig. 1b on Plate XII. Figs. 1 and 2 on Plate
XII represent worn specimens, and the comparison with
them is invalid. Fig 1), Plate XII, possesses unusually
heavy black markings, but the representation of the depth
of the red ground-colour is normal for a fresh individual, as
is that of the under-side in Fig. 1, Plate XIII. The more
complete row of black spots and the ereater conspicuous-
ness of the border, owing to the larger white and blue
markings in it, as described by Trimen, are also well seen
when the fioures are compared, but allowance must be

426 Mr. G. A. K. Marshall on

made for the fact that the border represented in Fig. 1,
Plate XII, is exceptionally broad, and is still more excep-
tional, indeed transitional towards sesamus, 1 the size of
the blue markings init. But the lighter character of the
markings and the more conspicuous appearance of the
under-side border is perfectly clear in Fig. 1, Plate XIII.
And there is one other point not expressly mentioned
although probably implied by Mr. Trimen, which is I
think the most convincing evidence of all in favour of
adaptation in the direction of conspicuousness ;—the fact
that the spots of ground-colour included in the basal black
patch of the hind-wing, and absent from the upper-side,
are distinctly lighter im tint than the rest of the ground-
colour, and thus afford a far more effective contrast with the
black. 'This difference in tint is well seen in Fig. 1, Plate
XIII. The spot in the basal black of the fore-wing which
represents a similarly-placed spot on the upper-side, is
also often lighter than the rest of the under-side ground-
colour, but the difference is far less marked than in the
hind-wing and is sometimes absent.

Now the basal area of the under-side of butterflies’
wings and especially of the exposed hind-wing is a part
specially seized upon by natural selection for the display
of conspicuous warning characters. It is seen in the red
patches of many Pierine genera, especially the distasteful
Delias (appearing also in ‘its Chaleosid mimics) in the Old
World, and several Pievine in the New, where Dr. F. A.
Dixey has shown that the character has probably been
adopted by Heliconinx in Miillerian association with them,
the relationship—an important discovery first made _

1894 by Dixey—being one of Bees assimilation ”

“ diaposematic resemblance ” (Trans. Ent. Soc. Lond., 1894,
pp. 296-298 ; 1896, pp. 72-74; 1897, pp. 326, 327, 331;
Proc. Ent. Soc., 1897, p. xxix). A stripe of bright yellow
or red bordering the basal part of the costal margin of the
under-side of the hind-wing of a large number of distaste-
ful tropical American butterflies of different sub-families
is another very characteristic synaposeme, rendering the
same part of the wing especially conspicuous. And in
Africa itself we have the most remarkable case of all, in
the triangular golden-brown, black-marked synaposeme
which is discussed at some length on pages 488 to 490
of the present memoir. Furthermore, there is the group

The Bionomics of South African Insects. 427

of large black spots on a light ground which renders this
part of the wing prominent in such large numbers of
Ethiopian butterflies.

In the wet phase of Precis sesamus this area is also
remarkably conspicuous, but by a method which is as
positive to negative in relation to other distasteful butter-
flies inhabiting the same part of the world, viz. by the
appearance of light spots on a black eround, instead of
black spots on a light eround,

Thus it is improbable that this particular element in the
conspicuous appearance of the under-side of the wet phase
of P. sesamus can be mimetic, and its existence, side by
side with a general resemblance in colour and pattern to a
large Acrwa, is evidence that such resemblance is Miil-
lerian or synaposematic rather than Batesian or pseudapo-
sematic. This argument is much strengthened by the
discussion of the wet phase of the allied P. archesia (see
pp. 428-430).

Much that has been said of P. sesamus applies with
greater force to the closely-allied P. antilope. On compar-
ing the under-side of the wet phase, shown in Fig. 4, Plate
XIII, with the upper-side of the same specimen, shown in
Fig. 3, Plate XII, it is at once seen that the difference in
tint of the ground-colour and in conspicuousness of the
marginal band on the two wing surfaces is far more pro-
nounced than in the species which has been just described.
In fact, with an upper-side which is much less conspicuous
than sesamus GB (compare Figs. 3 and 10, Plate XII: it must
be remembered that the ground-colour of 3 is merely
tawny, while that of 1/ is salmon-red), the wet phase of
antilope combines an under-side which is distinctly more
conspicuous than that of the corresponding form of the
allied larger species (compare Figs. 4 and 1, Plate XIII).
The increased conspicuousness is “especially clear in the
relative size of the spots in the basal black patch and the
inclusion in it of a very large piece of the ground-colour of

t=)
the fore-wing. Here too the increased lightness of the
spots of ground-colour in the black area is often distinct,
as it is in Fig. 4, Plate XIII, but in a large proportion of
the individuals I have had the opportunity of examining
it is only feebly marked.

In P. antilope @ there is probably some considerable syn-
aposematic approach towards the Acrva type, but to a less
extent than in /P, sesamus; while the conspicuous basal

428 Mr. G. A. K. Marshall on

character which is non-acrseiform and purely aposematic
is far more emphasized than in the latter species. It is
probable that sesamus represents a later development, and
that im it the synaposematic clements have been gradually
strengthened and the peculiar aposematic character cor-
respondingly reduced.

We now pass to the consideration of a species in which
the conspicuous characters of the under-side of the wet
phase are probably entirely peculiar and aposematie.

Mr. Marshall’s suggestion in 1896 that Precis pelasgis is
the wet phase of P. ‘archesia has never been confirmed by
breeding the one form from the other. It is, however,
certain that his conclusion was sound. The two forms
have often been captured in ae The female pelasgis
represented in Plate XII, fig. 4, was captured by Mr.
Marshall in coitw with the male archesia shown in Fig. 5 of
the same plate. Intermediate forms are much commoner
than in the case of seswmus and antilope; and above all the
relationship of wet phase to dry is far closer in archesia, so
that it is possible to see how the one was derived from
the other more fully thanin any of the species with mark-
edly-different seasonal forms. The under-side of one of the
commonest forms of the dry phase is represented in Fig. 6,
Plate XIII, and opposite toit that of the typical wet phase
in Wig. 5. At first sight they appear totally different, and
certainly the latter is as conspicuous as the former is well
concealed. An uncoloured illustration cannot do justice
to the varied shades of brown and grey on the under-side
of archesia (Fig. 6), and a long series of specimens would be
required to show the immense range of individual variation
by which all kinds of common appearances presented by dead
leaves are reproduced. Among such variations is one in
which the dark-brown eround- -colour is almost uniform
and unmottled imside the mid-rib-like stripe (Fig. 7). From
this we pass to forms in which the stripe widens into a
light band (Fig. 8), clearly showing its homology with the
still more conspicuous band of pelasyis (compare Figs, 5 and
8). Such a variety as that shown in Fig. 8 is still a long way
on the archesia side of a form intermediate between the
wet and dry phase, and would certainly be cryptic rather
than conspicuous in nature, although not so well concealed
as the form shown in Fig. 7, and still less so than that
shown in Fig. 6. Truly intermediate varieties between
the wet and dry phases are not uncommon, in which

The Bionomics of South African Insects. 429

the broad band becomes sharply defined on its outer border,
but lacks the light tint of the full wet phase.

These considerations and the careful comparison of Figs.
5 to 8 on Plate XIIT will show the essential nature of the
changes by which the eryptic under-side of the dry phase
is converted into the startlingly conspicuous under-side of
the wet phase, or vice versdé. The mid-rib-like stripe
widens, lightens in tint, becomes sharply defined along its
outer border, and is now the “ warning band” of pelasgis.
The row of ocellated spots, many of which, with sem1-
transparent white centres and specially-coloured borders,
suggest holes in the apparent leaf of archesia, become
entirely or almost entirely black upon the hind-wing, and
gain intensely black borders upon the fore-wing of pe lasyis,
and, placed upon the light ground of the “ warning band,”
render this feature still more conspicuous. The mottling
disappears, and the ground-colour, both within and without
the borders of the “warning band,” becomes an almost
uniform very dark brown, forming a most effective contrast
with the band. Finally, the dead-leaf-like margin of the
wing of archesia is rendered conspicuous in pelusgis by a
black-and-white fringe and two parallel series of light
markings just within and parallel to the much less deeply
indented outline.

Furthermore, the comparison of Figs. 6 to 8 on Plate
XII with Fig. 5 on Plate XII shows clearly enough that
the under-side of the dry phase of archesia differs from its
upper-side in being cryptic, while the comparison of Fig. 5
on Plate XIII with Fig. 4 on Plate XII shows that the
under-side differs from the upper-side of the wet phase
(pelasgis) in being more conspicuous, thus in both respects
acting like the two phases of seswnus and antilope. The
under-side of pelasgis is more conspicuous than its upper-
side because of the increased lightness and greater sharp-
ness of the borders of the band and the greater contrast
afforded by a darker ground-colour, also because of the
more pronounced light marginal markings.

I have described the relationship between the phases of
archesia at some length, because it was the consideration of
this species which first convinced me of the validity of the
interpretation here set forth, that we have convincing
evidence of natural selection acting in two opposite direc-
tions in the two phases—in the one to produce the maxi-
mum of concealment, in the other a very efficient form of

430 Mr. G. A. K. Marshall on

conspicuousness. When I realized that it was the mid-
rib-like stripe—the character which more than any other
gives meaning to the cryptic resemblance to a dead leaf;
that it was this very character which, transformed into the
“warning band,’ became the conspicuous feature of the
wet phase—the operation of natural selection seemed as
clear in the one case as the other.

When we examine the species of the whole genus
Precis and those of the genera allied to it, the conclusion
is forced upon us that the dry cryptic phases are ancestral
as compared with the conspicuous wet phases. I do not
mean to imply that the cryptic forms have not altered,
but that the original form of the species possessed a cryptic
under-side, which has been handed down with more or less
change as the cryptic under-side of the existing dry phase,
while the conspicuous under-side of the existing wet phase
is a new and comparatively recent development. This
question of the relative age of the two forms is most im-
portant and interesting, and from the very first occupied
Mr. Marshall’s attention. Thus the following passage is
extracted from a letter written a few weeks after his
discovery :—

“Salisbury, June 5, 1898.—I should be most interested
to learn your ideas as to the reasons for the singular
seasonal change in this species, for I must admit that I
cannot arrive at any really satisfactory conclusion on the
subject as yet. The blue scales of seswmus are my chief
stumbling-block, for I certainly cannot perceive what
utility they can possess, and considering its protective
under-side colouring there seems no reason why it should
not have retained its wet-season colours above, as in the
case of P. artaxia or P. ceryne. I suppose you will agree
that sesamus is a later development ?”

For the reasons I have indicated above it is difficult
to doubt that the cryptic character of the under-side
of sesamus is ancestral and the conspicuous under-side of
natalensis relatively recent, but with regard to the upper-
sides this conclusion is by no means so "evident. Indeed
on comparing the species with anti/ope and other allies, it
seems probable that the upper-side of natalensis is more
ancestral than that of sesamus, having been chiefly modified
in tint, thus falling into Miillerian association with the
larger Acrzeas. The upper-side of sesamus probably
shows cr yptic changes in the acquisition of the far darker

The Bionomics of South African Insects. 431

colours which render the phase less conspicuous in the
stations it frequents,

There does not seem to be any escape however from the
conclusion that the conspicuous under-sides of the wet phases
are relatively recent, and if this conclusion be considered
in relation to the comparison between the under-sides of
archesia and pelasgis, it leads inevitably to the conclusion
that the conspicuous appearance of the one has been
modified out of the older cryptic appearance of the other,
and not vice versd.

On what hypothesis can we believe that such a change

has taken place? In the existing state of our knowledge
there are only two possible interpretations: (1) that the
modification is mimetic of some other conspicuous dis-
tasteful form; (2) that it is a warning of some special
protection possessed by the Precis itself. The former
interpretation cannot apply to the case of pelasgis, because
its pattern is so unlike that of the well-known distasteful
Ethiopian Rhopalocera, although some advantage may be
gained by Miillerian association with black and white
aposematic genera such as Amauwris, Neptis, Planema, ete.
Furthermore, it has been shown that there are important
elements in the conspicuous under-sides of the wet phases
of sesamus and antilope which are not synaposematic,
although the appearance as a whole is probably to be
explained in this way. I therefore firmly believe that the
conspicuous appearance of pelasgis has been produced by
selection from the cryptic a7chesia as a warning character
indicative of some special protection, an aposeme pro-
claiming that it is less palatable or in some way less suit-
able as the food of insect- eating animals than an immense
number of other species which abound during the wet
season in the same stations.

I proved in 1887 (Proc. Zool. Soc., p. 191) that the likes
and dislikes of insect-eating animals are purely relative,
and that a conspicuous distasteful form will be freely
eaten under the stress of hunger, that the existence of
these forms depended entirely upon the co-existence in
their neighbourhood of an abundance of palatable species,
that under any other circumstances the warning colours
if freely exposed would be a danger and would lead to the
extermination of the species. As soon as I had studied
the case of archesia and pelasgis I felt convinced that the

432 Mr. G. A. K. Marshall on

extraordinary seasonal phases of Precis were to be inter-
preted along the lines suggested in 1887—that we have to
do with a set of somewhat distasteful species which can
only exist in the keen struggle of the dry African winter
when food palatable to insect-eaters is relatively scarce, by
a very high standard of protective disguise associated with
the appropriate instincts, but gain the recognized advan-
tages of aposematic colouring’ by producing markedly
conspicuous generations during the moist suunmmer, when
insect- -eating animals have a much greater variety and
abundance of suitable food.

F. The severity of the Struggle for Existence among Insects
in the African Dry Scason as compared with the Wet.
The relation of the Seasonal Changes in Precis to
those of other Butterflies.

As soon as the idea expressed in the concluding para-
graph of the last section of this memoir occurred to me, I
wrote to Mr. Marshall asking for his experience on the
subject, and also inquiring w hether any of the admittedly
unpalatable African butterflies exhibited seasonal changes,
such that the winter generations became comparatively
inconspicuous,

His deeply-interesting reply is printed iz extenso below.

“Salisbury, Jan. 8, 1899.—As to your query about the
keenness of the struggle for existence at the two seasons,
in my own mind I had never felt any doubt that the dry
season is certainly the more eritical period for insects, and
this I referred to incidentally in my paper on Precis (Ann.
and Mag. Nat. Hist., ser. 7, vol. 11, July 1898, p. 36). It
is true that insectivorous birds are far more numerous
during the summer, but this I think would be more than
outbalanced by the increase of such insects as Coleoptera,
Hymenoptera, Diptera, etc., apart from the fact that the
summer broods of the perennial butterflies are undoubtedly
larger and much more numerous (some Pierinw have a fresh
brood every four or five weeks), and that a number of
additional species make their appearance at that season
only. On the other hand, during the dry season, although
a number of migratory birds depart northwards, yet we
have a considerable number of resident insectivorous birds,
including rollers, drongos, shrikes, flycatchers, bush-king-

The Bionomics of South African Inseets. 433

fishers, ete., and owing to the warmth of the midday sun,
even in mid-winter, the lizards are always more or less
active, and the insectivorous mammals are probably in no
way reduced. With the insects it is very different ; owing
to the parching up of the vegetation the hosts of phyto-
phagous insects disappear almost entirely, and the diminu-
tion in insect-life is enormous, being most noticeable
among the Coleoptera and least so among butterflies, of
which latter almost two-thirds have winter broods; and
moreover their lives would be rendered even more pre-
carious by the generally adverse conditions of their
environment from climatic causes. It therefore seems
clear that the struggle for existence would fall pretty
severely on butterflies during the winter, owing to their
general conspicuousness, and that such is actually the case
is shown by the numerous instances of the development of
a highly-protective under-side coloration during the dry
season among Satyrine, Nymphaline, Lycenide, and
Pierine, That the strugele i is sufficiently keen, however,
to compel unpalatable species to adopt protective color-
ation I should not like to say. The following is, I take it,
a complete list of the South African genera possessing
more cr less undoubted distasteful qualities: Limnas,
Amauris, Acrva, Planema, Pardopsis, Neptis, Pentila,
Alxna, Mylothris, and Pontia hellica, and with the ex-
ception of Acrwa none of these exhibit any change of
colour durmg the winter which can possibly be construed
as protective. Dealing therefore with Acrea, I find that
even in this genus a considerable number of species such
as horta, neobule, anemosa, acara, encedon, cabira, ete.,
exhibit only a comparatively insignificant seasonal dimor-
phism or even none at all. There remains therefore a group
composing such insects as violarum-asema, nohara-halali,
petrea, doubledayi-axina, atolmis, buxtont, ete.,in which the
dimorphism is fairly strongly marked in one sex or the
other, and an interesting feature about this group is that
they are all, with the exception of petra, frequenters of
open country, having a low flight and frequently sitting on
the ground, It is also noticeable that this group, unlike the
other, presents a very marked difference in the sexes, and
wherever this is not the case, both sexes have a distinctly
obscure coloration as compared with their congeners, e. g.
axina and asema; further, that where the summer males
exhibit any exceptional brilliancy, as petrea, atolmis, or

TRANS. ENT. SOC. LOND. 1902.—PART II. (NOV.) 29

4.34 Mr. G. A. K. Marshall on

nohara, it is always compensated for by an exceptional
dulness on the part of their respective females. I fear I
do not feel sufficiently competent to attempt an explan-
ation of the above facts, but I think you will agree that
as a whole they hardly bear out the suggestion that
distasteful species are compelled to adopt protective
colouring in winter through the keener struggle for exist-
ence; and for the present I am tempted to incline to the
view that the less marked cases of dimorphism may be
attributable to purely climatic causes. The colouring of the
other open veldt Acrvex, viz. halali, axina, and asema, is
somewhat puzzling; for in the two latter it is far from
being very brilliant or conspicuous ; in halal, the male in
summer is very britliant, but the blackish or brownish
grey of the female is certainly protective, and the insect
when alarmed is very hard to follow with the eye in its
low dodging flight over the herbage. In the winter the
colouring of both sexes of all three species is certainly not
very conspicuous among the withered grass. Either their
unpalatability must be of a low order, or else they must
be more subject to attack by some particular enemies than
the woodland species. I should not be surprised if the
rollers, of which we have five species, or cuckoos (also
five) were to eat Acreas, as they are all far from particular
as to their diet.”

This hypothesis concerning certain of the smaller
Acrzeas had been in Mr. Marshall’s mind for a long time.
Thus he wrote in 1896 from Natal :—-

“ Hstcourt, Oct. 15, 1896.—I have an idea that all the
species of the genus Acrwa are not protected equally by
nauseous taste, etc., and some of them perhaps not at all;
for in many of the smaller species there is a marked
seasonal dimorphism which has clearly a protective value.
Now such a change seems hardly in keeping with warning
coloration, which must be constant to impress itself on the
minds of enemies, and moreover a species which requires
protection by seasonal dimorphism cannot be very much
protected in other ways, not to mention the fact that its
colouring cannot be both warning and protective at the
same time. There is, of course, nothing to show how
much of the seasonal change we can attribute to climate
alone. For instance, in comparing the slight alteration in
an A. aeara with the marked change in female A. petraa,
are we to suppose that the dark-grey female of the latter

The Bionomies of South African Insects. 435

is due solely to climatic influence, and that such a change
in the former is checked by the necessity for keeping the
warning coloration uniform, or are we to consider that the
slight change in the fer is all that climate can effect,
and that in the case of petrea this slight climatic effect
has been enhanced by some other cause—presumably
protection? Personally I incline to the latter view, but in
either case it is clear that there are varying grades of
protection by distastefulness in the genus.”

An extract from another letter states the same im-
portant conclusions as to the severity of the struggle
during the dry season.

“ Salisbury, March, 10, 1898.—There are very few butter-
flies (South Ninicank at least), exclusive of the admittedly
protected species and their mimics, of which the bright
colour cannot be explained on the Zeracolus-Kallima
basis. The most evident exceptions are Byblia, certain
species of Precis, as sesamus (form natalensis), ceryne, etc.,
which are practically coloured the same below as above,
and Belenois severina and mesentina. The first I will
admit has been so far a stumbling-block to me, though I
am not yet prepared to accept it. as a protected species.
Provided that my ideas on seasonal variation in Precis be
correct, these would also fall under the same heading as
Teracolus, for like them they only assume the protective
under-side colouring during winter, when attacks from birds
are no doubt a great deal more to be feared, owing to the
almost complete absence of easi/y-caught prey, such as
beetles and other small insects; the summer forms
probably are very little molested by birds, owing to their
ereat agility and alertness, and the profusion of other
insects at that period; they do, however, not unfre-
quently fall a prey to the rock- lizards, which stalk them
with much astuteness, as I have observed on several
occasions.

“With regard to Lelenois my mind is still open, for it is
a very curious genus, containing as it does the above two
species which might perhaps from certain considerations be
considered protected, and at the same time a species like B.
gidica which evidently comes under the Teracolus heading,
and lastly B. thy sa which, to my mind at least, is clearly 4 ei
Batesian mimic.’

After a consideration of the evidence “brought forward
above, it will be generally admitted that the struggle for

436 Mr. G. A. K. Marshall on

existence is far keener in the dry winter season, and that
butterflies are especially subject to it.

The most distasteful forms, many of which are the
models for mimicry, are sufficiently protected to retain
their conspicuous aposematic appearance throughout the
year, and either exhibit no change in the winter season or
a change which is not in any way cryptic.

While this is true of all the larger and most conspicuous
Acreas, some of the smaller Acrzas do exhibit changes
in a cryptic direction in their winter generations. These
are Acrzas which, from their colouring and habits, may
be inferred to possess only a moderate degree of unpalat-
ability as compared with the other species of the genus.

Cases in which colouring is “ warning and protective
[procryptic] at the same time” are quite common, e. g.
the protected larvee of many Zenthredinide which har-
monize sufficiently well with their food-plant to be con-
cealed at a little distance, but assume the most conspicu-
ous aposematic attitudes and movements as soon as they
are discovered and disturbed. But in the case of the
smaller Acrzeas suggested by Mr. Marshall, the colouring
which is most procryptic does not occur at the same time
as that which is less procryptic or probably aposematic.
Mr. Marshall’s numerous experiments upon the edibility of
the smaller Acrieas (see Sections 9, 18, 19) do not support
the view that any of them are palatable to the insect-
eating animals made use of. It has already been pointed
out that the refusal or evident dislike of insect food by
captive animals is trustworthy evidence of unpalatability,
while acceptance is not proof of palatability (see p. 317).
The smaller Acrzeas furthermore fall into beautiful syn-
aposematic groups (see pp. 492, 495); indeed a strong
Miillerian association can be recognized throughout almost
the whole of the Ethiopian representatives of the genus,
as was first suggested by Professor Meldola (Ann. and
Mag. Nat. Hist., ser. 5, vol. x, 1882, p. 425).

It is therefore probable that these smaller Acreeas are
still specially protected, although to a less extent than
other species of the genus, but “that the keener struggle
of the dry season has compelled them to produce gener-
ations which are inconspicuous as compared with those of
the wet season.

If these interpretations here suggested be correct, the
parallelism with Precis sesamus, ete., is very remarkable,

The Bionomics of South African Insects. 437

In the Acrvinx we find that the least unpalatable species
of an unpalatable and conspicuous family have been com-
pelled to produce relatively inconspicuous generations in
the severe struggle of the dry season: in the Nymphaline
we find that some of the less palatable species of a com-
paratively palatable and inconspicuous family have been
compelled to produce strongly conspicuous generations in
the wet season when more edible insect food is abundant.

The interpretation I have here suggested was put
forward very cautiously in a note, dated Nov. 1898, to a
short paper on Mr. Marshall’s results with P. sesamus in
the Proc. Ent. Soc. Lond., Oct. 5, 1898, pp. xxv, XXvV1.
The note points out that insects with warning colours
are not to be seen in an English winter. “Those such as
Coccinellidx, which exist in the perfect form, hide them-
selves. The reason probably is that the amount of palat-
able food available is not sufficient to make it safe to rely
on unpalatability, accompanied by warning colouring [see
also ‘Colours of Animals, London, 1890, pp. 179, 180].
Experiments with hungry animals support this view. It
is possible that the conditions are similar in South Africa
[it is perhaps unnecessary to state that organic conditions
were alone referred to], and that warning colours are more
characteristic of the wet than of the dry season, thus
affording greater opportunities for mimetic resemblance.
If it should hereafter be shown that Precis is to some
extent unpalatable, and that its resemblance to an Acrean
type is synaposematic rather than pseudaposematic, the
parallelism with our own fauna would be even closer, the
conspicuous species which hide and thus adopt procryptic
habits being represented by one which gives rise to
another brood with markedly procryptic colouring and
habits.”

Mr. Marshall in commenting on this note records the
following interesting observations on the habits of South
African Coleoptera as determined by damp and dryness.

“Salisbury, Feb, 12, 1899.—Do you think that the
English Coccinellidx really hide in winter owing to their
increased danger from enemies, and not from climatic
causes? I ask the question because in this country
Coleoptera are highly susceptible to weather. They
appear to be for the most part absolutely dependent on
moisture, this being especially noticeable among the ter-
restrial forms such as Cicindelidx, Carabide, Psammodes,

438 Mr. G. A. K. Marshall on

Anomalipus, etc. These insects appear with a rush as
soon as the early rains have saturated the ground, but
should a dry spell supervene, they disappear as rapidly as
they came, only to emerge again on the recurrence of a
good rain. The case of the dung-beetles has always
puzzled me, for here we have a large family of powerful
and apparently hardy beetles, which have a constant
supply of food all the year round, and yet they are unable
to stand out the winter in the imago state, although a
delicate butterfly can do so. In fact, the ‘Copridx are
quite as dependent on moisture as the large Carabidx, and
are only to be seen at work from November to March,
retiring even then during the dry spells.”

The interesting effects of dryness described above cer-
tainly cannot be produced in our damp winters, and it 1s
difficult to believe that our cold can be the cause of the
retirement of Coccinellidx, ete., when species of insects
closely allied to those of England can endure to be frozen
stiff and brittle in a temperature of 50 degrees below zero
(F.) in a Manitoban winter.

Another letter. from Mr. Marshall, received about the
same time, contains a different comment upon the inter-
pretation suggested by the present writer in 1898 and
here amplified.

“Salisbury, Jan. 8, 1899.—I can fully perceive that
any arguments that may be brought forward in support
of the contention that Precis sesamus (natalensis form)
is an example of inciptent mimicry are equally applicable
to the suggestion of incipient warning coloration, and
for the present it must remain a matter of opinion as to
which is the correct explanation, though the alertness of
the insect and its undoubted palatability, so far as lizards
are concerned, seems to lend more support to the former
view to my mind.” [See also Ann. and Mag. Nat. Hist.,
ser. 7, vol. 1, July 1898, p. 35.]

It has been here shown that there are important elements
in the under-side coloration of the wet phases of Precis
sesamus and P, antilope which cannot be explained as
mimicry, Batesian or Miillerian (see pp. 425-8), while the
entire appearance of the under surface of P. archesia form
pelasgis can only be interpreted as a warning character
(pp. 428-431).

The conspicuous appearance of the under-sides of these
forms is doubtless chiefly adapted to render them con-

The Bionomees of South African Insects. 439

spicuous during the attitude of rest. There is probably
a certain parallelism with eryptic under-sides, such as those
of our Vanesside, which have no particular meaning im
flight and when the insect alights, but still remains fully
on the alert. The resting attitude is specialized in relation
to the development of cryptic colours and patterns on the
under-side, and in this attitude cryptic insects are always in-
conspicuous. Apart from the evidence of adaptation in the
direction of conspicuousness on the under-side of the wet
phases of Precis—the strongest argument for the presence
of some distasteful quality--the mere existence of such
an appearance in a palatable species is inconsistent with
the explanation of cryptic under-sides as the product of
adaptation in the direction of concealment from enemies.

The successful attacks of a species of lizard may be
analogous to other well-known instances in which special
enemies, such as the cuckoo, are known to devour con-
spicuous unpalatable insects.

Two other arguments in Mr. Marshall’s paper (Ann.
and Mag. Nat. Hist., ser. 7,-vol. u, July 1898, p. 30)
must be met here. First, the suggestion that the brilliant
colours of natalensis are due to the impunity with which
such a development can arise in the limited struggle for
existence in the stations occupied by the species, and the
abundance in the wet season of other insect food (loc. cit.,
pp. 35, 36). Such a suggestion does not explain the
under-side coloration, and especially the evidences of
adaptation init. Secondly, Mr. Marshall meets de Nicé-
ville’s and Weismann’s contention, that both seasonal forms
“must be adaptive, otherwise the non-adaptive form would
be gradually supplanted by its more favoured relative,”
by the suggestion that the dry-season phase may be a recent
development which is even now actually supplanting the
wet phase (loc. cit., pp. 36, 37). It is, however, difficult
to believe, looking at the Nymphaline as a whole, and
especially the nearest allies of the species under discussion,
that the conspicuous under-side of the wet phase is an-
cestral, and the cryptic under-side of the dry phase recent
(see p. 430), so that the argument set forth above seems
to me untouched. Mr. Marshall has indeed shown that
the dry phase of P. artaxia has actually supplanted the
wet phase (nachtigalii) in forest regions, where the
struggle for existence is far more uniferm at all seasons
of the year than it is in the more open woodland country

4.40) Mr. G. A. K. Marshall on

in which the dry and wet phases alternate. The displace-
ment of one forrm of artaxia by another is however no
evidence of relative age, but only of relatively better
adaptation to the conditions which obtain in the area
where the displacement has occurred. Hence the ob-
servation recorded by Mr. Marshall seems to me strongly
to confirm de Nicéville’s and Weismann’s conclusion that
when both seasonal phases exist, both are adaptive.

Mr. Marshall also shows on pp. 421 to 423 that the
species of Precis entirely restricted to forest regions possess
cryptic under-sides and habits all the year round, although
the dry-season generations are more completely cryptic.

It is not difficult to understand the observations referred
to above,—viz. that the appearance and habits while
cryptic all the year round should be more cryptic in the
generations of greatest stress. Thus Mr. Marshall describes
the wet-season phase of the purple-tipped South African
Teracoli as having under-sides not specially well adapted
for concealment on the ground during the resting attitude,
and without the habit of suddenly settling when pursued,
modes of concealment adopted by the dry-season generations.
In such examples the success of the adaptations may be
equal in the two seasons because of the difference in the
intensity of the struggle. But the extreme seasonal phases
of Precis can never be thus understood, because the wet
forms are not merely less cryptic than the dry, they have
gone over into the opposite camp, and have developed a
very extreme, and, except in the examples of mimicry and
warning colours, an unknown degree of conspicuousness.

Another and very interesting form of seasonal dimor-
phism is that which has been well known for a long time
in the Satyrinw, and consists chiefly in the development
of conspicuous ocelli, especially upon the under-side of
the wet phases and their greater or less suppression in
the dry. No interpretation of the change has, so far as
I am aware, been attempted, except that of Portschinski,
which has been further alluded to and criticized on p. 898.
I think it is probable that a valid interpretation is sug-
gested by the result of an experiment made in 1887, and
witnessed by Professor Meldola as well as by me. A
specimen of Canonympha pamphilus was introduced into a
lizard’s cage. ‘‘ It was at once obvious that the lizard was
greatly interested in the large eye-like mark on the under-
side of the fore-wing: it examined the mark intently, and

The Bionomics of South African Insects. 441

several times attempted to seize the butterfly at this spot.
The observation scems to point to, at any rate, one use
of the eye-like markings which are common on the under-
sides of the wings of butterflies,” viz. in order to attract the
attention of an enemy, and thus divert it from more vital
parts (“Colours of Animals,” London, 1890, pp. 206, 207).
The same interpretation is suggested by the habits of
many species which expose an eye-spot as soon as they
settle, when they are likely to be seized by an enemy
which has marked them down to their resting-place, but
quickly lower the wings and conceal the spot, so that they
are far more likely to be concealed from an enemy which
has not been specially directed to the exact place by
seeing them alight. (Much confirmation will be found
on pp. 371-5, where Mr. Marshall’s injured specimens are
described.)

Such directive marks may well be an advantage in the
wet season, when enemies with an abundance of other
insect food are less keen in their pursuit of butterflies,
but in the far greater stress of the dry season we can
understand how they would become a danger, and how
the only chance of the survival of the species lies in the
adoption of a cryptic appearance, and cryptic instincts in
their most extreme and unqualified form.

This explanation has much in common with that sug-
gested for the seasonal phases of Precis. Indeed, it is of
much interest to observe that nachtigalii, the wet form
of P. artaxia, has precisely the same relationship to the
dry form as that described above in Satyring. It is far
less cryptic than the leaf-like dry phase, but it is not
conspicuous. The ocelli on the under-sides of both wings
and the strongly-marked hind margins, together with the
specially prominent apex of the fore-wing, are probably
directive characters which divert the attention of an
enemy from the vital structures, when the insect is at rest
with its wings closed.

The relationship of the interpretation in Precis to that
just suggested in Satyrinw, and to that offered in certain
smaller Acrzas (see pp. 433-7), renders it on the whole
improbable that there is any alternation in degrees of un-
palatability corresponding to the alternation in the seasons.
There is, however, no @ priori difficulty in the hypothesis
that a higher degree of unpalatability may be correlated
with the conspicuous colouring of the wet phase of Precis ;

442 Mr. G. A. K. Marshall on

and experiments specially undertaken in order to test the
suggestion would be of much interest. That the hypothesis
is improbable is further shown by a long series of experi-
ments (hitherto only published in abstract in the Report
of the British Association, Manchester, 1887, p. 768)
which I conducted in 1887 with lizards and the highly
insectivorous marmoset. Large numbers of the imagines
of Vanessa io and V. urtice were made use of, and I came
to the decided conclusion that both were somewhat un-
palatable. They were certainly only eaten when the
insect enemies under observation were hungry. Now the
strongly cryptic under-side of both species associated with
a fairly-conspicuous upper-side renders them in every
way comparable to the dry phases of Precis. The results
of my experiments suggest that if Vanessa wrtice appeared
on the wing in the teeming organic environment of Africa
in the wet season—with far more enemies but an even
greater preponderance of palatable insects—it would be to
its advantage with its present degree of unpalatability to
acquire a conspicuous under-side coloration, and thus to
ensure easy recognition and rejection with comparatively
little loss of life by experimental trials.

The considerations set forth above suggest what will
probably hereafter be proved to be true, that a degree of
unpalatability associated with a conspicuous appearance
in the tropics will often appear associated with a cryptic
appearance in the Holarctic Belt as well as in those areas
of the tropics in which for special reasons the amount
and variety of insect life is greatly restricted.

It is suggested on pages 475 to 477, that this is the
interpretation of the loss of much of the aposematic ap-
pearance of Limnas chrysippus, var. klugit, on desert areas
in the tropics.

To return to the seasonally dimorphic Ethiopian species
of the genus Precis, if the two phases have been produced,
as is here contended, by natural selection working in
opposite directions because opposite kinds of adaptation
are advantageous in the very different organic environ-
ments of the wet and dry seasons, the questions as to the
way in which the change is actually determined, and as
to the existence of any kind of susceptibility to external
influences connected with the seasons, are still unanswered.
The considerable amount of labour devoted by Mr. Marshall
to the solution of this problem has up to the present

The Bionomies of South African Insects. 443

yielded negative results. Before describing his experiments
and discussing the results, it is desirable to show the
mode of succession of the phases in the wild state. By
far the most complete evidence I have been able to obtain
relates to a single species, P. sesamus.

G. The suwecession of the two Seasonal Phases of Precis
sesamus 77 Nature.

The followimg extracts from Mr. Marshall’s letters from
1897-1900 give an account of his experience of the
succession of wet and dry forms of this species in the wild
state, and also show how the conviction was gradually
forced upon him that the early appearance of occasional
specimens of the dry phase in the heart of the wet
season is not due, as he thought at first, to exceptional
climatic conditions (see also his paper in Ann. and Mag.
Nat. Hist., ser. 7, vol. viii, Nov. 1901, p. 402).

“ Malvern, Natal ; March 12,1897.—You will notice that
the dry forms of several species made their appearance at
the Karkloof in the middle of February: this is most
unusually early. I do not know whether it is a feature
and characteristic of that locality or whether it is due to
the abnormally dry weather during that month, which is
usually one of the wettest in the year. The average rain-
fall for this February was considerably lower than it has
been for twenty years. It is true that here the insects
are still all of the true summer form, but the proximity of
the sea may account for that.”

“Salisbury, March 6, 1898.—Sesamus was unusually early
here this year, appearing at the beginning of February,
full six weeks before its usual time. This I am inclined
to attribute to the exceptionally dry January and February
we have had—normally our wettest months—though I
am aware that Weismann considers that exceptional
seasons have little or no effect on seasonal forms, which
certainly does not accord with my experience in §.
Africa.”

“Salisbury, Feb. 12, 1899.—I send a specimen of P.
sesamus © captured on Jan. 27, 1899, on which day I also
saw another. These two examples are of considerable
interest as bearing on the problem concerning the stimulus

5
which actually induces seasonal change in this species.

4.44 Mr. G. A. K. Marshall on

In a normal wet season (in which there are more or less
continuous heavy rains from the middle of December to
the end of February) P. sesamus © appears at the end of
March. Last season we had heavy rains up to the end of
December 1897, but January was unusually dry and
sesamus @ appeared on Feb. 6, being the earliest record I
had for it. This season the drought was still more severe
in January and commenced earlier, viz. about Dec. 12.
This has been accompanied by a still earlier appearance of
sesamus @. The evidence so far as it goes tends to show
that climatic conditions, 11 some cases at least, are directly
capable of inducing the change and upsetting the normal
alternation of the forms. Here, owing to highly abnormal
conditions, we have the dry form occurring at what is
normally the very height of the wet season. Moreover,
ever since I have observed seasonal dimorphism in butter-
flies I have noticed the effect of abnormal weather in
retarding or accelerating the appearance of either form,
and Barker has made similar observations at Malvern, near
Durban, Natal.”

“Salisbury, April 25, 1899.—The sesamus form is
evidently more dominant than the natalensis, for despite
the heavy rains in February 1899 the latter made very
little headway after the appearance of the dry-season
form; whereas among such insects as the Pierine the
result of the alternating extremes was much more
evident.”

“ Salisbury, Feb. 7, 1900.—In spite of our heavy rains
during January (16°75 inches for the month) the winter
forms of Precis are appearing just as early as last year,
which has puzzled me considerably. I shall send you the
first examples of each form captured.”

“ Salisbury, June 26, 1900.—I am afraid I am not yet
convinced as to the automatic alternations [viz. due to the
organism itself and not to external stimuli] of the seasonal
forms in P. sesamus ; there seems to be at present an equal
amount of eaidenec on either side, and until the matter
can be settled by an exhaustive series of experiments if
must retain an open mind on the question.”

I have given below a list of all the specimens of the two
phases of “Precis sesamus sent to me from Mashonaland by
Mr. Marshall.

The Bionomics of South African Insects.

4.45

LOCALITY.

OBSERVER.

Mazoe,
4000 ft.

G. A. K. Marshall.

Gadzima,
4200 ft.,
Umfuli River.

Salisbury, |
5000 ft.

j G. A. K. Marshall. .

DATE,

FORM OF Precis sesamus
IN HOPE COLLECTION.

Dec. 28, 1894.

1 natalensis, fresh.

G. A. K. Marshall.

|

i

Dec. 20, 1895.
Dec. 23, 1895.

March 20, 1895.

1 natalensis, fresh.
3 natalensis, fresh.

1 sesamus, fresh.

Feb. 13, 1898.

2 natalensis (in coitu),
3 worn, ¢ fresh.

Feb. 20, 1898.

Feb. 27, 1898.

1 natalensis, worn (par-

1 natalensis, fresh.

ent of 1 sesamus,
and 1 natalensis).

March 2, 1898.

1 sesamus, fresh.

March 6, 1898.

1 natalensis, worn (par-
ent of 1 sesamus).

March 12, 1898.

March 16, 1898.

April 2, 1898.

Jan. 27, 1899.

1 sesamus, fresh.
1 sesumus, fresh.

1 sesamus, fresh.

1 sesamus, fresh (the
first seen in 1899).

| March 11, 1899.

1 natalensis, worn.

Feb. 3, 1900.

Jan. 23, 1901.

Jan. 26, 1901.

| 1 natalensis, fresh.

1 sesamus, fresh (the
second seen in 1900).

1 sesamus, fresh (the
first seen in 1901).

Mareh 2, 1901.

April 8, 1901,

1 sesamus, fresh, trans-
itional towards natal-
ens Ss.

2 sesamus, fresh.

A study of the above list makes it probable that the
occurrence of occasional specimens of the dry phase of
sesamus in January and February is a normal overlap.

4.4.6 Mr. G. A. K. Marshall on

Very careful and numerous records over a large number
of years would be required to show that any change in the
relative time limits of the two forms is taking place.

Owing to the kindness of Mr. 8. L. Hinde, H. M. Sub-
Commissioner, Kast African Protectorate, and Mrs. Hinde,
I have received a most interesting series of the two forms
from British East Africa, probably near the northern
boundary of the range of the species. The numbers,
captured in a short time on a limited area, are sufficient
to enable us to judge of the relative proportions of the
two forms, and we see that in May and the beginning of
June the two occur mixed in about equal proportions,
while in December and January the wet phase greatly
predominates, although an occasional dry form appears, as
it does in Mashonaland, early in January. I have included
in the series two other specimens from near the northern
part of the range of P. sesamus, The whole list is wonder-
fully similar to that from Mashonaland, and supports the
view that Mr. Marshall’s observations record the normal
mode of replacement of the wet by the dry phase, although
the former persists in large numbers much later in the
north than it does in the south.

447

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448 Mr. G. A. K. Marshall on

The succession of the seasons is very different, in the
region in which Mr. and Mrs. Hinde captured the butter-
flies, from that which obtains in Salisbury. The two
forms of succession are shown in parallel columns below.

BRITISH EAST AFRICA MASHONALAND
(MACHAKOS, KITUT, ETC.). (SALISBURY, MAZOE, GADZIMA).

Mid-October > Small wet season , Karly November \ Wet season, aver-

to \ (about 17—18 age rainfall of
Mid-December J inches). to | Salisbury about
__ to \gmall dry season. F ; 35 inches.
Mid-March J. | Mid- April
Big wet season
5 to \ (about 17—18 |
end May J inches). to Dry season.

to 5 :
Mid-October J Big dry season. | Early November

It is to be observed that the rainfall of the small and
the big wet seasons are about the same, and also that the
country is not really dried up in the small dry season
except in unusually dry years. The country is always
dried up in the big dry season.

In spite of these great differences in the seasons, the
succession of the phases is wonderfully alike in the two
areas, as has been pointed out above. We must conclude
that sesamus can produce two seasonal phases annually
but not more, so that the small dry season of the north
is no more effective in producing the dry phase than the
simultaneous wet season of the south. The species is so
constituted that it produces a dry phase for the big dry
season and a wet phase for the rest of the year, some of
the dry-phase individuals being produced some months
before the normal change takes place, viz. at and just
before the beginning of the chief dry season. The differ-
ence between the date at which this great change of
seasons takes place in north and south is attended by a
corresponding difference in the date at which the wet
phase of sesamus gives place to the dry. Both lists are
unfortunately wanting at the transition from the big dry
season to the wet. There is indeed only a single record
for the period between the beginning of June and the
beginning of December. Speaking from memory, Mr.
Marshall thinks that the break from sesamus to natalensis

The Bionomics of South African Insects. AAD

at the beginning of the wet season is not sharp; indeed,
he has a distinct recollection of seeing them flying together
at that season fairly frequently. “T believe,” he writes
(1902), “that in some seasons one might take sesamus in
every month of the year. Certainly at Gadzima, in 1895,
the true winter broods of sesamus lasted right up to the end
of December. In a dry spring, that is when the rains are
late in starting, buttertly life appears to be less abundant
and the emergence of the wet-season forms seems to be
retarded. On such occasions an actual break without
specimens might occur in such a comparatively unfavour-
able locality as Salisbury. But I believe that this would
be an unusual occurrence, and even if it happened in one
locality I doubt if it would necessarily take place every-
where at the same time; for example, in the moister parts
of the low veldt the successton of the broods would
probably continue unbroken. I am quite satisfied that
there are at least two or three broods of seswmus during
the winter months, that is if the condition of wild specimens
can be taken as any criterion. Food is much less plentiful
in the winter, but it is obtainable in quite sufficient
quantities to keep the species going. The change of
seasons from wet to dry is of rather a “oradual character ;
the reverse change is more marked, but this depends a
good deal upon the total rainfall of the preceding year.
When this has been heavy, the ground retains a certaim
amount of moisture throughout the w inter, so that when
the frosts cease and the sun’s heat increases in the spring,
a large number of the earlier plants spring up and flower
before a drop of rain has fallen. But after a succession of
dry years this does not take place, and, with possibly a
few exceptions, none of the plants come out im response to
the heat, but require the rains to bring them out. In
this latter case the change in conditions is very strongly
marked, much more so than during a wet cycle.”

The discussion of the possible nature of the environ-
mental stimulus, if any, is better deferred until after the
description and consideration of Mr, Marshall’s experiments
in the next section.

TRANS. ENT. SOC. LOND. 1902.—PART Ill. (NOV.) 30

450 Mr. G. A. K. Marshall on

H. The attempt to control the Phases of P. sesamus and
P. archesia by the artificial application of Moisture
and Heat to the earlier stages. Suggested lines of

_ Hxeperiment.

All the experiments hitherto made by Mr. Marshall were
directed towards the production of the wet natalensis and
pelasgis phases in place of the dry sesamus and archesia
respectively. The whole of the specimens produced were
presented by Mr. Marshall to the Hope Collection, and all
are tabulated below, together with a statement of the
experimental conditions which were employed in each
case. All experiments were made at Salisbury.

The following extracts from Mr. Marshall’s letters refer
to some of the experiments on sesamws :—

“ Salisbury, June 5, 1898.—I kept two larvee in a damp
jar, but one did not attach itself properly when pupating,
and the resulting pupa fell down when soft and was
killed. The other larva produced a black pupa which
emerged as the wet form [April 13, 1898, in the Table
below], but this was rendered nugatory by the fact that
one of the larvae in normal conditions produced the same
form, though from a gilded pupa [April 20, 1898, in the
Table below].”

“Salisbury, Apri 25, 1899.—I have fourteen bred
specimens of P. sesamus which I will send later [specimens
in year 1899 in the Table below]; I tried some experi-
ments with them, but the results are mostly negative.
There are two interesting varieties, one with a red bar in
the discoidal cell and another with the red spots much
reduced.”

451

Insects.

UCan

tes of South Afr

The Bionom

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4.52 Mr. G. A. K. Marshall on

The experiments on the power of adaptation of the
pupal colours to their environment are very interesting,
and prove that the susceptibility resembles that of the
allied British species Vanessa wrtice. They also show
that there is no essential difference between the colours of
the pupe of the two phases, but merely an adaptive
response to environments which differ in colour at the
two seasons, as suggested by Mr. Marshall (Ann. and Mag.
Nat. Hist., July 1898, p. 33). It is clear, from the experi-
ments, that if withered leaves of the usual winter yellow
appeared, owing to exceptional circumstances, in the
summer, larvae suspended to them would produce gilded
pupe instead of the usual dark summer forms, and,
mutatis mutandis, larvee would produce dark pupze upon
dark leaves in the winter.

The highest form of procryptic defence, viz. the power
of each individual to respond adaptively to any of its
different normal environments, here exists in the helpless
pupal stage, although the under-side of the wet phase of
the imago can only be interpreted on the supposition
that natural selection has developed a conspicuous appear-
ance. Our own Vanessidx however offer examples of the
same kind of association in the different stages of a single
life history. Thus the pupe of V. urtice and V. 20 have
the same specialized power of concealment, while their
gregarious black larvee are excessively conspicuous and the
imagines themselves by no means palatable to certain
enemies of insects (see p. 442).

No special significance appears to attach to the varieties
of the imagines produced in these experiments. The red
bar in the cell of No. 5 is a common variety which indeed
appears to be universal in the dry phase of the West
African P. octavia, and red scales can be detected in this
region in a large proportion of the individuals of P. sesamus.
The red spots of No. 5 are not specially developed. The
latter were largest in an individual exposed to normal
conditions (No. 16), while Nos. 14 and 17, also exposed
to normal conditions, were among the specimens with the
smallest spots. The bright blue shade of the ground-colour
of No. 9, exposed to damp heat, is well known in captured
specimens from the most northern part of the range of the
species as well as the south.

The specimens were weighed on an Oertling’s balance,
each pinned on a small cork foot which weighed -06275
grammes on June 29, 1902, and ‘00025 grammes more on

The Bionomies of South African Insects. 453

June 30. The weight of the No. 16 pin (D. F. Taylor’s)
was obtained by weighing three sets of ten similar pins.
The weight of the first ten was *7960 grammes, of the
second and third ‘7950 grammes. The average weight of
a pin was therefore 07953 grammes, and this number
added to ‘06275 was deducted from each of the specimens
weighed on June 29 (see p. 456): added to °063 it was
deducted from the e specimens of the experiments recorded
above, and all others weighed on June 30 (see pp. 451,
456). On the latter date the cork foot was weighed
at the beginning of work, in the middle, and at the end.
On all three occasions it weighed ‘063 grammes.

The consideration of the experiments on Sesamus 1s
better deferred until after describing those upon archesia,
although it is at once evident that no positive conclusions
can be drawn as to the nature of the environmental
stimulus. The negative character of the results obtained
induced Mr. Marshall finally to form the opinion quoted
below.

“Salisbury, Feb. 26, 1902.—I do quite agree with you
now that in the case of Precis the evidence is sufficiently
strong to show that climate has ceased to operate as the
stimulus which calls forth the seasonal change. But I do
not think that this view is applicable to “other genera
whose changes coincide closely with the changes in
climate. The theoretical proposition I would suggest is
that at its inception seasonal change was but slight and
then due entirely to climatic action, such cases doubtless
occurring at the present time. Any markedly useful
variations of this kind would then be preserved and
accentuated by natural selection, but climatic causes would
still remain the controlling factor. Finally, as in Precis,
the influence of natural selection would attain its maximum,
and the seasonal changes would then take place solely as
a result of this principle and irrespective of the influence
of climate. It remains to be seen whether this can be
proved by experiment.”

On pp. 455 to 458 it will be seen that there are still
hopes that the operation of some environmental stimulus
may yet be discovered in the case of Precis.

The results obtained from the smaller series of experi-
ments upon 2. archesia are even more negative than those
yielded by P. sesamus, as will be seen by a glance at the
Table below, giving a complete account of all that has
been as yet done.

Mr. G. A. K. Marshall on

454

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The Bionomies of South African Insects. 4.55

Concerning the specimens which emerged on May 7 and
14, and April 28, 1899, Mr. Marshall wrote as follows :—

“ Salishury, Aug. 29, 1899.—The case of pelasgis and
archesia was very puzzling, as the results were just the
opposite of what one would expect—the forced pupa
emerging as the dry-form archesia, and the normal ones
as the wet-form pelasgis, though this latter has disappeared
for some time, being replaced by archesia.”

The negative results from these fairly-numerous experi-
ments tempt us to believe that the change from sesamus
to natalensis and natalensis to sesamus may be fixed in the
constitution of the species, and may form an alternating
series contemporaneous with the alternating seasons but
not causally connected with taem. Such a view is how-
ever rendered improbable, as Dr. Dixey has pointed out to
me, because there would be nothing to prevent a gradual
shifting and finally an entire want of parallelism between
the two series. That, however, the change is essentially
constitutional in the species and merely requires some
external stimulus to set it going may be taken as certain.
Furthermore, it is not necessary to suppose that a stimulus
is required for both changes, the return to one of them,
and presumably the more ancestral, may be in the nature
ofa rebound. The slight but distinct difference between
the succession of the forms of sesamus in British East
Africa and in Mashonaland also probably indicates a
causal relation with the inorganic environment, and the
same conclusion is supported by the fact that arfaxia has
been observed without its wet-season phase in a forest
region (see pp. 422-3).

After Mr. Marshall’s experiments it is difficult to believe
that the application of heat or moisture or the two com-
bined to the pupal stage can determine the production of
natalensis or pelasgis in place of sesamus or archesia,
respectively, at the period when the latter forms are
becoming abundant in nature. It is possible that here
we are merely witnessing the return to a more ancestral
phase due to purely internal causes. The reverse experi-
ment, viz. the application of cold, or dryness, or both com-
bined, to pupe of the earlier generations of natalensis, might
produce more positive results and cause the appearance of
sesamus at a time of the year when it is very rarely seen,
although the occurrence of occasional individuals of
sesamus in nature in the depth of the wet season seems to

456 Mr. G. A. K. Marshall on

be quite unrelated to dryness or cold (see pp. 445-8). But
it would probably be necessary to apply artificial conditions
to the larval stage. Indeed, the fact that the winter
phases of certain species of Precis are so very much larger
than the summer phases seems to require the conclusion
that the change is pre-determined during or previously to
the stage in which material is accumulated.

The differences in weight are well shown in captured
individuals of two species in the following list: the method
of procedure has been already described on pp. 452-3.
It is seen that the dry phase always weighs more and
sometimes over twice as much as the wet one.

WEIGHT.
DATE OF CAPTURE. | PHASE. | FIRST 2 ON JUNE 29,
{REST ON JUNE 30, 1902.

SPECIES. SEX. | LOCALITY.

|

Precis antilope.| & | Salisbury. | March 2, 1898. | Wet. | °03422 grammes.

Feb. 27, 1898. | Wet. | -03747 grammes.

Precis antilope. | Salisbury.

April 3, 1898. | Dry. | 07472 grammes.

| Precis antilope.|

2
Precis antilope.| @ | Salisbury.
2

| :
Salisbury. | March 9, 1898. | Dry. | 04947 grammes.

| Precis artaxia. | Unmtali. Dec. 27, 1900. | Wet. | 05597 grammes.

Precis artaxia.

Umtali. | Dee. 30, 1900. Dry. | (09672 grammes. |

2
Precis artavia.| g | Umtali. Dec. 30, 1900. | Dry. | 08447 grammes.

Precis artaxia.| 9 | Gadzima, | Dec. 29, 1895. | Wet. | 06622 grammes,
| Umfuli R. |

| Precis artaxia,| g | Gadzima, | Dec. 30, 1895. | Wet. | °05422 grammes.

| Umfuli R. |

| Precis artavia.|~@ | Gadzima, | July 29, 1895. | Dry. | 06997 grammes.
| | Umfuli R.

Precis artaxia. | Mazoe. Dec. 28, 1894. | Wet. | ‘04522 grammes.
| poe Tal ee ee : = || abe as SS
Precis artaxia | @ Mazoe. | Oct. 23, 1894. | Dry. | ‘09547 grammes.

(worn). |

It may be argued that the results from captured speci-
mens are untrustworthy because some females will have
laid their eggs, some males will have paired, and others

not. The five series of specimens of sesamus and natalensis
bred by Mr. Marshall are not open to this objection and

The Bionomics of South African Insects. 457

are therefore of especial value. The weights of the
eighteen bred specimens are given on p. 451, and it will
be seen that the difference between the phases is very
marked, although not nearly equal to that between the
two forms of artaxia.

There is no escape from the conclusion that the larvee of
the dry phase of these species must be much larger than
those of the wet, and must eat a great deal more food.
This inevitable conclusion suggests that in experimenting
on this most interesting of all known examples of seasonal
change, it will be well to keep an open mind on all con-
ceivable stimuli: on the abundance and character of the
food-plant as well as the inorganic conditions of humidity
and temperature, the latter of which has been proved by
Dorfmeister, Weismann, Merrifield and Standfuss to be
an effective stimulus in the case of certain Palearctic
seasonally dimorphic species. It is possible that the
parched state of the food-plant towards the end of the dry
season may be the stimulus which determines development
in the direction of the smaller summer phase. The different
sizes and weights render it nearly certain, as I have argued
above, that the phase is predetermined in the larval stage.
Now the larval stage of the first dry-season brood is passed
in the wet season, and that of the first wet-season brood
probably in the dry. We must look to some condition
affecting one or both of these larval stages, or the eggs
from which they arose, as the stimulus which sets in
motion the organic processes resulting in a change of
phase. Some colour of support is lent to the suggestion
that the condition of the food-plant may afford the neces-
sary stimulus by the fact that the wet phase of P. artaxia
is unknown in certain forest regions, where it is probable
that the food is not subject to the same alternation of
condition as in more exposed stations. But forests would
also act as moderating influences for extreme differences
in temperature and humidity, and thus tend to prevent
these from acting as stimuli for the species in question ;
for we know that some stimuli must be effective in pro-
ducing such seasonal changes as occur in other forest
species of Precis (see p. 423). Finally, quantity as con-
trasted with condition of food would be well worth trying.
The unusually low weight of the imagines bred from the
egg (Expts. 11, 12, and 13 on p. 451) was a probable

65
result of difficulty in obtainig a constant supply of fresh

458 Mr. G. A. K. Marshall on

food in an entirely normal state, and it is noteworthy that
one out of the three was natalensis. The extremely low
weight of the ? natalensis in Expt. 10 also suggests some
unfavourable condition in the larval state.

With the facts before us I do not see that any further
suggestions can be made at the present moment; but I
think the tables of weights clearly indicate the period at
which the stimuli should be applied, while temperature,
humidity, quantity or quality of food, or some combination
of these, seem to exhaust all probable influences in the
direction of a change of phase.

T. The Bearing of the Seasonal Phases of Precis upon the
Science of Insect Systematics.

The results which have been described and illustrated
in this section of the present memoir are so startling that
they may well shake the confidence of naturalists in the
whole fabric of insect systematics. If such forms as
natalensis and sesamus, as simia and antilope, as pelasgis
and archesia, are nothing but the generations of two
alternating phases of a single species, approximately
synchronized with the heat and cold or humidity and
dryness of the alternating seasons, naturalists may feel
driven to ask, “ What becomes of the validity of specific
distinctions?” Between the two phases of Precis sesamius
there are extraordinary differences in colours, pattern,
shape of wings, relation of upper- to under-side, nay, even
in instinctive habits, including the choice of particular
stations. This latter distinction between the phases is
but the outward expression of some profound difference in
the intimate structure of ganglionic centres and inter-
communicating strands in the central nervous system.
Important differences in venation are incidentally brought
about by the great differences in the shape of the wing.
The extreme rarity of intermediate varieties furthermore
recalls the abrupt transitions which are so common,
although very far from universal, between species of
animals which are assumed to be distinct. Under the
shock of Mr. Marshall’s discovery that sesamus and
natalensis are two forms of the same species, the systematist
may well feel doubts about the foundations upon which
his science has been erected. In these distracting circum-

5
stances a firm belief in natural selection will be found to

The Bionomics of South African Insects, 459

exercise a wonderfully calming and steadying influence.
The structures which are adopted as the conventional
criteria of specific distinction are of course modified by
natural selection and brought into adjustment with new
conditions of the struggle for existence as one species is
gradually changed into another; but they are also capable
of modification in one and the same species as it passes
through various conditions during its life-history and in
sexual and other dimorphism. ‘The species frequently
requires that the female sex should be more protected than
the male, and hence we often witness a more perfectly
cryptic appearance and habits in the female, and mimicry
in the female alone. In many kinds of di-, tri- and
polymorphism we see a species more perfectly protected
at one and the same time by extending the area over
which it must be sought by its enemies—in cryptic
resemblance, earth and bark as well as leaves and shoots—
in mimetic resemblance, Danaine or other distasteful
models not of one species alone but two or more. In the
di-, tri- or polymorphism of the social Hymenoptera and
Neuroptera we see the specialization of the individual for
the good of the community. In the extreme cases of
seasonal dimorphism, exhibited by the genus Precis, there
is a far less common modification of a species into two
series of generations respectively adjusted to the con-
ditions obtaining at two seasons of the year. But less
marked cases of the same kind are probably not uncommon.
There is however nothing revolutionary or subversive in
any of these interesting facts. ‘The conventional marks of
specific distinction remain just as they were, convenient
indications to the systematist, enabling him provisionally to
separate groups of individuals into the assemblages we call
species. When his work is done carefully subsequent
breeding experiments will, we may be sure, confirm his
conclusions in the majority of cases. But here and there
startling exceptions will be found when it is to the
advantage of a species to appear in two or more very
different forms. In such cases the reason for the differ-
ence can generally be satisfactorily explained on the
principles of natural selection ; and when such an explan-
ation is possible or even probable it is quite unnecessary
to assume that the exceptions possess a numerical import-
ance sufficient to shake the foundations of systematics.
Certain species are cryptic while others are aposematic

460 Mr. G. A. K. Marshall on

or pseudaposematic; certain stages in the life of an indi-
vidual may be cryptic, others aposematic or pseudapo-
sematic. ‘There is nothing subversive in the thought that
certain species exposed to different organic environments
in two seasons of the year may appear as cryptic genera-
tions at one of these, aposematic or pseudaposematic at
the other. The explanation is at any rate sufficiently
probable to enable us to contemplate Mr. Marshall’s wonder-
ful discovery with equanimity and with an interest un-
disturbed by the thought that he has laid in ruins the
whole edifice of insect systematics,

29. THE GREGARIOUS INSTINCT IN HYBERNATION AND
EMIGRATION OF Insects. (E. B. P.)

The interesting observation that idividuals of Precis
sesamus are apt occasionally to congregate in large numbers
as they go to roost, led me to reflect on the possible
meaning of such an instinct, Mr. Marshall records other
examples of the same kind “in species of Huralia, also
in Belenois, Herpenia eriphia, and Teracolus eris” (Ann.
and Mag. Nat. Hist., loc. cit., 1898, p. 34). It is possible
that one interpretation does not explain all these cases,
but I think it is probable that the observed instances of
the congregating of Precis and Huralia are sporadic ex-
amples of an instinct which is associated with hybernation
or, at any rate, a prolonged period of rest during a time of
relatively excessive cold, heat, or dryness. Objection may
be taken to this interpretation on the ground that large
companies undergoing a prolonged rest ought to be well
known in these species. It is possible however that the
extreme conditions which render such a state desirable
or even necessary for the species are not common, and,
when they occur, do not conduce towards the active pursuit
of natural history; furthermore, such prolonged rest would
probably be passed through in some hidden recess which
could only be found by accident.

Large numbers of naturalists for hundreds of years have
been interested in the doings of Vanessa io, but, so far as
IT am aware, it is not generally known that. this species
may display a gregarious habit in hybernation.* My friend,

* Edward Newman recorded the occurrence of a company of more
than forty V. io in a hollow oak (British Butterflies and Moths,
London, N.D., p. 16), and the Rey. Joseph Greene disturbed three

The Bionomics of South African Insects. 461

Dr. W. Hatchett Jackson, the Radcliffe Librarian at
Oxford, permits me to publish the following observation
made by him at Weston-super-Mare in the second week
of January 1895. Dr. Jackson found, in the garden of
his house, on the side of a hill sloping south, about twenty
peacock butterflies hybernating in the heart of a bramble-
bush. The butterflies were arranged in rows on two or
three approximately horizontal runners about a foot from
the ground. All rested with their wings hanging down-
wards. When the butterflies were first disturbed they
made no movement, but on repeated disturbance they

specimens in the hollow formed by the arching roots of a large beech-
tree, in Dee. 1852 (G. C. Barrett, British Lepidoptera, London,
1893, vol. i, p. 139). W. 8, Coleman (British Butterflies, London,
1862, p. 88) quotes Doubleday in the Zoologist :—* Last winter some
large stacks of beech faggots, which had been loosely stacked up in our
forest (Epping) the preceding spring, with the dead leaves adhering
to them, were taken down and carted away, and among these were
many scores of io, wrtice, and polychloros.” No reference is given,
and I have failed to find the original statement. An observation of
Mr. Banning of Monte Video, Ballacraine, Isle of Man (also quoted
by Coleman, |. c. p. 91), is recorded in the Zoologist (1856, p. 5000) :—
“Whilst standing in my farmyard on the day following Christmas
Day [1855], it being unusually fine and warm, I was suddenly
astonished by the fall of more than a hundred of the accompanying
butterflies [Vanessa urtice]. I commenced at once collecting them,
and sueceeded in securing more than sixty... .” This observation
apparently points to the emergence of a hybernating assemblage in
consequence of exceptionally warm weather. It also indicates con-
ditions which at a normal time of the year would be favourable to
pairing.

Mr. J. W. Tutt states that the imagines of V. io feed largely during
August, disappearing at the end of the month or in September
(Entomologist’s Record, 1895-6, vol. vii, p. 3). It is therefore
probable that the butterflies produced by one company of larvee do
not keep together, or the fact would certainly have been noticed
when they are in search of food. It is probable that the products of
all companies scatter and become thoroughly intermingled before
again assembling into groups for hybernation. Another lne of
evidence may perhaps yield incontrovertible proof of the existence
of this intermixture before reassembling—a probable adaptation to
prevent in-and-in breeding. Dr. W. H. Jackson and Mr. O. H.
Latter, F.E.S., have found that the pupze obtained from different
batches of larvee of V. io “were principally, but not entirely, of one
or of the other sex” (Trans. Linn. Soc., London, vol. v, 1890, p. 156).
It would not be difficult to obtain a numerical statement of the
average constitution of a company in this respect, so that it would
be available for comparison with that of a hybernating group. A
marked difference would prove intermixture before hybernation,
while a similar constitution would yield negative evidence.

462 Mr. G. A. K. Marshall on

flicked their wings and the movement passed along the
row. It must be remembered that the climate of Weston
is extremely mild, and the great frost of 1895 had not
then begun.

The advantages of a period of rest during excessive
heat and dryness may be as great as those which follow
from excessive cold. In the former the food-plant may
be parched and dry or confined to very few and widely-
scattered damp spots, and the perfect insect may pass
through its life without the chance of laying eggs in
places where the larvae would be able to survive. But
quite apart from this, the continuous excessive drought
may be injurious to the perfect imsect itself. At the
driest and hottest part of the African dry season a great
scarcity even of common butterflies has been noticed, and
it is not unlikely that many individuals of some species
pass through the most critical period of very dry and hot
years concealed in a state of rest. It is significant that
the congregating instinct has only been observed in the
dry phase of Precis sesamus.

This does not solve the problem of the gregarious instinct
itself. It is clear that Dr. Jackson’s observation on V. 70
and Mr. Marshall’s on Precis, etc., indicate the existence of
an instinct which must be a real and great danger to the

5
species. The less the individuals congregated and the

oO

more widely they scattered, the greater would be their
chance of safety. A fortunate enemy finding one of the
peacocks in the bramble-bush at Weston would have
secured the whole. It is therefore certain on the principles
of natural selection that some great advantage is gained
by the instinct, an advantage which more than compensates
for the increased danger. I would suggest that this
advantage is the facility ¢ riven for pairing -and the laying
of eggs Without any loss of time, as soon as the period of
rest comes to an end. The advantage would be quite as
great or even greater after the rest during drought than
after ordimary hybernation, because of the rapidity with
which the food-plant recovers with the first moisture. It
would be interesting to consider from this point of view
the food-plants of the African species in which the instinct
has been observed.

This suggestion naturally leads to a consideration of the
gregarious instinct in the peculiar form of emigration
which has been observed in insects. The same increased

The Brionomiecs of South African Insects. 463

dangers attend the phenomenon, and I think it is very
probable that they are more than compensated by the
analogous benefits. The instinct to emigrate probably
exists in a dormant state in all species lable from their
powers of rapid multiplication suddenly to outrun the
food-supply in any part of their range. The stimulus
which evokes the instinct is, in such insects as the locust,
or such mammals as the lemming, probably merely the
direct and obvious incentive of hunger (A. R. Wallace,
“ Geographical Distribution,’ London, 1876, vol. i, p. 18).
In the majority of perfect insects, however, we cannot
accept this interpretation, and we are compelled to look
for a stimulus in some other result of undue increase—the
crowds of individuals everywhere, the food-plant covered
with eggs and young larvee, and females laying still more
eggs. Then probably arises the imperative instinct to
move, perhaps in both sexes, perhaps only in the female,
the males accompanying them (in many species in far
larger numbers). And the instinct further compels the
individuals to move together in vast masses in the
same direction, rather than to scatter and fly in all
directions. The increased danger from enemies is of
course lessened, as compared with the hybernating com-
panies, by the enormous number of emigrating individuals;
but there is, I believe, the solid advantage that fresh food-
plant may be found in another uncrowded area; that the
limits of the normal range of the species may be overpassed ;
that areas from which the species has been driven may
be regained :—not by single individuals or by a very few
pairs, but by immense numbers of both sexes without any
of the dangers of in-and-in breeding when once they have
established themselves as a fresh colony. In this way
the range of many species has probably been extended
in the past, and, although the emigrating crowds so often
described may again and again be landed in a foodless
desert or the sea, the instinct is advantageous in that it
utilizes individuals which are at the moment useless and
even injurious to their kind, in a manner which may be
in a high degree beneficial (see also Trimen, “South
African Butterflies,” vol i, 1887, p. 31). The suggestion
is made that the crowded masses, resulting from over-
production and inability of enemies to cope with the
Increase, are injurious to the species, because it is likely
that food-plants would be checked for years or even killed

464 Mr. G. A. K. Marshall on

out altogether in certain localities, while the heaps of dead
individuals would encourage the attack and rapid spread
of bacterial foes. Indeed, the advantages to be derived
from the removal of the surplus from an overcrowded
area may probably overweigh those which accrue from
the occasional successes in colonization, and may more
than the latter account for the development by natural
selection of the instinct to move. The massing of the
moving individuals and their flight in the same direction
seem, on the other hand, to have arisen by selection from
the beneficial results conferred by spreading into less
crowded areas. It is difficult to imagine any other possible
means by which such animals as insects could overcome
the effects of asudden increase too great for the restraining
influences of their natural enemies—effects which in-
sufticiently checked for a few generations would inevitably
lead to the destruction of the species in the area of over-
production.

We may well inquire why it should be necessary for
such emigration, with a possible successful issue in colon-
ization, to require the services of countless individuals
when the importation of half-a-dozen rabbits or a few
specimens of Pieris rape will, for the naturalist, change
the face of a continent. The results of these unintentional,
or intentional but ill-considered, experiments do indeed
shake the behef in the paramount necessity for crosses
and the dangers of in-and-in breeding ; but the end is not
yet, and the teeming colonies which have arisen from such
small beginnings may in time vanish from the operation
of deep-seated causes. The varied adaptations for cross-
fertilization and the prevention of in-and-in breeding are
so evident in nature, that we are compelled to believe that
they meet and counteract serious dangers which sooner
or later would menace the very existence of the species.
And among other adaptations it is significant that the
instinct under discussion should lead to the streaming
of large populations, and not of small batches of individuals
from an area of high pressure.

The gregarious instinct in emigration has been observed
in many groups of insects beside the Lepidoptera. I need
only mention here the hundreds of Amimophila hirsuta,
ordinarily a solitary species, found by Fabre under a large
flat stone on the summit of Mont Ventoux at a height
of 6000 ft., and the crowds of ladybirds witnessed by him

The Bionomies of South African Lnsects. 465

on the same mountain and on the tableland of St. Armand
(“Insect Life,” English translation, London, 1901, p. 193).
A valuable account of a large number of observations will
be found in Mr. J. W. Tutt’s numerous papers on “The
Migration and Dispersal of Insects” (Ent. Record, 1898—
1902). The author recognizes the dangers of over-
multiplication as a cause of migration (1. c. vol. xii, 1900,
p. 238: see also vol. xin, 1901, p. 200). Numerous
examples quoted by him prove that movement in vast
bands, often at great distances from land, has been
observed again and again in those very species which
are remarkable for their wide geographical distribution
and occurrence upon oceanic islands. The appearance
of two species of Hylbernia, H. defoliaria and H. auwranti-
aria, observed in large numbers in Heligoland by Gitke,
cannot be explained on the hypothesis here suggested
because, the females being flightless, males only appeared.
In several other instances recorded by Mr. Tutt the
presence of both sexes is either specially affirmed or
implied. In the great majority of cases, however, no
observations of sex were made, and it is to be hoped
that careful attention may be paid to this point in the
future. The proportion of females to males would also
be deserving of careful investigation.

The limits of this memoir are perhaps too wide already,
and it 1s impossible to attempt any discussion of these
observations in detail, but I have taken the opportunity of
making a suggestion as to the possible essential meaning
of the instinct.

I have deliberately used the word “ emigration” of
insects, because this term probably expresses the exact
state of the case. In response to some stimulus connected
with undue increase, immense masses of individuals move
out of an overcrowded area. The line of movement may
carry them to destruction or to plenty, in bath cases
benefits are probably gained, although they are of course
much greater in the latter. True “ migration” as of birds
and perhaps of fishes implies different and far higher
faculties—the memory of the individual summed up by
tradition into what may be called the collective memory
of the species.

TRANS, ENT, SOC. LOND. 1902,—PART II (NOvV.) 31

466 Mr. G. A. K. Marshall on

30. DESCRIPTION AND DIScUSSION OF MATERIAL BEARING
on Mimicry IN SourH AFRICAN RHOPALOCERA COL-
LECTED BY Guy A. K. MARSHALL, AND THE RECORD
OF OBSERVATIONS MADE BY HIM. (HE. B. P.)

The splendid material which is described and discussed
below has gradually accumulated as the result of Mr.
Marshall’s kind and generous response to my desire for
specimens for the Hope Department illustrating the fact
that mimetic species and their models, and the members
of large convergent or synaposematic groups, not only in-
habit the same areas but fly together at the same time.
The study of this material naturally led to conclusions and
suggestions which it is hoped possess a general interest in
relation to the doctrine of evolution and the important
part which mimicry plays in it, as one of the chief evidences
of the operation of natural selection. ‘These more general
discussions are placed under separate headings immediately
after the groups whose study gave rise to them.

The last sub-section is placed under Mr. Marshall’s name,
being quoted in extenso from his letters.

A. Black-and-White Amauris-like Group.

The central model for the group described below is
probably Amauwris ochlea, but 1t was not captured on
March 27, 1897, when five convergent individuals were
taken at Malvern, near Durban, Natal. The group as
captured is as follows :—

Planema aganice &.

3 esebria 2, var. with white markings.
Neptis agatha €.
2 Nyctemera lewcono€.

The male Planema aganice is but an imperfect member
of the group, the lighter markings being buff instead of
white, as in the female. We thus find that the latter sex
forms closer synaposematic resemblances than the male,
when the two sexes differ. It is probable that this rela-
tionship between male and female will be found to be
generally true of Miillerian mimics in which the sexes
exhibit different degrees of likeness to the type of some
group characterized by Common Warning Colours. Fur-
thermore, the culmination is often reached in Miillerian

The Bionomics of South African Insects. 467

mimicry, just as it is in Batesian where it has long been
recognized, in species of which the female enters “into a
more or less well-marked membership of a group towards
which the male has made no apparent approximation.
Numerous examples will be found in the present memoir.

This interesting similarity between Miillerian and
Batesian mimicry was probably unrecognized until 1894,
when it was discovered by F. A. Dixey,* because of the
fact that in the first-known examples of Miillerian mimicry
in tropical America, which are the most wonderful instances
in the world, the convergent pairs and groups contributed
by the Heliconinx and Ithomiine and by different genera
within each of these sub-families, are made up of species
with males and females which are superficially alike.

Now, however, tliat the principle has been recognized by
Dixey in many Neotropical Miillerian mimics with differ-
ing sexes and here in many Ethiopian, the explanation is
doubtless the same as that suggested by Wallace (Trans.
Linn. Soc. xxv, Pt. I, 1865) in the case of Batesian mimics,
viz. the great importance for the species that the female,
with her slower flight and the necessity to pause and lay
her eggs, should gain to the full the advantages of that
extra advertisement of warning coloration which is con-
ferred by membership in a synaposematic group. This is

=) .
the interpretation offered by Dixey in his 1894 memoir

(q. v.).

Neptis agatha exhibits in an interesting manner that
concentration of white markings into four large patches,
one upon each wing (save that the fore-wing is invaded
by a small portion of the hind-wing patch), and that
disappearance of the other bars and markings, except
for traces on the under-side, which are characteristic of
many Ethiopian species of this genus, and doubtless indi-
cate a synaposematic approach to the black-and-white
species of Amauris and Planema of the Region.

It is too wide a subject to introduce into the present
memoir, but I cannot forbear to allude to the evident
synaposematic sensitiveness of the genus JVeptis, leading it
to form associations with local conspicuous Rhopalocera.
Among the most beautiful of these are WV. venilia and NV.
lactaria, which resemble the remarkable Danaine genus
Hamadryas, especially upon the under-side: Again, the

* Trans. Ent. Soc. Lond., 1894, p. 298, note ; 1896, pp. 70, 71;
1897, pp. 319, 326-328, 3380,

468 Mr. G. A. K. Marshall on

likeness to Athyma and Limenitis must have struck every
naturalist who has looked through the drawers of a toler-
ably large collection. Col. Swinhoe has recently called my
attention to a Huplwa-like Neptis from China, NV. imitans.

The resemblance of the genus Neptidopsis to Neptis
seems, on the other hand, to have been due to mimetic
approach on the part of the former towards the type set by
the latter.

The Hypsid moth Vyctemera lewconoé seems to have in-
dependently adopted the same aposematic scheme of
colourmg as the genus Amauris, the only change in the
direction of the latter dominant type being a slight broad-
ening of the white bar crossing the fore-wings, a broaden-
ing which is at once recognizable when this and other
African species of the moth are compared with their nearest
Oriental allies. The conspicuous and almost certainly
specially-protected Hypsidw strongly tend to enter into
synaposematic association with other specially-defended
forms in various parts of the world. Thus one species
approximates towards Hamadryas, while, in tropical
America, the smaller forms become transparent and
resemble the smaller Jthomiine, while the larger (Pericopis)
possess the warning coloration of species of Melinwa and
LHeliconius.

B. Limnas chrysippus-like Groups.

The first of these groups was captured on March 6, 1897,
at Malvern, Natal. It consists of the eight following
individuals :—

Limnas chrysippus § .
as z 2, var. alcippoides. _

Hypolimnas misippus %, with pale hind-wings like
the last-named insect.

H. misippus 2, var. maria.

Planema esebria §, chrysippus-like type-form with
white sub-apical bar to fore-wings and reddish-
brown black-bordered hind-wings, the ground-
colour extending on to the fore-wings.

Acreva encedon &.

A, serena, var. buxtoni, ¢.

A, doubledayt §.

The latter individual, bemg a male, is not really a
member of the group, inasmuch as it lacks the oblique

The Bionomics of South African Insects, 469

sub-apical white bar present in the female of typical
doubledayt. The presence of the male indicates, however,
that the female flies with the other members of this
chrysippus-like group, of which it forms an imperfect and
outlying constituent. The male of serena also does not
resemble chrysippus, while the female is an even more
imperfect Miillerian mimic than the female of doubledayi.
Nevertheless such cases are of the highest. interest, inas-
much as they enable us to understand how mimicry arose
in species which now exhibit a startling likeness. A.
encedon, one of the most perfect Miillerian mimics of
chrysippus, presents an equally close approximation in
male and female.

The fact that the female of P. esebria should present
two well-marked varieties, one of which falls into a black-
and-white group convergent round the species Amauris,
while the other, the type-form, enters the combination
which surrounds ZL. chrysippus, recalls a principle already
well known and probably correctly understood in the case
of Batesian mimicry. When an abundant well-protected
Acrva thus approximates to two very different Danaine
patterns it is obvious that we are not necessarily driven to
a Batesian interpretation of the forms of the female Papilio
eenea, Which approximate to the appearance of Amawrts
echeria as well as to the two other Danaine types alluded
to above. The enemies of chrysippus and the species of
Amauris are certainly not precisely the same, and it may
well be an advantage to a Miillerian mimic to secure that
increased protection from insect-eating enemies which is
conferred by belonging to two or more groups.

Furthermore, the Planema has come to resemble the
Danaine and not the Danaines the Planema, and this
probably indicates that the Danaine is on the whole the
less attacked and the better known. It is probably of
advantage to the whole group that the Danaine which set
the pattern should still be the dominant member of the
assemblage of which it is the centre. This dominance is
favoured by the individuals of an abundant species joining
two or more groups instead of throwing the whole of their
number into a single one. In the case of Batesian mimicry,
where the mimics are comparatively palatable and would
be freely eaten if recognized, the advantage of this di- or
trimorphism and the likeness to two or three models is
even more obvious.

470 Mr. G. A. K. Marshall on

A second group of the same type was captured in the
same locality on March 30, 1897, and consists of six
individuals :—

LL. chrysippus §.

Hf, misippus 2, a pale patch in the centre of each hind-
wing.

P. esebria J, buff sub-apical bar to fore-wing.

A. encedon &.

A. serena, var. buxtoni &.

A. petrva 9.

The lack of correspondence between the varieties of the
females of HZ. mzsippus and those of the central member of
the group is well seen in these two sets. Thus one of the
three females is the inaria form, although the klugit var.
of chrysippus is almost unknown in 8. Africa, while the
other two suggest the appearance of the alcippoides var.,
which does indeed occur not uncommonly, but is not
nearly so abundant as typical chrysippus. The female of
A. petrexa is another outlying member of the group, while
the male is altogether outside it.

A third group, captured by Mr. Marshall at Salisbury on
April 10, 1898, contains these species :—

L. chrysippus ° |
H. misippus § ¢ all typical forms.
2 A. encedon f |

A fourth group captured at the same locality on April
9, 1899, contains :—

2 L. chrysippus ¢, 2 (Plate XIV, figs. 1, 1a).
2 Mimacrea marshalli § (Plate XIV, figs. 2, 2a).

This beautiful Lyczenid mimic presents in some respects
a closer approximation to Acrwa encedon (Plate XIV, figs.
3, 38a) than to LZ. chrysippus, the primary model of both.
Thus the character and contour of the sub-apical white
bar of the fore-wing suggests that of the Acraea rather
than the Danaine. In the two former the bar is more
continuous, in the latter more obviously broken into dis-
continuous spots, attended by outlying smaller spots.
Another far more important similarity between Lyczenid
and Acrza is brought about by the numerous conspicuous
black spots which in both are scattered over the basal part
of the under-side of the hind-wing. In other respects the

The Bionomics of South African Insects. 471

under-side of the Lyczenid presents a much closer approx-
imation to the Danaine than does the Acraa, In well-
marked individuals there are fourteen of these spots in the
Lyceenid, nineteen in the Acrea. In place of these, the
Danaine model possesses, in addition to the marginal
white-marked spots and a single spot at the extreme base
of the wing, only a row of three spots on the outer
boundary of the cell in the female, with an additional
white-centred black patch, marking the scent-pouch, in the
male. The three spots along the outer margin of the
cell are encircled with white, as are all the spots in the
Lyceenid ; while the veins of the under-side of the hind-
wing in both are more or less emphasized with white. In
these minor but distinct points the Lyczenid approximates
to the Danaine and not to the Acrzea; and furthermore in
the fact that the black spots of the under-side of the hind-
wing are hardly visible on the upper-side of the Lyceenid,
while all except the basal ones are as distinct upon the
upper- as upon the under-side of the Acraea. In the Danaine
they are very distinct although much smaller on the upper-
side, but as there are only three in the female and four in
the male, the Acreea is in this respect much further from
the other two than they are from each other, its distance
being still further increased by a few (4—6) large conspicu-
ous black spots on the ground-colour of the basal half of
both surfaces of the upper wing. It is noteworthy that
these points of divergence on the part of A. encedon are
characters which it shares with a large number of related
species. All the points mentioned above can be well seen
in the six upper figures of Plate XIV, which should be
compared with the six corresponding figures on Plate XV,
showing the prevalent form of the Danaine model and
its Acreine and Lyczenid mimics much further north in
British East Africa.

In the dark shade of the brown ground-colour the
Acrza is much nearer to the Danaine as developed in 8.
Africa than the Lycenid, and upon the wing the black
spots would probably make the Acraa appear still darker.
The bright fulvous tint of MZ, marshalli is more of the
shade of the Oriental specimens of chrysippus. This rela-
tionship appears to exist between many of the other African
Miillerian and Batesian mimics of L. chrysippus and their
model, and suggests that the Oriental bright shade is
ancestral, although the Oriental intrusion is comparatively

472 Mr. G. A. K. Marshall on

modern, as proved by the relatively small amount of
mimicry, and that little very imperfect, in species peculiar
to the Region. The fact that the bright Oriental shade
still persists in many of the specimens of chrysippus from
the north-east and probably other parts of Africa, supports
the same conclusion.

Mr. Roland Trimen points out that the Mimacrea also
resembles the female of the type-form of Planema esebria
(Trans. Ent. Soc. Lond., 1898, p. 15).

Mr. Marshall gives the following account (1902) of the
habits of this interesting insect :—‘ In its general habits
Mimacrzea marshalli, Trim., like Acrea encedon, is essen-
tially a woodland (but not a forest) insect, and shows a
marked fear of venturing out into open country. Limnas
chrysippus, on the other hand, frequents both open and
woodland stations. When quite undisturbed it flits about
ina limited area of the bush with a slow flight exactly
resembling that of L. chrysippus (see also pp. 481, 482),
but when alarmed it is capable of flying with considerable
speed, and dodges with great dexterity. When hard pressed
it will occasionally rise right over the tops of the trees,
descending into the bush again further on. But its usual
method of escape is by dodging in and out among the tree-
trunks, then settling suddenly on the far side of one of them,
which makes it extremely difficult to follow. It is in its rest-
ing habits that it differs most from the mimetic group to
which it belongs, for I have never seen a specimen settle
anywhere except on a tree-trunk, and then always with its
head downwards, just like a Libythea or the brown species
of Crenis. The Danaine and Acreine, on the other hand,
always rest with their wings hanging down, and usually
in more exposed positions; indeed, I think it may be
said that no species of these groups (in South Africa at
least) ever settles upon tree-trunks. This habit is also a
very unusual one among our Lycsenide, one or two species
of Teriomima being the only cases of its occurrence which
I can recall. Despite the great resemblance of this
Mimacrva on its under-side to Acrwa encedon, yet its
colouring when at rest is very far from conspicuous, and
harmonizes a great deal better with its surroundings than
might be supposed from an examination of the insect in
the cabinet, especially as the fore-wings are so much
depressed between the hind-wings as to quite conceal
the sub-apical white patch. The species seems to be of

The Bionomics of South African Insects, 473

considerable rarity; I know of only fourteen or fifteen
specimens, all of which, except two or three, were captured
by myself.”

C. The Origin and Meaning of the Three Chief Fornis
of Limnas chrysippus.

I have often discussed the question set forth in the title
of this sub-section with my friend Colonel J. W. Yerbury,
who has observed this insect carefully in many of its local-
ities, and is deeply interested in it.* He believes that the
appearance of the various forms is controlled by environ-
mental influences—dryness or moisture—acting upon the
pupa at some critical period of special sensitiveness. The
facts recorded below do not seem to be consistent with
this interpretation.

My friends Mr. and Mrs. 8. L. Hinde, who have kindly
collected many specimens throwing light on problems to
which I have given much thought (see also pp. 446, 447),
sent me two series of forms of Limnas chrysippus, which
are of special value in relation to this discussion.

The first set (of 15) was captured, almost on the sea-
level, in the uniform damp heat of Mombasa, on May 6,
1900, and consists of four of the type-form of Limnas
chrysippus (1 § and 3 @), and eleven of the form klugii (9
d and 2 9). All were taken in less than an hour on a
spot of ground a few yards in extent. They thus afford a
fair criterion of the proportionate numbers of the two
forms.

The second set (of 13) was taken, at a height of about
5400 ft., at Machakos Road, on the Uganda Railway, on
May 22, 1900, and consists of four of the type-form (3
and 1 ), one alcyppoides ( f), seven klugii (2 f and 5 9),
and one dorippus (f). These also were taken on a spot
of ground a few yards in extent, in less than an hour.

Mr. Hinde has given me information as to the climate
of the period in which the latter capture was made. It
is printed on pp. 447, 448, but it is well to re-state here
that May 22, 1900, was at “the end of a very dry wet-
season in an exceptionally dry year.’ The specimens show
the effect of these conditions, for they are on the average

* J. W. Yerbury, Journ. Bomb. Soc. Nat. Hist., 1892, p. 207.
Col. Yerbury’s observations on the species are also quoted by Dr.
A. G. Butler in Proc. Zool. Soc. Lond., 1884, p. 478; 1885, p. 756.

474 Mr. G. A. K. Marshall on

very much smaller than those bred from larvee which fed
on the more luxuriant food-plant in Mombasa. Examining
the two series, we are driven to the conclusion that the
Machakos larvee were partially starved, probably by feeding
on parched food-plant. Interesting and important ferences
may be drawn from the comparison.

In the first place the specimens, so far from supporting
the conclusion often arrived at from incomplete and, as I
think, ill-regulated experiments, that males are produced
by starvation, actually show a larger number of females in
a smaller total of specimens than the set from Mombasa,
viz. 6 out of 13, as against 5 out of 15. Even if the
females had been very scarce at Machakos, nothing would
have been proved in the direction of the determination of
the sex of the individual by diet, for starvation pushed to
the extreme of preventing the completion of development
of many individuals is certain to kill off the heavier sex
far more freely than the lighter. The results, however,
show no abnormal excess of males, and in every way
support a prediction firmly founded on the anatomical fact
that the essential organs of sex, the testis and ovary, are
already present, rudimentary, but perfectly distinct, in the
larval stage.

A comparison of the two series furthermore indicates
very strongly that the various forms of the species are not
in any way due to environmental causes, but are inherent
and hereditary. It is believed that Alugii is due to
drought, but there is a larger proportion of this form in
the series bred in the moist heat of the coast than in
that reared at high and dry Machakos. The great differ-
ence in conditions which is manifest in the different
average size of the two series was powerless to effect any
change in the inherent hereditary tendency of the indi-
vidual to become either /wgii or its modification dorippus,
the type-form or its modification a/eippoides.

This comparison of forms from adjacent localities under
different climatic conditions leads to an inference which is
precisely the same as that drawn from the comparison of
forms from different localities under the same climatic con-
ditions. The tropical forests of West Africa and the
Malayan Islands are very similar as regards climate: in
the first chrysippus occurs as the white-hind-winged
alcippus, in the second as the type-form, a peculiar dark
form inhabiting Java (L, bataviana). It is not necessary

The Bionomics of South African Insects. 475

to pursue such comparisons further. So far as chrysippus
is concerned, I know of no facts which support the hypo-
thesis of the environmental production of the forms, and
many which are inconsistent with it.

The only alternative hypothesis which presents itself is
that of the operation of natural selection in determining
the very different distribution of the various forms of
chrysippus in the different parts of its range. And in
attempting to solve this difficult problem I have been
guided by the same principles which enabled me_ to
suggest a meaning for the two widely-different seasonal
phases of Precis, viz. the relation between insects and their
enemies, the value of warning colours under certain con-
ditions, their weakness and danger under other conditions.
I believe that the condition of desert areas corresponds to
that of the dry season, only differing in that they are more
rigid, so that cryptic colouring is still more imperative. I
therefore suggest that the klugit form is a development in
a procryptic “direction in areas where the struggle is so
severe that even this most unpalatable and widely-
mimicked species must put off some of its aposematic
appearance, viz. the conspicuous black-and-white apex of
the fore-wing.

There is also a peculiar faintly greenish-orange shade in
the area of the apex of the fore-wing under-side beyond the
sub-apical white bar of chrysippus which is wanting from
the corresponding part of klugii, the difference tending to
bring about a further uniformity in the ground- colour of
the under-side of the latter.

Furthermore, many specimens of //ugii have a ground-
colour quite different from that of even light individuals
of the type-form, gaining a distinct sand colour. This is
all the more striking in Africa, where the type-form
commonly develops a dark rich fulvous ground-colour
very different from the paler Oriental type.

This interpretation is based on the assumption that
klugu has developed from chrysippus and not chrysippus
from klugii. and no escape from this assumption seems
possible. The main lines of argument are these. Island
individuals, which are so generally ancestral, are chrysippus
and only very rarely klugii, except near the metropolis of
the latter form in Somaliland. Perfect and imperfect
mimics, Batesian and Miillerian, are very-large in number,
especially in Africa. Probably not one of them mimics

476 Mr. G. A. K. Marshall on

klugit and not chrysippus, a few mimic both, while the
great majority mimic chrysippus alone. Again, klugii
stands alone among Danaina, while the pattern of chrysip-
pus is closely related to that of several other species and
genera, such as Salatwra. The distribution of klwgii can
be understood by supposing the desert region of Somali-
land to be its centre. From this point it radiates,
towards the east becoming gradually rarer, although it is
well known in Karachi, and Colonel Yerbury even captured
a single specimen in Ceylon, towards the south finally
disappearing in South Africa, towards the Nile Valley,
here also probably disappearing towards Central Africa,
Chrysippus, or at least forms with a black-and-white apex to
fore-wings, on the other hand, occur over the whole vast
range of the species with the exception of certain parts
of Somaliland.* The strongest argument is, however, one
which is developed at the end of the section, because wide
conclusions of great interest spring from it (see pp. 482-484).

The white-hind-winged alcippus was for a long time a
great difficulty to me, but Mr. Marshall’s suggestion (see
p. 479) that it is a further development in the direction of
still more efficient warning colours than the type-form
seems to me to be sound, especially considering its distri-
bution in the abundant life of the tropical West Coast,
and considering the fact, of which Mr. Marshall assures
me, that it is much more conspicuous on the wing.

I have for a long time thought that this great develop-

5
ment of white, combined with the darkening of the
fulvous ground-colour so common in African specimens
and marked in aleippus, may indicate an incipient synapo-
sematic approach to the black-and-white Danaina of the
genus Amauwris, and the large black-and-white Acrvine
of the genus Planema.

It is in favour of this view that the darkened ground-
colour appears to be a recent development, although it has
arisen in the Hthiopian region—the ancestral home of
the species, if we may judge by the much larger number
of mimics which resemble it in this part of its range. I
believe the lighter ground-colour of chrysippus in India
and, with certain exceptions, the Hast generally, formerly

* Consult Dr. A. G. Butler’s map of the distribution of the forms
of chrysippus, on Plate XLVI, accompanying his paper in Proc.
Zool. Soc. Lond., 1884, p. 478; also by the same author, Proc.
Zool, Soc. Lond., 1885, p. 756.

hey

The Bionomics of South African Insects. ATT

prevailed in Africa, because so many of its African mimics
retain this shade rather than that now borne by their
model, and because chrysippus itself often exhibits the
paler tint, especially to the north and east * of the African
continent (see pp. 471, 472.).

I quote below extracts from several of Mr. Marshall’s
letters dealing with the hypothesis which I have here set
forth, and also referring in other ways to this interesting
butterfly and its forms.

“ Malvern, May 14, 1897.—I was interested to hear
that the Z. chrysippus, var, alcippoides, | sent resembles
the West Coast specimens. Personally I have never come
across that variety commonly, and those I have taken have
mostly been very shghtly marked with white, but Mr. G. H.
Burn, who has collected for some years in the Tugela Valley,
near Weenen, says they are not uncommon there, but appar-
ently are most prevalent in the early winter (it is a very
hot dry district), and they are frequently marked quite as
strongly as the one I sent you.”

“Salisbury, Jan. 24, 1900.—Referring to your very inter-
esting remarks on L. chrysippus, 1 was much struck by your
theory with regard to//ugiz. The only point, however, on
which I do not feel satisfied is whether we can consider the
colouring of this variety to be really protective. Of course
I have never seen it in life, but I have seen many of the
anariva form of Hypolimnas, and I must confess that the
insect is very far from being inconspicuous; and even apart
from colour it must be remembered that slowness of flight
is a very important factor in rendering an insect conspicu-
ous (compare our large and powerfully-armed but soberly-
coloured wasps of the genus belenogaster). Again, it would
seem hardly consistent to regard the colouring of klugii as
protective if we rank that of the smaller Acreas, which
inhabit some at least of the same areas, as among the
warning colours. It would therefore strengthen the
theory if some other use could be assigned to the klugii
coloration, though none occurs to me at the moment. Of
course the correlation argument might be brought
forward; but while thoroughly appreciating Meldola’s
masterly defence of this principle, 1 must admit that I
have a distinct distrust in its use in such cases, as it
means virtually begging the question. - I cannot. alto-
gether gather from your remarks what are your reasons

* See F. A. Dixey in Proc. Zool. Soc., 1898, p. 373, note.

478 Mr. G. A. K. Marshall on

for supposing that the lighter Asiatic form is the older
[see pp. 471, 476]. On general principles it would seem
that swamping would be likely to keep the species more
or less constant in its ancestral home, whereas those speci-
mens that wandered further afield would probably tend
to vary along slightly different limes; but perhaps I have
not properly caught your idea. The case of alcippus
would be a great deal more difficult to explain satisfactor-
ily, seeing that it occurs also at Aden ; and Butler says
that examples sent from such places as Monbuttu, Wade-
lai, ete, by Emin Pasha, showed every gradation from
chrysippus through aleippoides to alcippus; further, if I
remember rightly, you wrote me that an example I sent
you from the Tugela had the white developed as strongly
as in any West Coast specimen, and Burn said they were
by no means uncommon there.”

[« A. G. Butler records (Proc. Zool. Soc., 1896, p. 243)
Captain Nurse’s statement that in Somaliland he bred
all four forms of chrysippus from quite similar larve.”
Goa M3025]

“ Salisbury, June 26, 1900.—Referring to the question
of Limnas klugii, although I fully appreciate the value of
your arguments, yet I must confess that when looking at
the matter from the point of view of an opponent of
mimicry, it seems at least open to criticism, The difficulty
seems to lie in the fact that the same coloration would
thus have to be regarded as both protective and warning.
Now you have said that in desert regions insects would be
more liable to attack owing to the paucity of insect life,
and I should be glad to know whether you have any
special reasons for adopting this view, as I have no ex-
perience of what the conditions of life really are in such
localities. But don’t you think that it is more likely that
the struggle for existence would be principally against
climatic conditions and not so much a competition with
other organisms, and that thus probably insects would
have a better proportionate chance of finding a living than
would the vertebrates as compared with more fertile
regions? If this were so it would follow that insects
would be comparatively freer from attack in desert regions,
and this would afford us another explanation of the klugia
phenomenon. We might presume that the less conspicuous
klugwt colouring was the more ancestral (as seems not
unlikely), but that in the more fertile regions where

The Bionomres of South African Insects. 479

insectivorous vertebrates and invertebrates are so much
more plentiful, this coloration was not sufficiently striking
to guard it from the tasting experiments of these enemies,
and thus the white bar would be gradually developed.
The typical form would therefore supplant //ugii in all
places where there was greater need of more efficient
warning colouring, and the latter form would only survive
in those tracts where only a limited number of insectivorous
enemies could exist. Such an explanation would further
throw some light on the additional development of white
in the hind-wing of alcippus in the prolific West Coast belt,
where the increased number of vertebrate enemies re-
quiring to learn by experience renders an additional con-
spicuousness advantageous. It seems to me that if such
an explanation could be maintained it would be more
consistent with our general views; but of course the
matter hinges on the conditions of life in desert tracts,
which is merely an assumption on my part.”

“ Salisbury, Sept. 21, 1900.—I was much interested in
your remarks on the subject of klugii, and I quite agree
that now that you have shown that my premise as to the
conditions of life in desert countries is erroneous my con-
tention falls to the ground. I should like to know how
the range of the aria form of mzsippus falls in with your
proposition, and also whether you find the variety of Acrwa
encedon without the white bar to follow the same range as
Klug. I have found this variety [daira] extremely rare
in 8. Africa, but a correspondent wrote me from Beira the
other day that it was not uncommon there and promised
to send me specimens.”

“ Estcourt, Natal ; Oct. 15, 1896.—I had thought Acrea
encedon might be mimetic, but it must be a case of con-
vergence—the type towards Limnas chrysippus and the
variety /ycia towards esebvia. That this latter is the case
I am led to believe by the fact that in Mashonaland only
the type-form occurs and there esebria is also absent,
whereas along the South-east Coast where the latter is
plentiful, /ycia occurs, and when I met with it in Durban
I was struck with its resemblance on the wing to the
whiter specimens of esebria, though this is not so apparent
in the cabinet.”

“ Malvern, Natal; Feb. 21, 1897.—The case of JZ.
misippus is however more puzzling than the Huralias
which mimic Amauris. The inaria form of the female is

480 Mr. G. A. K. Marshall on

often cited as a mimic of ZL. chrysippus, var. klugi. Now
this is by far the commoner form of the female misippus in
S. Africa, whereas klugii appears to be extremely rare, in
fact the single specimen recorded by Trimen is the only
one I know of. How then can it be said to be mimicked
by wnaria? Again, misippus is recorded in several places
in South America, where I believe chrysippus does not
occur. It seems to require further imvestigation.”

“Umkomaas Mouth, Natal ; Sept. 3, 1897.—I certainly
think that I have more frequently seen Hypolimnas
misippus (female) in company with ZL. chrysippus than
with its own male. The latter is fond of haunting the
tops of kopjes in company with various species of Precis
(which always occur in such localities), but I have never
seen the female do so, neither does chrysippus.”

The range of the forms of encedon corresponds remark-
ably well with the forms of chrysippus. Mr. Marshall
states above that the Alugii-like form daira is extremely
rare in the south where k/ugii is absent. Passing north-
ward on the east side of the continent it gradually im-
creases in proportionate numbers till it preponderates over
encedon where klugii preponderates over chrysippus. On
the West Coast all forms seem to occur, but recently the
white-hind-winged alcippina (Plate XV, fig. 7) has been
found there in greater numbers than elsewhere. (Auri-
villus, Rhopalocera Ethiopica, Stockholm, 1898, pp. 533,
534; Poulton, Proc. Linn, Soc. Lond., 113th Session, p. 6,
Report of Meeting Dec. 20, 1900, where however the name
encedon 1s erroneously printed unicolor .) The distribution
of the Lyceenid mimic corresponds equally well, marshall
with chrysippus in the south (Mashonaland), dohertyi with
the predominant //wgiiin British Kast Africa. H. misippus
2 shows upon the whole an almost complete lack of corre-
spondence, for «na7ia 18 common nearly everywhere, while
klugit is confined to the range described on p.476. In British
Kast Africa, however, misippus ? corresponds well with the
two forms of its model; while on the west, where alcippus
is the only form, the want of geographical coincidence is
most striking, for the inaria form is relatively abundant,
while neither in it nor in the type-form, so far as I am
aware, is there any special tendency towards the develop-
ment of white in the hind-wings. It is a striking fact
that the Acreine mimic should exhibit so close a co-
incidence with the geographical range of its Danaine

The Bionomics of South African Insects. 481]

model, while the Nymphaline mimic shows such a marked
want of correspondence. The comparison may help
naturalists to realize the great importance of Miillerian
mimicry and the searching selective process which has
brought it about.

I have for many years attributed this want of corre-
spondence between the commonest mimic of chrysippus
and its model, to the wide-ranging powers of the former
butterfly and its great tendency to wander, combined with
some special protection which there is reason to believe it
possesses, rendering its resemblance synaposematic rather
than pseudaposematic. There are in the Hope Depart-
ment three females (two of them inaria) and two males of
H, misippus captured out of a swarm through which the
ship Winefred passed in May 1893, when she was on the
Atlantic over 500 miles from land (Ent. Record., vol. xii,
No. 11, p. 315). The Miillerian resemblance of imisippus 2
to chrysippus was suggested by the present writer at the
meeting of the American Association for the Advancement
of Science in 1897 (see vol. xlvi, p. 242, where arguments
in support of this conclusion may be found). Extracts
on this subject from Mr. Marshall’s letters are printed
below :—

“ Malvern, Natal; Oct. 7, 1897.—I fear I cannot at
present accept your suggestion that Hypolimnas misippus
is itself protected. I may be wrong, but in these matters
I depend more than anything on the habits and actions of
the insects as I have seen them when undisturbed and
when frightened. There is to my mind a radical difference
between mimics and their models (as opposed to convergent
forms) which is often very difficult to define. There is
also a structural difference which appeals to me, so that I
believe I could almost tell one from the other with my
eyes shut merely by the feel of it in the net. To give an
instance: when on a short holiday trip to the rich Mazoe
Valley in December 1894, I started out on Christmas Day
with the set purpose of catching something “good” to
commemorate the occasion. While strolling along the
narrow belt of thick bush which there fringes the river, I
saw flying leisurely in front of me what I took to be a
very small and brightly-coloured specimen of Limnas
chrysippus. 1 coveted it, and a few seconds later it was
in my net, through the folds of which I could but indis-
tinctly see it, so that I was still deceived. But no sooner

TRANS. ENT. SOC. LOND. 1902.— PART II. (NOV.) 32

482 Mr. G. A. K. Marshall on

had my finger and thumb met across its thorax than my
heart beat high with that keen excitement that every
ardent entomologist feels when he has found some un-
expected treasure—for I knew I had got a new mimic of
chrysippus! A short inspection showed it to be a new
Lycenid—a Mimacreva (since named J/. marshalli, Trimen).
On the other hand, when I caught my first specimen of
Aletis here in Malvern in March 1898, I remember it
puzzled me much, for I felt sure it was not a mimic of
chrysippus, and yet I could not understand the reason of
the colouring, for I was then unaware of Miiller’s theory.
“To return to misippus, although I admit it is a some-
what difficult case, yet the fact that it has elected to mimic
what I take to be the hardiest and best-protected butterfly
in Africa, combined with the general adaptability to varying
conditions which one would expect it to share with its
nearest allies the Junonias, must I fancy go a long way
towards explaining its wide range and comparatively large
numbers. There is another factor that must not be over-
looked, and that is that through a considerable portion of
its range in South-east Africa at all events it 1s the only
butterfly which shows mimicry of chrysippus. This is
particularly noticeable on the rolling grass veldt of the
inland plateaux where chrysippus is particularly abundant.
“JT think your idea as to the latter insect having originated
in Africa is excellent and in every way most probable.”
“Salisbury, March 6-10, 1898.—The facts that you
mention with regard to Hypolimnas are certainly very
curious, and would seem to be only explicable by presuming
the species you mention to be protected. But in the case
of Hypolimnas misippus, after reviewing the general habits
and attitude of the female, I cannot bring myself to believe
that it is anything but a true Batesian mimic. Might it
not be a similar case to that of the genus Papilio, in which
we have the distinctly protected and distasteful P. coon
and at the same time the clearly mimetic P. cenea?”

D. A Study of Mimetic Forms may enable us to reconstruct
the Lost Stages through which the Older Model has

passed,

If klugii has been derived from chrysippus we should
expect to find traces of the markings of the latter upon
the wings of the former. And as a matter of fact faint

The Bionomics of South African Insects, 483

indications of the white sub-apical bar of chrysippus can
be detected in klugii, especially at the points on the costa
and the hind margin which the two ends of the bar would
have reached. Very faint traces of the course of the bar
between these two points can be made out in certain
individuals (Plate XV, fig. 1), while occasionally they are
very distinct, especially upon the under-side (Plate XV,
fig. la). Looking at these two figures, and comparing
them with Figs. 1 and 1l@ on Plate XIV, it is impossible
to resist the conclusion that we see before us the vestiges
of a fading character and not the rudiments of a developing
one. It is interesting to note that one of the shehtly
intermediate varieties of k/ugii here represented (viz. Fig. 1,
Plate XV) was an individual captured by Mr. and Mrs,
Hinde at Machakos Road, and that three or four others of
the same set showed similar tendencies. It may be that
the unfavourable conditions (see pp. 473, 474), although
unable to change one form into another, nevertheless
administered a shock which caused a slight reversion
towards the ancestral type in some individuals,

The three great mimics of both forms of chrysippus, the
female of the Nymphaline, Hypolimnas misippus with its
inaria form mimicking klugii; the Acrzeine, A. encedon *
with its Alugii-like form daira; the Lyczenid Mimacrea
marshalli with what I believe to be merely its k/wgii-like
form dohertyi, all these show precisely the same thing as
their model only im an exaggerated form, because the
mimie follows its model and therefore still exhibits stages
which the latter has left behind. Comparing the upper-
and under-side of the cirysippus-like Lycenid on Plate
XIV (Figs. 2 and 2a) with those of the klugii-like form
on Plate XV (Figs. 2 and 2a), there can be no doubt
that the latter developed from the former. The white
bar of marshalli (Plate XIV) can still be distinctly
traced in doherty. (Plate XV), not mdeed as a white
bar but as a very faint paling of the ground-colour
over a sub-apical area, the outline of which exactly

* The first recognition of the mimicry of chrysippus by encedon,
and indeed of the existence of Miillerian mimicry in the Ethiopian
Region, was first brought forward at the meeting of the British
Association at Toronto in 1897 (Report, p. 689). Aurivillius (Rhop.
Eth. 1898, p. 533) states that the resemblance had not been previously
noticed. The account given by Aurivillius is however far more com-

plete than that in the brief abstract here referred to, and is also
accompanied by illustrations,

4.84 Mr. G. A. K. Marshall on

corresponds to the bar itself. The comparison to a pseudo-
morph suggests itself; the bar is indeed absent but its
shape is there. The case of the Acraine mimic is still
clearer, Figs. 3 and 3a on Plate XIV bearing the same
relationship as that above described in marshalli-dohertyi,
to Figs. 3 and 3a on Plate XV. The same “pseudo-
morph” ” of the white bar can be seen in the latter, while
in Figs. 4, 5, and 6, on Plate XV, some by no means un-
common intermediate varieties between encedon and daira
are represented. Fig. 7 shows the form aleippina which
resembles alcippus, the white-hind-winged chrysippus.

When a geologist finds a recognizable fragment of one
rock included in a stratum of another, he is usually safe in
inferring that the latter is the younger. With equal
confidence the zoologist may conclude that the mimicking
species 1s younger than the species it mimics. The latter
must have been in existence before the former attained a
resemblance to it. From this point of view the comparison
between chrysippus-klugiv and their mimics is of intense
interest. Chrysippus and klugii are now well defined the
one from the other, and it is probably impossible or at
least extremely difficult to get a series of intermediate
forms between them. If we had not the mimics we might
well believe that kluwgii arose ready-made from chrysippus
by a process of discontinuous or transilient evolution.
But two of the younger mimics are very common and
widespread, and both misippws-inaria and encedon-daira
present us with abundant varieties showing every grade
of transition from the one form to the other. Of the
Lyczenid less can be said. It is still extremely rare (sce
pp. 472, 473) and at present only known in two widely-
separated areas. But even in it the gap marshalli-dohertyi
has been shown above to be much less wide than that of
chrysippus-klugii. Weare led to believe from this com-
parison that in some earlier age the two forms of the
Danaine model existed in the stage now reached by their
commonest mimics, and, like these, were connected by a
series of abundant intermediate varieties which have’ since
been obliterated by selection.

E. Amauris echeria-like Group: Marked Secondary Resem-
blances between the Forms mimicking echeria.

This species of Amawris, with its very characteristic
rectangular buff patch on the hind-wing and buff or white-

The Bionomies of South African Insects. ASD

spotted fore-wing, is the dominant Danaine of South
Africa, and extends in considerable abundance right up
the east and east central parts of the continent well into
British East Africa. It is a centre of convergence for
several Nymphaline and Papilionine in the same district.

An interesting group, captured by Mr. Marshall at
Malvern, on March 25, 1897, consists of the following
species :—

Amauris echeria, var. albimaculata 2.

2 Huralia mime §.

Papilio cenea 9, cenea-form, with white spots on fore-
wing, like the var. albimaculata of A. echeria.

Papilio leonidas, var. brasidas.

The last-named Papilio is a somewhat outlying member
of the group, being separated from the others by the
patch on the hind- wing, which is white with a faint
greenish tinge, instead of buff. The general arrangement
of the light markings on the black ground is however
similar, and Mr. Trimen states that “it was in the
habit of settling precisely in the way affected by the
Amauris, viz, on a projecting leaf or twig, with the wings
closed and hanging downward, and in this exposed position
remaining motionless for a considerable time” (“South
African Butterflies,” vol. 11, 1889, p. 216). In two points,
viz. size, and contour of the wings, it is more like the
Danaine model than any of the other above-mentioned
species.

The most interesting point about the group as a whole
is, however, the undoubted secondary resemblance between
the species which primarily resemble the Amauris. The
secondary resemblance is, moreover, even stronger between
P. cenea and the Huralia than between either of these and
P. brasidas, that is to say, the species exhibiting a closer
primary mimicry also exhibit a closer secondary mimicry.
This fact suggests that the secondary resemblance is of
permanent value and not a mere phase which will ulti-
mately be lost in the primary resemblance. Mr. Marshall
informs me that these secondary mimetic resemblances are
still more marked upon the wing, so that a naturalist may
often be sure that he sees before him a mimic of echeria
or of a black-and-white Amauris, but cannot im any way
distinguish the mimic itself as Papili io, Pseudacrewa, Hypo-
limnas, ete,

486 Mr. G. A. K. Marshall on

The points in which the mimics of A. echeria converge
together and diverge from their primary model are as
follows :—

(1) Size: They are much larger than their primary
model. Srasidas is in this respect intermediate.

(2) Scalloped outline of hind-wing; feebly marked
in brasidas. Slight imdications of scalloping are
intensified in echeria by the fringe bemg marked
by two white spots in each of the shallow concavities.
This is only distinct in some individuals, appar-
ently chiefly from the northern part of the range.
The marked concavities of the mimics are also
intensified by whiteness.

(3) Elongated oval shape of largest spot in fore-wing,
viz. the spot below the cell, while that of the
model is nearly circular. The long axis of the
oval spot furthermore assumes the same direction
in each of the three species, while that of echeria
is entirely different.

(4) The much greater prominence in all three mimics
of the sub-marginal row of white spots on the
upper-side of both wings.

(5) Strongly-marked black internervular rays pass
inwards from the margin of the hind-wing of
Huralia and invade the periphery of the ochreous
patch, uniting with the black ves to make up
a pattern of radiating dark lines. The radiate
appearance of the under-side is even more promin-
ent than that of the upper-side. Papilio cenea ¢
is very similar, except that the upper surface
exhibits only faint indications of the character
(which however is strongly marked in the hippo-
coon form resembling Amauwris dominicanus).*

Some of these differences between mimics as a whole

* In this respect, viz. the prominent development of internervular
rays on both surfaces of the hind-wing, the hippocoon form of the
Western P. merope presents a far closer resemblance to its co-mimic
Ewralia anthedon than to the primary model Amawris niavius, and
similarly the hippocoon form of the Southern and Eastern cenea to
Ewralia wahlbergi than to Amauris dominicanus. Not only is there
the conspicuous radiate appearance wanting in the Danaine model,
but the white centre of the upper surface of the wings deepens
eradually at its margin into black in both Papilio and Nymphalid,
while the margin of the corresponding white area in the Danaine
exhibits an extremely sharp and abrupt transition into black,

The Bionomics of South African Insects. 487

and their primary model are less pronounced in the
northern part of the range, in British East Africa, where
the Amauris is often larger and commonly possesses far
more distinct sub-marginal spots on the upper-side. It is
interesting to compare other mimics of echeria with the
members of the group captured by Mr. Marshall at
Malvern on March 25, 1897.

The female of Pseudacrva tarquinia is a poorer mimic
of echeria than Huralia mima and Papilio cenea &, although
upon the wing the resemblance is doubtless strong. In size
it closely approaches the model: it has a slightly- scalloped
border, and an irregular oval spot with a direction similar
to that of the other mimics; ; the marginal spots are small
as in the southern echeria. On the under-side of the
hind-wing is a basal brown patch with conspicuous round
black spots as in many Planemas. In the development of
internervular black stripes invading the squarish ochreous
patch on the hind-wing it resembles Huralia and diverges
from the Danaine model.

The female of Papilio jacksoni, in the shape and direction
of the spot below the cell on the upper-side of the fore-
wing, far more closely resembles echeria than any other
mimic I have seen. On the other hand, the hind-wing ‘1s
deeply scalloped, the effect being much intensified by
white-margined concavities, the sub-marginal spots are
as a whole larger than those of any other mimic, while
the under-surface of the hind-wing exhibits very little
approach towards the Amauvis, retaining the basal, black-
spotted brown triangle of the male, that well-known
synaposeme and pseudaposeme of many Ethiopian
Papilios, Planemas, Acrzeas, Elymnias, Pseudacrieas, ete.,
unknown in the Ethiopian Danainx. Thus, in addition
to its primary mimicry of echeria, var. albimaculata,
jacksoni manifests secondary mimetic resemblance to the
former group of Rhopalocera, especially the Planemas, and
also an approach to other mimics of echeria in its con-
spicuous sub-marginal spots and deeply-scalloped border.
The general effect of the spotting of the fore-wing is also
more like that of P. cenea, and even of Huralia mina,
than its primary model.

The female of the south-eastern Papilio echerioides is
very similar to that of jacksoni, but the spot below the
cell of the fore-wing is oval and in shape ‘and direction
more nearly resembles that of Papilio cenea.

488 Mr. G. A. K. Marshall on

F. Lhe Origin of the black-marked, golden-brown Triangle
at the base of the Hind-Wing wnder-side in many
Hihiopian Butterflies,

Mr. Roland Trimen, F.R.S., considers that the black-
marked, triangular, golden- brown basal patch on the under-
side of the hind-wing of the female P. eynorta which
mimics Planema gea, and P. echerioides [and we may now
add P. jacksoni| “ points to the inference that mimicry of
the Planema group was in both these Papiliones the
earlier tendency, and has only more recently been diverted
in the direction of Amauris in the case of the Southern
species”; for “this character is in the ¢s of cynorta and
echerioides even more developed than in the 9s, and is in
direct mimicry of the Planema” (“South African Butter-
flies,” vol. ii, 1889, p. 258). But if in the female of
cynorta, which is admitted to be an excellent Planema
mimic, this very character is reduced, how can it be
believed that its greater development in the male is a case
of direct Planema mimicry, seeing that in every other
respect this sex, if a mimic at all, is a most imperfect one ?

A comparison of the Papilios and Planemas with the
most remarkable development of this warning character at
the base of the under-side of the hind-wing renders it
probable that in this respect the latter have acted as
Miillerian mimics rather than models. The character is
far more highly developed and specialized in a section of
Ethiopian Papilios than in any of the Planemas: it also
appears in forms which are unknown, and accompanied by
other warning characters which are also unknown in the
Planemas ; it reaches its highest development in species
which do not mimic Planemas. It is also probable that
the conspicuous, sharply-outlined white band of the male
echeriodes, cynorta, etc., nearly alike on both upper- and
under- sides, is a warning character peculiar to this group
of tailless Ethiopian Papilios. In the most strongly-

marked species, with sexes nearly alike (zenobia, cypreofila,
etc.), the outer margin of the band on the fore-wing is
coarsely serrated in a very characteristic manner on both
surfaces. Furthermore (in cyprvojila, gallienus, ete.), another
warning character of great interest is added in the row
of large conspicuous marginal white spots on both surfaces
of the hind-wing and smaller ones on the fore-wing. In
this respect this group of Papilios presents an exact

The Bionomics of South African Insects. 489

negative of the positive form of synaposeme character-
istic of the Pierme genus Mylothris. Thus very striking
warning characters are peculiar to this section of Papilios,
the fifth or*Zenobia Group of Aurivillius; and, further-
more, the character we are specially considering, the black-
marked basal patch of the under-side of the hind-wing,
assumes a form which is unknown in Planema, bemg tra-
versed by blackened veins and broader black internervular
lines. In the species of Papilio last mentioned there are no
spots upon the brown triangle, only these strongly-marked
radiating lines. In zenobia and still more in the male
cynorta « few spots are added by modification of some of
the other markings, and it is probable that this shght
change is a late diaposematic response to Planema, made
after the latter had gained the golden-brown triangle in
Miillerian mimicry of these dominant Papilios.

Another important point is the fact that the golden-
brown triangle is /arger in the females than the males of
eypreofila and zenobia which do not mimic Planemas,
smaller, as has already been pointed out, in the females of
the species which strongly exhibit this Miillerian approach
to either Planema or Amauris,

It must also be remembered that Papilios may be
excessively unpalatable to insect-eating animals as a
whole. Thus Mr. Frank Finn concludes that P. avisto-
lochiv is more distasteful to birds than Danaine, Acrea
viole, and Delias eucharis (Journ, Asiat. Soc. Beng., lxvii,
pt. a1, 1897, p. 614).

The facts and arguments set forth above render it
probable (1) that the golden-brown triangular patch first
arose in the Zenobia Group of Ethiopian Papilios; (2) that
it was later reproduced on a smaller scale by the Acreine
genus Planema, the Acreine round black spots contrasting
with the ground-colour in place of the radiating black lines
of the Papilio; (8) that, later still, other widely- separated
genera reproduced the character in the form’ it had as-
sumed in Planema, e.g. Pseudacrva, Elymnias, ete., while a
reciprocal (diaposematic) tendency (see p. 426) to approach
the Planema form is seen in certain species of the Zenobia
Group of Papilios. Several species of the group have
females mimicking Acrzines or Danaines. In the three
of these which were examined the males exhibited the
above-described diaposematic tendency, while the females
possessed a greatly-reduced but otherwise similar triangular

patch.

490 Mr. G. A. K. Marshall on

Hence this characteristic widespread Ethiopian synapo-
seme and pseudaposeme has probably originated in a
diaposematic fusion of the triangular golden-brown patch
of the Zenobia Group of Papilios with the scattered
circular black spots which are characteristic of Ethiopian
Acreas. I have made much use of Aurivillius’ admirable
“ Rhopalocera Ethiopica” in this section which is devoted to
the discussion of an under-side synaposeme, although the
distinguished author himself maintains that mimetic
resemblance is almost confined to the upper-side of butter-
flies’ wings—a very strange conclusion (/oc. cit., p. 585).

G. Compound Group containing Representatives of all the
L ues
three previously described. Species probably entering
two Groups.

The groups described above fly together, and thus repre-
sent in a compound group the chief types of butterfly color-
ation which a young insect-eating animal of South and
Eastern Africa requires to learn, by a trial of one or more
representatives. The following members of the three
groups were captured by Mr. D. Chaplin at Berea, a
suburb of Durban, on April 5, 1896, and are now in the
Hope Department.

BLACK-AND-WHITE GROUP. Echeria-L1KE GROUP.
Amauris ochlea. Amauris echeria, var. albi-
Planema aganice &. maculata.

Huralva mina.
Chrysippus-LIKE GROUP.

Limnas chrysippus §.

2 Hypolimnas misippus f, 2 type-form.

2 Acrea petrxa § 9.

2 Acrwva encedon, type-form and var. Lycia.

That the same species may produce two or more forms
entering as many groups is well known, but, as a rule,
such polymorphism 1 is confined to the female sex. In the
polymorphism of <Acrwa encedon, however (see pp. 483,

484), we have a case in which both sexes are present in
the various forms, and although the relative numbers of
the forms are very different and certain of them may
perhaps be absent from a district, | know of no case in

The Bionomics of South African Insects. 491

which one alone is found in any part of the total range of
the species. Hence the polymorphism, although partially a
distributional phenomenon, is not entirely so. On the
other hand, I know of no example among the Lepidoptera
in which a species is at the same season divided into two
sub-equal sections throughout its range, each containing
both males and females, and each “section mimicking
a very different model. Among Diptera, the European
Volucella bombylans and its form mystacea supply good
examples; and now Mr. Marshall has collected evidence
which makes it in the highest degree probable that the
Lepiduptera are not without such cases. He brings
convincing support for the belief that Buralia wahlbergr
and H. mima are the two forms of a single species. It is
greatly to be hoped that Mr. Marshall may be as success-
ful in establishing this interesting and unique case, as he
has been in the marvellous seasonal transformation of
Precis, His evidence is set forth in the following quota-
tions from his letters :—

“Umkomaas Mouth, Natal; Sept. 3, 1897.—In my own
mind I am pretty well convinced that Huralia mima and
wahlbergi are one and the same species which has developed
two mimetic forms as in Papilio cenea, but that in this case
both sexes are concerned. My reasons for so thinking are
that they have been taken in coitu several times, “that
specimens occur presenting intermediate coloration, and
also that the two forms are always found together whenever
they are met with in any number. I have not often been
fortunate enough to see these congregations, but I re-
member seeing some thirty or forty specimens, comprising
about equal numbers of each form, collected together on
the side of a steep shady kraantz along the Palmiet River.
Mr. A. D. Miller, who has collected for many years in
Durban, tells me this is by no means uncommon, and
that they congregate particularly in the afternoon when
going to roost. Mr. C. N. Barker tells me that some years
ago he came across a large number of both forms on a
large tree on which they had evidently bred, for many of
them had only just emerged, and some had not their
wings fully developed. This shows that they are also
associated in their earlier stages.”

About the time when this passage was written Mr.
Marshall presented to the Hope Department a set of
eight individuals of these Euralias, viz. four of #. mima (2 f,

4.92 Mr. G. A. K. Marshall on

2 2), and four of £. wahlbergi (3 J, 12), which he had cap-
tured on the Umbilo River, near Durban, Natal, on June
28,1897. His account of the habits of this little com-
pany is given below.

“Malvern, Natal; Oct. 7, 1897.—The specimens of
Luralia mima and wahlbergi were captured going to roost
together on a small clump of ferns under a steep kraantz
between 3 and 4 p.m. Although disturbed a good many
times in my efforts to catch them, they always returned
after some minutes. There were two others which were
too tattered to keep, and two more that I failed to catch.”

If Mr. Marshall’s conclusion be established, it follows
that the corresponding and closely-allied mimetic West
African forms Huralia anthedon and E. dubia, connected
like wahlhergi and mina by intermediate varieties, are
similarly the dimorphic forms of a single species.

H. Groups of Synaposematic Acrvas captured at the same
Place and Time.

Professor Meldola first suggested the use of Fritz
Miiller’s principle to explain “ the prevalence of one type
of marking and colouring throughout immense numbers
of species in protected groups, such as the tawny species
of Danais, the barred Heliconias, the blue-black Hiupleas,
and the fulvous Acrwas” (Ann. and Mag. Nat. Hist.,
ser. 5, vol. x, 1882, p. 425). As an example of Miillerian
mimicry in the last-named group, I was anxious to obtain
convergent species captured in one place and at one time.
Mr. Marshall very kindly obtained two such groups for
me. The first was captured by him on Dec. 31, 1898,
at Salisbury, and contains the following species :—

6 Acrva doubledayi, var. axina, 4 $2 2 (¢ Fig. 1, 2
Fig. 2, Plate XVI).
4 Acreva caldarena, 3 f 1 2 (¢ Fig. 3, 2 Fig. 4, Plate
XVI).
4 Acrea nohara, var. hala, 2 § 2 2 (¢ Fig. 5, ¢
Fig. 6, Plate XVI).
2 Acrea violarum, var. asema, 2 9 (Fig. 9, Plate
XVI).
2 Acrva rahira, 1g 12 (¢ Vig._7, 2 ieee late
XVI).
All these species are of approximately the same size
and shape of wing, colour, and pattern, rahira being furthest

The Bionomics of South African Insects. 493

removed from the average appearance of the group, while
caldarena is an outlying member in one respect only, viz.
the pronounced apical black patch on the fore-wing. The
brightly-coloured males and the brightest of the ‘females
of all these species would closely resemble each other on
the wing, including the male of violarwm, which was not
captured on that particular day; and similarly a uniform
effect would be produced by the darker females. The
under-sides of all species except rahira are superficially
alike. The strong superficial resemblance is well shown
in Figs. 1 to 9 on Plate XVI.

A second group was captured at Salisbury on Jan. 7,
1899, and contains the two following species :—

2 Acrva anemosa f 2 (2 in Plate XVI, fig. 10).

i=)
Zz ,, natalica 2 (Plate XVI, fis. 11).

These large Acrzeas are obviously very different in
the details of coloration, but the positions in which the
black marking of the upper surface are massed on the
fulvous ground-colour are almost exactly the same, the
only marked difference being the presence of numerous
black spots in the first-named species which are almost
absent in the second. Mr. Trimen speaks of aneimosa as
“in habit and general colouring very near natalica” (loc.
cit., vol. i, p. 156). The under-sides are far less alike, but
there can be no doubt about synaposematic convergence
having occurred, It is probable that the approach has
been chiefly, perhaps entirely, on the side of natalica,
which has adjusted markings of a type usual among
Ethiopian Acrvinw in such a manner as to produce super-
ficial similarity to anemosa, an Acrva in which a very
remarkable and unusual appearance is the warning sign
of exceptional defence against insect-eating animals (see
p: 413):

Mr. Marshall informs me that the two species are very
similar upon the wing, and that the resemblance is much
closer in the case of the female natalica than the male,
thus following the rule in mimicry, and confirming still
further the opinion expressed above that the approach

has been from the side of natalica.

I. Mimnetic Species of South African Lycenide and Hesperidx
captured with their Models. .

Exclusive of Mimacreva marshalli and its form dohertye

4.94 Mr. G. A. K. Marshall on

which were discussed in relation to their models chrysippus
and Alugit, the groups containing Lycenide or Hesperidx
are considered below.

A group of the smaller Acreeas with a single mimetic
Lycxnid was captured at Salisbury on Sept. 28, 1900.
It contains the following species :—

2 Acrxa violarwin, var. asema ff.

1, = doubledayt, var. axina &.

1, wduna fo.

1 Catochrysops mashuna f (figure of 2 on Plate XIV,
fig. 5).

A. induna falls into the first-mentioned group of small
Acrias, resembling ca/darena in the possession of a black
apical patch to the fore-wing. The strong development of
black spots upon an ochreous ground on the under-side of
the Lyczenid is doubtless mimetic in the position of rest,
especially when in the company of Acraas or in places
where they are likely to be found. Mr. Marshall informed
Mr. Trimen “that on October 20, 1894, he saw two of this
Lycena sleeping on the end of a stem of dry grass among
a number of Acrewa nohara and A. caldarena, and was
struck with the general similarity of their under-side to
that of the Acrzas; he also noticed that in the attitude
of repose the fore-wings of the Zycena were well depressed
between the hind-wings, giving the insect the elongate
outline of an Acrva” (Trans. Ent. Soc. Lond., 1898, p. 6).
Mr. 'Trimen also suggests that the heavy black spotting of
L. (C+) gigantea and L. (C) perpulchra (Trim.) [ = peculiaris,
Rog.| is of the same significance, although the ground-
colour is not Acrwa-like in these species.

A paragraph from one of Mr. Marshall’s letters is quoted
below.

“ Umkomaas Mouth, Natal ; Sept. 3, 1897.—The under-
side of the Acrwa-like Lycenid Z. mashwna is ochreous
yellow with large black spots, but I did not realize its
resemblance to an Acrexa until I saw them roosting to-
gether. The mimicry, however, is in a very incipient stage,
for the yellow under-side fades somewhat rapidly, thus
much lessening the resemblance ; and, as it does not occur
in its near allies Z. hypolewca and the fine L. gigantea, it
is clearly a recently-acquired character.”

In another group captured at Umtali, 3700 feet, in
December 1900, the Acrieas are only represented by a single

The Bionomies of South African Insects. 495

species, and that not specially suitable as a model for the
other members. The great interest of the assemblage is the
presence of a rare Hesperid, A. fetfensis, in which black
spots and a pale pinkish tinge on the under-side of the
hind-wings appear to indicate strongly-marked mimicry
of Acrzeas on a line along which the above-named Lyczenids
have advanced to a greater distance. In another respect,
however, viz. the strongly-marked black-and-white margin
of the hind-wing under-side, the Hesperid is a much closer
mimic of a general Acrwva type. Mr. Trimen also speaks
of the spotting and tinting of the under-side of both wings
of this species as very peculiar and strongly recalling the
aspect of some of the smaller Acraeas (Joc, cit., vol. 111, p.
338).

I have received from Mr. Marshall the following refer-
ences to the two Hesperids in the list of species making
up this group :—

“ Salisbury, Jan. 11, 1901.—I have recently obtained at
Umtali another Hesperid, the very rare Abantis tettensis,
showing strong Acrzoid coloration on the under-side,
which I wili send you together with a Aedestes, which is of
interest as it shows the incipient stages of such mimicry.”

“ Salisbury, Sept. 27, 1901—I have never seen Abantis
tettensis with its wings closed over its back ; all that I have
captured rested with wings expanded horizontally. Indeed
I cannot at the moment recollect ever seeing any Abantis
settle with vertical wings. I think it is highly probable
that ¢etfensis would sleep in that position, but under the
circumstances I should not like to assume it as a fact
without actually seeing it.”

The group 1s as follows :—

3 Acrxa encedon, type-form.
2 Catochrysops peculiaris (Plate XIV, fig. 4).

2 Abantis tettensis (rs Tae ce <0
1 Kedestes macomo, var. (_,, RS BE

In the latter Hesperid the brilliant ochreous under-side
affords an effective background for the small but distinct
black spots on the under surface of both wings. The
general effect is somewhat Acrwva-like, but the tint of the
under-side of most smaller Acraeas is pinkish when they are
fresh.

A. more perfect Hesperid mimic is seen in the rare
Baoris netopha, of which a female (Fig. 13, Plate XVI) was

496 Mr. G. A. K. Marshall on

captured by Mr. Marshall at Salisbury on April 6, 1898,
together with a male of dAcrxa doubledayi, var. axina
(Fig. 12, Plate XVI), which is one of the many smaller
Acreeas, to which it bears a somewhat generalized resem-
blance on the under-side of its wings in the natural posi-
tion of rest, which is nearly but not quite represented in
Fig. 13 (see the description of the Plate). Mr, Marshall
has sent the following account of the attitude :—

“ Salisbury, Feb. 12,1899.—The Baoris netopha rest with
closed wings, and the fore-wings pressed well within the
hind-wings so as to hide the white spots; they then look
much more Acrwva-like.”

The curious reticulate under surface of the hind wings
of the isolated and remarkable Hesperid Cyclopides willemi
is mimetic of the probably distasteful Alena nyasse, which
possesses a somewhat similar but much coarser reticulation.
My. Marshall states that the resemblance is much enhanced
in the resting attitude of both species by the concealment
of the fore-wings, with the exception of the apex, within
the hind. Both species frequent the same localities, and
both rest upon grass-stems. One of each species was
captured at Salisbury on Feb. 23, 1901, and another similar
pair on March 8, 1901.

In thus bringing together Mr. Marshall’s examples of
mimicry in Hesper ide, it is appropriate to include the
following interesting case of mimicry on the part of a
Hesperid for a Danaine larva.

* Salisbury, March 10, 1898.—The larva of the large
‘skipper’ Rhopalocampta forestan possesses a colouring
wonderfully similar to that of Z. chrysippus, though it lacks
the filaments. I only know the larve of three other
species of Hesperide and they are all green; moreover,
they form shelters for themselves, and never come out to
feed except after dusk; whereas, although /orestan also
forms a shelter, yet it frequently comes out and feeds in
broad daylight, when it is a very conspicuous object.”

The upper-side of Alena nyasse appears to fit in with
the strong combination of black-and-white Ethiopian
butterflies belonging to the Danainwy and Acreine and
their Batesian and Miillerian mimics. Among the smaller
of the latter Neptis agatha is probably to be placed, and
this species is on the wing with the much smaller Lyczenid.
Thus Mr. Marshall has sent to the Hope Department
specimens of the Veptis captured at Salisbury on March

The Bionomies of South African Insects. 497

19 and May 1, 1898, and of the Alena taken on March 20
and April 3 of the same year.

It occurred to me that Castalius calice might also belong
to the same group, but Mr. Marshall points out, in the
passage quoted below, that its habits do not support
this view.

“Salisbury, Jan. 8, 1899.—I should very much doubt
whether Castalius calice is convergent with or even a
mimic of Alwna nyasse. Their habits and stations are
very different, and moreover C. calice (of which I believe
C. melena will prove to be the summer form) is common
in Natal and the Transvaal, where A. nyasse does not
occur. I should not regard C. calice as an unpalatable
species, and its colouring is by no means conspicuous owing
to its small size: it 1s an active little insect resembling
T. plinius, Lycenesthes, and other arboreal Lycenide in its
habits. In the intense light and shade of this climate its
black-and-white markings are rather protective as it rests
on the shiny leaves of its food-plant (Zizyphus), just as are
the brilliant white under-sides of some Jolat. The con-
vergence you suggest between A. nyassev and Neplis agatha
and Nyctemera lewconoé is highly probable, but Amauris
and the black-and-white Acreas are all absent from the
Mashona plateau, being all coast or low-veldt forms.
Alxna, Pentila, and perhaps Delonewra, are in my opinion
the only unpalatable South African Lycwnide, and the
latter is more likely to be a mimic of some day-flying
moth. Catochrysops mashuna used to be very abundant
here, but only occurring in September and October. I only
saw two or three this season and always when I had no
net.”

Three specimens of another interesting and probably dis-
tasteful species of the same Lyczenid genus Alena amazoula
captured on the same day, Sept. 26, 1897, as the conspicu-
ous day-flying and probably unpalatable geometrid moth
Petovia dichroaria were presented by Mr. Marshall to the
Hope Department. Mr. Marshall had taken the group in
the same locality at Malvern, Natal, and, as the passage
from his letter quoted on p. 498 indicates, he believes that
the resemblance is synaposematic. In the cabinet the
likeness is stronger on the under than upon the upper
surface, but is probably strongest of all upon the wing.

Alxna amazoula is a Lyceenid of great interest, probably
exhibiting a generalized Miillerian resemblance to the

TRANS. ENT. SOC. LOND. 1902.—PART II. (NOV.) 33

498 Mr. G. A. K. Marshall on

Acreas. Although unlike any single species of Acrwa, the
likeness to this group and unlikeness to the Lycwnide was
sufficient to deceive Boisduval and at first Roland Trimen
(Rhop. Afr. Austr., 1862-66, p. 111). In the Hope Col-
lection also I found it had been placed among the Acrzeas
by Professor Westwood. Its undoubted Lyczenid affinities
were finally established by Roland Trimen (“South African
Butterflies,” vol. 11, 1887, p. 222).

The above-mentioned errors as to the affinity of this
aberrant Lyczenid certainly support the opinion that it
bears a general resemblance to the Acrwinx. It would be
interesting to know its resting habits. With the under-
side exposed and the long narrow wings it would probably
bear some general likeness to a very small Acrwva. Mr.
Marshall wrote concerning it as follows :—

“Umkomaas Mouth, Natal; Sept. 3, 1897—Alxna ama-
zoula is certainly a protected species, but I do not think it
is In any way convergent towards the Acrzeas, for it is in no
way suggestive of them on the wing, being by no means
conspicuous, but rather difficult to follow. Its length of
wing has been attributed to relationship with Acrza, but
this seems open to doubt. I should prefer to consider it
as a parallel development to Acrva, though it is worth
noting that the allied genus Lachnocnema has also some-
what elongate wings.”

“ Malvern, Oct. '7,1897.—The day-flying moths [Petovia
dichroaria| captured on the same day as Alena amazoula
can, I think, be well regarded as convergent in coloration.”

J. Mimicry in Lycenide and to a less extent in Hesperidx
a Character of the Ethiopian Region. Possible Inter-
pretation.

The instances of mimicry in South African Lycenide
recorded here, and the much larger number known in
other parts of the region, especially the tropical West
Coast, led me to inquire how the total number of species
of this family compared with that of other parts of the
world. My friend Mr. Hamilton Druce kindly made an
approximate calculation of the number of described species
in the two other great tropical south-extending land
masses. From Australia, the Malay Archipelago, and the
continental portion of the Oriental Region, over 1000
species have been described. From the Neotropical Region

The Bionomics of South African Insects. 499

about 700 species of Z’hecla have been described. In Auri-
villius’ catalogue of Ethiopian Rhopalocera only 582 species
are recognized. The predominance of Lycsenid mimicry in
this latter Region is therefore in no way connected with
richness in the number of species. The chief reason is
certainly the existence in the Region of the sub-family
Liptenine, with nearly all its species mimetic. In addition
to the general Acrzine appearance of Alena amazoula, the
Neptis-like A. nyasse and the chrysippus-klugii-lke Mim-
acrea marshalli-dohertyi, there is represented in the Hope
Collection mimetic resemblance to Zerias or other small
Pierines on the part of Larinopoda lircea, L. tera, Liptena
libyssa, and L. undularis ; to Mylothris by Pentila abraxas,
P. phidia, and Citrinophila erastus ; to chrysippus and the
chrysippus-like Huphedre and Aletis by Telipna bimacu-
lata and T’. sanguinea ; to a general Acreine type by two
or three species of Pentila. Many other cases of mimicry
are known in the sub-family, especially towards models of the
genera Planema and Acrwva (see Aurivillius, loc. cvt., p. 530).
But this remarkable group does not by any means exhaust
the Ethiopian Lyczenid mimics, for many species of the

Lycenine mimic Acreas, Terias or other small Pierines
and Mylothris. The general Acreine mimicry of species of
Catochrysops has already been described and illustrated.
Furthermore, Aurivillius considers that there is a certain
amount of mimetic approach between species of Liptenine
and Lycening in which the former probably always act as
models.

It is very difficult to understand this predominance of
Lycenid mimicry in the Ethiopian Region, and I can only
suggest the possibility that the number of feasible models
of moderate and small size furnished by the abundant
Acreine of Africa may furnish an explanation. In such
cases as Catochrysops peculiaris and mashuna we see at
once how naturally and easily the Lycznid under-side
adapts itself to the characteristic appearance of the Acreu
type, especially when it is further assisted by similar habits.
And this suggests another equally important principle
which has doubtless been fertile in bringing about Lyczenid
mimicry, viz. the habits of the models being such as to
bring them within the range of the forms which were to
mimic them. The numerous low-flying and low-settling
Acreas, resting at night on grass-stems, have precisely the
mode of life which is well known to be characteristic of

500 Mr. G. A. K. Marshall on

such a large number of Lycenide. The Liptenine are
probably a specially-protected group, and the rarity of
many of the species may be only apparent, and due to
their (Miillerian) mimicry of extremely common forms for
which they are constantly mistaken by naturalists.

In other parts of the world Lycenidx which are evidently
specially protected and extremely conspicuous are well
known, such as Yalicada of the Oriental Region and
Humeus of the Neotropical. These genera do not enter
into synaposematic association with the best-defended
butterflies of their localities, but each adopts an aposematic
appearance peculiar to itself. It is probable that in the
Neotropical Region, where mimicry is more striking and
more fully exemplified than in any other part of the world,
the habits of the Theclas are the chief obstacle to their use
of this means of protection. For models of all sizes abound
in this part of the world. On the other hand, in the
tropical East it may be the want of a sufficient number of
models of an appropriate size and habits which has acted as
the barrier.

The explanation which has here been thrown out as a
suggestion may also enable us to understand the cases of
mimicry in Ethiopian Hesperide, a family im which such
resemblances are rare. Here, however, the facts may be
paralleled in the Neotropical Region where there are a
few mimetic “skippers.” Mimicry in this group and the
want of it in the Lycewnide may follow from the difference
between the habits and stations of the tropical American
Hesperids and Theclas.

K. Mimicry in the Nymphaline Batesian or Miillerian ?

I have had much controversy with my friend Mr.
Marshall over this difficult and interesting problem, and I
propose to bring forward a résumé of the arguments which
seem to support the latter interpretation as opposed to
the former, and then to quote his weighty objections and
the interesting observations of the habits of mimetic
species and genera which he has made.

1. It is of interest although probably not of extreme
importance to reflect that all the great groups of un-
palatable, conspicuous, and much-mimicked butterflies
belong to the Nymphalide—the Ithomiine, Danaine,
Heliconine, and Acrvinx, and that the two latter are so

The Bionomics of South African Insects. 501

closely related to the Nymphaline that it is difficult to
draw a line between them. The argument is not of much
weight, because the intensely procryptic habits and colours
of many Nymphaline genera have certainly been brought
about by selection due to the great keenness and suc-
cess of insect-eating animals in their pursuit. I have
however suggested and brought evidence in support of the
view that some of the procryptic Nymphaline species are
to a certain extent unpalatable (see p, 442).

2. Mimicry in the Nymphalinew does not appear in
isolated forms but in all or nearly all the species of a
genus. Such mimetic genera are usually very large,
dominant, and wide- spread. The species themselves are
also often wide-spreading, and may have an enormous range
far exceeding that of the model (ypolimnas misippus).
Allowing for the fact that the mimetic species resemble
the commonest types in the world, and so are hable to
escape notice, it is probable that they are rich in indi-
viduals. In many instances we know that this is so.
The more we investigate it the more does Rhopaloceran
mimicry seem to be associated with dominant genera and
species, rather than the feeble and hard-pressed forms
which H. W. Bates presupposed in his well-known theory.

3. The dominant tendency towards mimetic resem-
blance in any genus cannot be explained by hereditary
transmission of the mimetic form of a single parent species,
or from the tendency of closely-related species to vary along
nearly the same lines, because the species of a mimetic
genus, as a matter of fact, mimic in many different direc-
tions. Thus Pseudacrva resembles Acrewa, Planema,
Amauris, and Limnas chrysippus; while Hypolimnas,
including Huralia, is even more protean.

4. The non-mimetic species of a mimetic genus are
often markedly conspicuous, exhibiting what has all the
appearance of an aposematic pattern peculiar to them-
selves (Hypolimnas, Pseudacrea). This is also frequently
true of the non-mimetic males of a species with mimetic
females (Hypolimnas). Such aposematic patterns are
especially displayed on the under-side, where procryptic
colours are developed in other butterflies.

5. The converse of the last argument is also true, viz.
some of the species in a genus, which is as a whole
markedly conspicuous and itself mimicked, are often
mimetic of quite other groups. Many instances of Neptis

502 Mr. G. A. K. Marshall on

have been given on pp. 467, 468, and mimetic species are
also well known in Li menitis, Cethosia, ete.

6. The non-mimetic species of mimetic genera are
sometimes mimicked; e.g. the mimicry of ‘the nerina
form of female Hypolimnas bolina by a rare Danaine in
Celebes, etc. The resemblance of the upper-side of cer-
tain species of Protogoniomorpha to some of the larger
species of /Hypolimnas may be another instance of the
same tendency.

7. The fact that mimetic species resembling some
primary model nevertheless in certain respects resemble
each other rather than the model. This deuterosynapo-
sematic resemblance, as it may be called, is a very wide-
spread phenomenon, and several striking instances of it are
discussed in the present memoir (see pp. 470, 471, 485-7).
It will also be shown to occur in Coleoptera (pp. 513-515).

I have given merely an outline of the chief evidence
which has induced me to believe that the mimetic
Nymphaline genera are to some extent specially pro-
tected, and thus have developed a beneficial synaposematic
association with far better protected forms belonging to
other sub-families.

This evidence has been sought and obtained under the
guidance of the principles discovered by Dr. F. A. Dixey
and ably presented by him in 1894-97 (Brit. Assoe.
Reports, 1894, pp. 692, 693; Trans. Ent. Soc. Lond.,
1894, p. 2935 1896; “p. 655) 1897, p. Sif), aihese
memoirs mark one of the few important advances made
in our attempt to understand the complex and diff-
cult phenomena of mimicry. I will quote one preg-
nant paragraph, which it will be seen contains the
essence of what I have here described as primary and
secondary mimicry (see pp. 513-515, also the above para-
graph 7). “Every conspicuous and distasteful form is a
centre of attraction for other forms, whether edible or
inedible; but in the former case (Batesian mimicry) the
mimetic attraction is limited in operation, and acts only
in one direction, influencing nothing but the mimic;
while in the latter case (Miillerian mimicry) the mimetic
attraction is unlimited and mutual, acting reciprocally in
both directions, and influencing each member of the group”
(Trans. Ent. Soc. Lond., 1897, pp. 324, 325).

Mr. Marshall’s valuable notes on the habits of Nymphaline
mimics and his discussion of the conclusions described

The Bionomies of South African Lnsects. 503

above, are contained in the following quotations from his
letters.

“ Malvern, Natal; Feb. 21, 1897.—As regards the Euralias
my experience of them is somewhat limited, but from what
I have seen of them I feel pretty sure that their coloration
is due to mimicry and not to convergence. Their range
seems in all cases to agree with that of the Amauwris they
resemble. In Durban A. ochlea is by far the scarcest of
the three, and its mimics ZH. deceptor and Pseudacrea ex-
pansa are also very rare ; further up the East Coast, how-
ever (Delagoa Bay and Beira), ochlea becomes one of the
commonest of the genus, and the two other species are
likewise much more numerous. At this place, it is true,
Huralia wahlbergi is certainly more plentiful than A.
dominicanus, but they are both uncommon, and all the
specimens we see are practically visitors from the thick
bush along the immediate coastline, (This is six miles

inland.)”

* * % % x

“TY may mention that in the last few years Mr. Ball has
caught two Huralia wahlbergi [in the Karkloof Forest
twenty miles N. of Maritzburg], but has never seen
Amauris dominicanus.”

“ Umkomaas Mouth, Natal; Sept. 3,1897.—With regard
to the Euralia, ete., I must admit that I have never been
fully convinced by the contention that in Batesian mimicry
the mimetic species must of necessity be a feeble one and
very fewin numbers. It seems to me that it would be quite
reasonable to suppose that such a mimic might well equal
or even exceed in numbers the protected species, though
this would of course depend entirely upon the degree “of
inedibility of the latter. For example, in the case of
Luraha mima and Amauris echeria (probably the best-
protected butterfly here), supposing they occur in equal
numbers in a given area, and that certain birds by chancing
to catch three or four of the former in succession were
induced to prey upon butterflies with that coloration,
then, from a mathematical standpoint, every alternate
specimen caught by any bird would be A. echeria. Now
I think we are quite safe in assuming that the fact that
every other butterfly caught had a nauseating taste
and smell would be far and away more likely to create
a strong and lasting impression upon .a bird’s mind

t=)
than the fact that every second one proved to be edible,

504 Mr. G. A. K. Marshall on

and would be quite sufficient to deter the bird from
attempting to eat butterflies of that colour. This is from
a mathematical point of view solely, but, from what I
know of these two forms in life, I believe that presuming
them to occur in equal numbers, a larger proportion of
echerva would actually be captured, for mima is a much
more shy insect, and although it has the same slow sailing
flight (when undisturbed) it does not keep on the wing
nearly as long as echeria ; moreover, it is much more wary
and always on the alert for danger, going off at a smart
pace when frightened, and not returning to the same spot
as echeria frequently does after being struck at; altogether
it is a much more difficult insect to capture. Indeed I
do not see why the mimic should not even somewhat sur-
pass the mimicked species in numbers, without upsetting
their relations to one another, provided the taste of the
latter be sufficiently unpleasant, and particularly if the
flavour be of a dasting nature.”

“ Malvern, May 14, 1897.—I feel quite satisfied that
Pseudacrxa trimenii is a mimetic and not a_ protected
species. In spite of its larger size it looks wonderfully
like Acrwa acara on the wing, and the first few examples
I caught completely took me in. Their flight is lke that
of all Pseudacrzeas and Euralias—slow and _sailing—so
long as they are not disturbed ; but if struck at and Tae
they are off like a shot and do not often give one a second
chance. At this particular spot (Malvern) they are a good
deal commoner than 4, acara, which is only a rare visitor.
The latter is however common on the immediate coast,
where P. trimenii is I am told pretty plentiful in good
seasons.”

“ Salisbury, Jan. 12, 1901.—I quite agree with you that
the resembiance between the under-sides of Delias pandemia
and Isbarta pandemia is the most remarkable case yet
brought forward, and one cannot but marvel how such
exact similarity can have been arrived at. Although I
should certainly incline to the belief that the mimicry is
Miillerian, judging by the congeners of both forms, yet its
very exactitude seems to be a difficulty, for although one
can readily understand how in an edible and “much-
persecuted species the resemblance might be brought up
to so high a grade, yet it is hard to understand how this
could be effected in a species which is comparatively
immune from attack. For it seems to be an inevitable

The Bionomies of South African Insects. 505

deduction from the theory of mimicry that a high grade
of resemblance must imply excessive persecution, either
now or within recent times, unless we are prepared to
admit some other convergent force.”

“Salisbury, Sept. 217, 1901.—With regard to Miillerian
mimicry I quite agree that the slow flight and the bright
colours of protected forms would lay them open to much
experimental tasting from inexperienced birds; but the
extreme toughness of their integuments (for example in
the Acraeas) and their great vitality seem to have been
specially developed to minimize thisdanger. And whereas
the Batesian mimic, if thus experimented with, would
promptly be eaten, a Miillerian butterfly would run a
very good chance of surviving its injuries and propagating
its kind. Do not suppose that I in any way disbelieve in
the action of Miillerian mimicry, for 1 certainly think it
must be a very strong factor. But this point seems to me
to be a real difficulty, and I should hke to be able to
answer it satisfactorily if it were brought up by an
opponent.”

L. Miscellaneous Observations on Mimiery in South
African Rhopalocera. (G. A. K. M.)

a, Nymphaline.

“ Malvern, Natal; Feb. 21, 1897.—I do not remember
ever having scen it suggested that the female of the hand-
some Charaxes xiphares mimics A. echeria, but I have
little doubt that such is the case. It is a fairly common
species at the Karkloof, but difficult to catch, and I only
took one. I believe there are more instances of mimicry
in this genus, e.g. achwnenes and guderiana, of which the
females are very scarce (though the male of the former is
one of the commonest Charaxes in Mashonaland) and very
differently coloured from the males, being remarkably
hke the common widespread C. satuwrnus. Again, some
years ago I pointed out to Trimen the strong resemblance
of the female of C. whytei to Neptis agatha on the wing.
I hope to be able to prove before long that Neptis is a
distasteful genus (by the way, is Limenitis edible 2), as its
appearance ‘and habits certainly point that way.”

B. Pierine. ‘
“ Estcourt, Natal; Oct. 15, 1896.—In Durban Mylothris

506 Mr. G. A. K. Marshall on

agathina was fairly abundant, and I was interested to
notice the very close resemblance between it and Belenois
thysa, though, as is the case with most mimics, the latter
has a much quicker flight when disturbed. &. thysa, so
far as my experience goes, is confined to the warm coast
belt, whereas JZ. agathina is common everywhere through-
out South-east Africa.”

“ Malvern, March 12, 189'7.—It is curious to note that
although Nepheronia argia was common at the Karkloof,
I never saw a single specimen of Mylothris agathina. My.
Ball has in his collection a ver y fine variety of the female
of the former, which clearly mimics Mylothris trimenia, the
upper-side of the hind-wings being lemon-yellow, and the
red mark on the under-side of the fore-wings absent. This
is the only one I have seen, although I took. one or two
females showing an approach towards it, one of which I
send you.”

“Umkomaas Mouth, Natal ; Sept. 3, 1897.—I am afraid
I can hardly bring myself yet to believe in Dixey’s theory
that the Pierinw are for the most part protected. As
regards our South African species, the only ones which I
feel confident are protected are Mylothris agathina,
ruppellit, and trimenia, and Pontia hellica. For the
remainder I think we must find some other cause to
account for their numbers and wide range. In Terias the
larva possesses wonderfully assimilative colouring, and is ex-
tremely difficult to detect on the food-plant, and so far as the
experience of Hutchinson and myself goes with 7. brigitta
we have never yet observed a single. case of parasitism.
The protective coloration and comparative freedom from
parasites of the larva, and the protective seasonal colouring
of the imago, would no doubt go a long way to explain
their number, though I fancy there must be some other
factor.”

“ Salisbury, May 1, 1899.—Query : are the black bands
in the females of Teracolus acquired in mimicry of the
common and widespread species, Herpenia eriphia?” *

y. Papilionine.
“ Salisbury, Dec. 26, 1897.—Papilio brasidas I consider

* Dr. F. A. Dixey informs me that Professor Westwood probably
suspected a relationship between the markings of H. eriphia and
T’. eris, for he had removed a specimen of the former species from
the others, and placed it next to a dry form of the Teracolus.

The Bionomies of South African Insects. 507

to be merely a local race of P. leonidas, as every one must
do who has seen a long series of the two forms. I have
found typical /eonidas pretty plentifully in the low veldt
of Mashonaland (Mazoe and Umfuli Rivers) and I also saw
it at Delagoa Bay. I have always been struck with its
marked difference in habit from the Southern brasidas.
Its flight is strong and rapid, and it always goes straight
ahead like P. policenes and P. anthews, which it somewhat
resembles on the wing in spite of its very different shape.
Brasidas,on the other hand, has a slow sailing flight, going
backwards and forwards over the same ground and often
frequenting one spot for days. Now there is absolutely
nothing suggestive of protection in the flight of /eonidas,
none of that slow sailing movement to show off its coloration
which is so characteristic of the protected Danainx and
Acrvinz. Moreover, there is no Danaine occurring south
of the Zambesi which is anything like it at all, and this
is very significant. I cannot therefore resist the conclu-
sion that in this country leonidas 1s one of those un-
protected species which has succeeded in the struggle for
existence by its strong rapid flight, and perhaps by pro-
tection in the larval stage hke P.demodocus and P. corinneus,
whereas in Natal it has found it advantageous, owing to
the abundance of Amauris echeria, to adapt its coloration
in mimicry of that species by the reduction in size and
number of the spots in the fore-wing and the toning down
of the colour from glaucous green to greenish-white,
accompanied by the marked change in its mode of flight.
It does not seem to me that convergence would explain
the facts, for if /eonidas is itself protected it should exhibit
throughout its range that slow flight which is the ‘hall
mark’ of protection, which it certainly does not in
Mashonaland. I believe in Central Africa it is said to
mimic 7’. petiverana, and it would be most interesting to
find out whether it has there assumed the Danaine flight.”

“ Malvern, Feb. 21, 1897.—1 have been collecting in the
Karkloof Forest some twenty miles north of Maritzburg for
the last three weeks. The only Amauwris occurring there
is echeria, which is very common, though not this year,
which is a curiously abnormal one, and as usual the
typical female of Papilio cenea is common, the dominicanus-
like form occurring only very rarely. But last year, so my
host Mr. Jas. Ball informs me, the latter was very
abundant—quite as common as the typical one, and he

508 Mr. G. A. K. Marshall on

caught a long series of them. It seems clear they were
not immigrants from Durban, but what caused their
appearance in such unusual numbers it is difficult to
understand. I may mention that in the last few years
Mr. Ball has caught two £. wahlbergi but has never seen
dominicanus. While there, I saw six females of P. cenea,
two of each of the three forms.”

31. WARNING COLOURS AND MIMICRY (ALMOST WHOLLY
MULLERIAN) IN SouTH AFRICAN COLEOPTERA.
(GAN MEE BE)

[In the groups described below, Coleoptera play a
dominant part, either making up the whole or, except in
the case of the Mutilloid group, acting as models for other
insects. In this one exception the chief interest centres
in the Coleoptera, and therefore the group is included
here. A certain number of mimetic Coleoptera will be
mentioned elsewhere in other groups which have collected
round various types of Hymenopterous models.

In the present section the extraordinary predominance
of Miillerian associations in South African Coleoptera stands
out as the most prominent conclusion.—H. B. P.]

A. Peculiar Warning Patterns and Directive Marks in
Carabidze and Cicindelide. (EK. B. P., G. A. K. M.)

Some of the warning patterns of the large Carabidx of
the genus Anthia are very remarkable and effective, and
their development and relationship in the different species
extremely interesting.

Six illustrative examples are figured on Plate XVII.
In Fig. 21 we see the ancestral appearance, the uniform
black of so many large Carabids, in Anthia massilicata.
Mr. Marshall’s account of the habits of the South African
members of the genus, printed on page 510, shows that
such a beetle is highly conspicuous. It is no doubt an
advantage, however, to gain easily-recognizable distinctive
marks on the black ground of the exposed dorsal surface,
and we find that the species of Anthia do, as a rule, possess
two or more white patches upon some part of this area.
The pair of elongated thoracic white patches, in A. peters:
(Fig. 22), are borne upon the sides of a thorax which is very
like that of massilicata, while in A. thoracica (Fig. 23) this
part of the body is greatly widened and the white patches

The Bionomics of South African Insects. 509

become a broad oval in shape. <A pair of white spots (as
well as a white line towards the outer border of the
posterior half of the elytra) has also arisen on the anterior
part of the elytra in 4. omoplata, var. mellyi (Fig. 26), and
it is of deep interest to note that these spots at a little
distance or when the insect is moving would resemble the
entirely different thoracic spots of thoracica far more closely
than these latter resemble the entirely homologous spots
of petersi. The comparison of Fig. 23 with 22 and then
with 26 will bear out this conclusion. Perhaps one reason
for the development of the remarkable structures in which
the spots are situated in thoracica (they are placed in the
concavities of special lateral outgrowths of the thorax)
may be in order to favour the synaposematic approach to
an arrangement like that of medlyi; for by this means it
is possible for the spots to attain approximately the same
size and shape, and at the same time to retain an interval
between them which corresponds to that obtaining in the
very different position upon the elytra. The concavities
exist however in a much smaller form in A. mawillosa, and
the white markings in them are inconspicuous.

In A. nimrod from West Africa (Fig. 25) we see a further
development of the medlyi pattern in the appearance of
another pair of spots on the posterior region of the elytra,
while in A. sex-guttata, from India (Fig. 24), this appear-
ance is combined with the two spots of thoracica, thus
building up a warning pattern of remarkable simplicity
and effectiveness, being an almost exact negative of the
six of dominoes. The success of the aposeme is much
enhanced by the approximate equality of the shorter
intervals between the spots of each pair, and the longer
ones between the spots of one pair and the corresponding
spots of the next. The great breadth of the thorax
permits this symmetry in one direction without the
development of outgrowths like those of thoracica, while
symmetry in the other direction has been rendered possible
because the middle pair of spots occupies a more posterior
position on the elytra than does the corresponding pair of
nimrod or mellyi. It will be seen by a glance at Figs. 23 to
25 that the addition of the spots of Fig. 23 to the thorax
of Fig. 25 would produce a very inferior warning pattern
as compared with that of sex-guttata (Fig. 24).

Mr. Marshall gives the following account (1902) of the
habits of these formidable beetles :-—

510 Mr, G. A. K. Marshall on

“The warning character of the large spots and stripes
of the Anthias (Plate XVII, figs. 22-26) is well borne out
by the appearance of these insects in their natural haunts.
They are purely terrestrial in their habits, and prefer open,
treeless country, where owing to their large size and
striking coloration they are very conspicuous objects.
When alarmed they adopt a very characteristic warning
attitude, raising themselves high on their legs, walking in
a quick jerky manner and often twisting sharply from side
to side; but, as in the case of other aposematic insects,
when they find these demonstrations are of no avail they
endeavour to make good their escape and are then capable
of running at a very fair pace. Their acid secretion is
very powerful, and causes a strong stinging sensation when
it touches the skin of the face or the more tender parts of
the hands, and as it can be projected to a distance of some
four or five feet, the insect would have to be captured with
considerable caution even by an enemy which might be
aware of its powers. The liquid is always ejected upwards,
and the insects seem capable of controlling its direction
to a limited extent. A very similar warning attitude is
observable in the huge Cicindelids of the genus Mantichora,
at least I have seen it in IZ. herculeana. The habits of
this insect are very similar to those of Anthia, but it cannot
project its protective secretion, which merely exudes when
it is handled; the lquid also is not acid as in Anthia, but
possesses a strong smell.”

Mr. Marshall brings forward the very probable hypothesis
that the posterior white spot or spots of the small and
medium-sized Carabide are directive. The fact that they
form an important element in the Mutilloid appearance of
these Coleoptera is no objection whatever to this hypo-
thesis, which is explained in detail below ; for it is probable
that the posterior white spots of the female Mutillide may
have a similar function, directing attention to the sting.
Mr. Marshall states (1902) that the abdomen is the most
conspicuous part of a Mutillid (see p. 512).

“Among the diurnal Carabidw in South Africa the
frequent occurrence of a conspicuous white spot at the
apex of the elytra is very noticeable, and there is good
ground for believing that this spot is of a directive
character. Unlike such insects as Cantharide, Lycide, etc.,
the Carabide do not appear to possess any general dis-
tastefulness, but depend for protection solely upon their

The Bionomics of South African Insects. 511

power of squirting a strongly acid liquid from behind. In
the case of the largest species, such as Anthia, the great
strength and large quantity of this liquid render it a very
efficient protection; but in the smaller species my experi-
ments and observations lead me to suppose that this is not
the case, but that the utility of the secretion lies rather in
the fact that it enables the msects momentarily to dis-
concert their enemies, and this, owing to their great
activity, gives them an opportunity to escape. Under
these circumstances it is evidently of importance that an
attack from an enemy should be directed to the anal
portion of the body in order to ensure its receiving the
discharge. The anal white patch is especially noticeable
in such genera as Polyhirma (semisuturata, bennett [see
Appendix, pp. 547, 548], notata, rutata, macilenta, etc.)
and Piezia (marshall and mashuna); it also occurs in the
Cicindelid Myrmecoptera polyhirmoides which consorts with
many of the above species. It is probable that the two
white or yellow spots which are found in so many Cara-
bide and Cicindelide (see Plate XVII, figs. 7-11, 14-19)
towards the apex of the elytra have also a similar
significance, that of the Cicindelide being of course
mimetic (of Mutilide and Carabidx).”

B. Mutilloid Coleoptera: Cleridx, Caralide, and Cicindelidx :
Primary and Secondary Synaposematic and Pseudapose-
-matic Associations, (EK. B. P.)

When forwarding the following mimics of Mutillide,
Mr. Marshall also sent a record of habits :—

“ Salisbury, Jan. 8, 1899.—The Mutilloid type is not
uncommon among Carabidex, Eccoptoptera cupricollis being
a beautiful instance of mimicry. But the markings are
most developed among the Cleridx, of which I have no
less than twelve species all exhibiting the Mutilloid
marking more or less distinctly; I expect they will be
found to be parasitic on Mutillide. I have also caught a
little spider which is an excellent Mutilloid mimic.”

This species of Mutilloid spider and another sent later
have been described by the Rev. O. Pickard-Cambridge,
E.R.S., as Prosthesima albomaculata and Titus lugens. (Proce.
Zool. Soc. Lond., 1901, p. 11, figs. 2, 3, Plate V.)

“Salisbury, Feb. 12, 1899.—The Mutilloid group is
interesting though not so satisfactory from a cabinet point

512 Mr. G. A. K. Marshall on

of view, the resemblance being much less marked than in
the field, except perhaps Hecoptoptera which is a splendid
case, and it has caused me to hesitate more than once
before venturing to handle it. At the present time I
would not venture to express an opinion as to the use or
significance of the colours in this group. The Mutdllidx
of course are armed with a powerful sting, which however
they are slow to use, and besides they are very hard; the
red prothorax is by no means conspicuous when they are
running on the ground, the abdomen being the part that
catches the eye, ‘and when hard pressed this is elevated in
the air evidently as a warning. I have noticed that it is
very difficult to distinguish the pattern while the insect is
running, the general impression being merely that of a
black body with white spots. The same applies to the
Cicindelide and Carabidx, which are all fast runners and
most of them very difficult to distinguish inter se in the
field at first sight. The exact resemblance of Graphipterus
antiokanus to Piezia selowsi is marred by the shrinkage in
the former of the pygidium, which bears two white patches.
Atractonota, despite its markings, very much resembles
one of our larger black ants, especially in its manner of
running. A good many other species (especially of Cleride)

might also be included in the group.”

The Mutilloid group of Coleoptera from Salisbury may
be arranged together with their models as follows, the
plan being that adopted in Plate XVII, figs. 1-12 and

14)

MUTILLID#.
Mutilla M. M. M. M. M.
purpurata tettensis cepheus | leucopyga? sycorax horrida
? . a = ers
Fig. 1. Fig. 2 Fig. 3. Fig. 4 Fig. 5. Fig. 6.
(Noy. 1898— (Nov. 1898—
Jan, 1899), Jan. 1899),
CARABIDE. CLERID&.
| Atractonota | Atractonota | Polyhirma ; Polyhirma | Eecoptoptera | Graptoclerus,
mulsanti | mulsanti enigma enigma eupricollis sp.
Fig. 7. Fig. 8 Fig. 9. Fig. 10 Fig. 11. Fig. 12.
| (Nov. 1898—
Jan. 1899). {|
CARABID#. CICINDELID.-£.
2 A ——
Graphipterus Piezia | Polyhirma |Myrmecoptera | Myrmecoptera| Myrmecoptera
antiokanus selousi | bilunata, Boh.) marshalli invieta bilunata
Fig. 14 Fig. 15. Fig. 16. Fig. 17. Fig. 18. Fig. 19.
(Noy. 1898— (Nov. 1898—
Jan. 1899). Jan. 1899). |

The Bionomics of South African Insects. 513

All were captured at Salisbury, in Nov. 1898, with the
exception of those specially noted as taken between Nov.
1898 and Jan. 1899.

Other species are not included in the Plate for want of
space. Thus Piezia marshalli has a single median white
spot, of which half is on one elytron and half on the other,
in place of the posterior well-separated pair of spots on
P. selousi ; and there are other species, outlyimg members
of the group under discussion, in which the Mutilloid
appearance becomes less marked, while in Polyhirima semi-
suturata it fades away altogether, although this intensely
black beetle, with its white dorsal line anteriorly and white
patch posteriorly, seems to be very conspicuous. The
appearance of the group and the relation to the outlying
species suggest a strong and very complex Miillerian
association. The large Caralide of the genus Anthia
are either entirely black or possess a peculiar synapo-
sematic appearance, described on pp. 508-510, and tigured
on Plate XVII, figs. 22 to 26. The sinellles Carabude:
depending upon a less development of the same defence
—the power of discharging a strong acid secretion—
have gained an appearance, due, hike that common in
Anthia, to white markings on a black ground, but arranged
so as to suggest more or less strongly the likeness of a
Mutilla. Ina single species, as Mr. Marshall points out,
the resemblance is extraordinarily exact (Plate XVI
fig. 11). This may be on account ef habits and a mode
of life which render the likeness especially beneficial.
The smallest Carabide, the Atractoncta and Polyhirma
enigma (Plate XVII, figs. 7-10), in shape resemble large
ants, and Mr. Marshall states above that the movements of
the former aid in producing this effect. On the other
hand, their white spots appear to be certainly Mutilloid or
perhaps rather to resemble the Mutilloid white spots of
the other unant-like Carabidy. Nothing is more char-
acteristic of a Miillerian (synaposematic) group than the
complexity of likeness which is thus revealed, and yet in
the light of the great hypothesis which we owe to Fritz
Miiller it is not difficult to understand the general principles
which account for its existence.

The Caralidx are a powerful, specially defended group,
and it is of advantage to be recognized as belonging to
the group, even though it is no doubt-of still greater
advantage to be mistaken, as may happen at a distance,

TRANS. ENT. SOC. LOND. 1902.—PART IIL. (NOV.) 34

514 Mr. G. A. K. Marshall on

or on a superficial view, or during rapid movement, for
the still more formidable Mutillide and ants. Hence,
although the smaller species of this group resemble the
latter, and the larger the former, markings are neverthe-
less retained which serve to connect these Coleoptera
together, and enable the experience gained in an attack
on one of them to be of service im preventing the waste of
life in many other species. For the same reason we can
understand another curious inter-relationship, viz. that
there is a superficial resemblance between different genera
of Carabide and between Cicindelide and Carabidae.
Running through, but not concealing the resemblance to
the Hymenoptera, is a resemblance between the Cole-
opterous mimics themselves. ‘Thus there is the likeness
described by Mr. Marshall on p. 512, between Graphipterus
antiokanus and Piezia selousi (compare Plate XVII, figs.
14 and 15), while the remarkable likeness of the Cicin-
delidx to the Carabide will be at once appreciated when
Figs. 17, 18 and 19 are compared with 15 and 16. The
same explanation is doubtless valid. The Cicindelide are
less powerful than the Carabidx, but many of them emit
a peculiar scent, and the genus T'ricondyla is closely
mimicked in Borneo by the Locustid Condylodera tricondy-
loides. We can well understand that it is to the advan-
tage of Cicindelide to be one with another and more
powerful set of Coleoptera, even though it may be a
further and probably greater advantage to resemble the
Mutillide, a resemblance which is also involved in the
appearance they have gained. If this explanation be
sound we shall expect also to find examples of the same
kind of likeness between Cicindelide and Carabidx which
do not resemble these Hymenoptera. Plate XVII shows
that this is undoubtedly a fact, for the all-black Polyhirma
boucardt (Fig. 13) is seen to be strongly resembled by a
Cicindelid, Myrmecoptera polyhirmoides, var. mashuna
(Fig. 20), belonging to the same genus as the three white-
marked species figured in the same Plate.

The Cleride everywhere tend to resemble MJutillide,
but they also resemble Cantharidx, and more rarely
Tycide and even Coccinellide, while one genus is beauti-
fully mimicked by a Longicorn in Borneo. The first-
mentioned likeness is probably a marked example of
synaposematic colouring, and I should expect that the
character of the original structure and warning pattern,

The Bionomics of South African Insects. 515

rather than the parasitism which Mr. Marshall suggests,
rendered resemblance to Jfutillide above all other
specially-defended insects particularly feasible for these
Coleoptera. A good example of such mimicry is seen in
Plate XVII, fig. 12, representing Graptoclerus, sp.

I believe that the principles discovered by F. A. Dixey,
which are here employed to explain the curiously complex
inter-relationships of Mr. Marshall’s Mutilloid group, will
hereafter be found to have an important bearing upon
many superficial resemblances of mimicry and common
warning colours in all countries, and in many orders of
insects and probably other animals.

The double (or treble, etc.) resemblances which may be
perceived one underlying the other in the appearance of
a single form may be conveniently spoken of as Primary,
Secondary, Tertiary, ete., Common Warning Colours (Proto-,
Deutero-, Tritosynaposematic Resemblance), or as Prim-
ary, Secondary, Tertiary, ete., Mimicry (Proto-, Deutero-,
Tritopseudaposematic Resemblance). Thus the resem-
blance of the Atractonota to an ant, or of MWyrmecoptera to
a Mutillid is Protosynaposematic, while their resemblance
to the Carabid genera Polyhirma, Piezia, ete., is deutero-
synaposematic. On the other hand, the resemblance of
the black Myrmecoptera to Polyhirma boucardi is proto-
synaposematic, or rather, synaposematic, since there
appears to be no other underlying or overlying resemblance
in this case.

Many examples of this kind will be found in the section
on Mimicry in Lepidoptera; see especially pp. 470, 471,
485-487.

I must here again refer to Dr. F. A. Dixey’s important
memotrs, alladed to in greater detail on p. 502, as contain-
ing the first account and interpretation of the extremely
complex inter-relationships which may exist in Miillerian
associations. The principles which he laid down are here
found to supply the interpretation of many puzzling and
subtle relationships, not only among Lepidoptera, but
also in other Orders in which the phenomena of mimicry,
warning colours, ete., have been far less fully investigated.

C. Lycidx as Models for other Coleoptera and Insects of
many Orders, (G. A. K. M.)

[The splendid and complex convergent group, repre-
sented on Plate XVIII, figs, 1-42, is by far the most,

516 Mr. G. A. K. Marshall on

complete illustration hitherto known of the power of
mimicry to attract forms of all kinds irrespective of
affinity. It is of the deepest interest to observe that the
association is almost entirely Miillerian (synaposematic).
The following passages are quoted from letters written by
Mr. Marshall about the time when the first consignments
were sent; but the group became very much larger when
the additions of later consignments were included.—
BeBe Pa)

Salisbury, Jan. 8,1899.—The two most prominent types
of colour among mimetic Coleoptera are what I call. the
Lycoid and Mutilloid types. Of the former I have put
aside for you a series comprising fifteen species, including
several species of Lycus, three species of Longicorns, a
Reduviid bug, a fly, a wasp, an Arctiid moth, a Mylabris, a
Hyperacantha, ete., and I shall be able to add more im all
probability.

The Lycoid Arctiid moth isa day-flyer, but the deceptive
resemblance is not good on the wing, being best shown
when the moth sits, as is its wont, on the ends of grass-
stems, etc., after the manner of a Lycus; it is perhaps
even more like Prionocerus dimidiatus, a Lycoid unpalat-
able Malacoderm which has a similar habit.

Salisbury, Feb. 12, 1899.—The six species of Lycus, the
Prionocerus, Diacantha, Zonitis, Mylabris, and Eletica, I
proved by experiment to be distasteful to baboons and a
kestrel. The Zygenid I presume to be so likewise, as it
emits a strong smell; the Zelephorus will also probably
prove to be unpalatable. The four species of wasps have
all got very eftective stings; thus the only unprotected
insects are the fly, which is an admirable mimic of one of
the wasps, and the three Longicorns, though I am not quite.
certain about Philagethes. As to the Reduvius I do not
know what to say, there are certainly some very remarkable
cases of mimicry in this family. * * * *
In flight the Zygenid [Newrosymploca, Fig. 52] is aided by
its very brilliant hind-wings, and the Hymenoptera have a
flight very different from, and far swifter than, that of
Lycus.

[A complete list of the species arranged in their respec-
tive families is given below. Large as the group is it
could certainly be made much larger, especially if the
whole of South Africa were put under contribution. Thus
an obvious addition to the Longicorn mimics is Dyenmo-

The Bionomics of South African Insects.

517

nus apicalis, which Mr. Marshall has not yet taken in

Mashonaland.—E. B. P.]

Group with Lycorp MARKINGS (Plate X VIII, figs. 1-52).

/ Copride

Melyridx
CoLEOPrERA (
Phytophaga

Lagriide

Cantharidx

Longicornia

Braconidx

HymeEnoprera / Pompilide

Crabronide
Larride
HBumenidex
Pyrrhocoridx

Layxidz
HEMIPTERA IY:

Reduviide

Arctiide

SS fe

LEPIDOPTE
SP OEARS on Zygenidx

DIPTERA Asilidx

~

Malacoderimata

Aphodius holubi (fig. 18).
Lycus (Merolycus) rostratus (figs. 1, 2).
Lycus (Acantholycus) constrictus (fig. 3).
Lycus ampliatus (figs. 4, 5).
Lycus (Lopholycus) zonatus (fig. 6).
Lycus (Lopholyeus) haagi (figs. 7, 8).
Lyeus (Chlamidolycus) subtrabeatus
(figs. 9, 10).
Lycocerus mimicus (figs. 11, 12).
Prionocerus dimidiatus (fig. 13).
{ Peecilomorpha fasciaticollis (fig. 14) ;
Diacantha conifera (figs. 16, 17).
| Peploptera zambesiana (fig. 15).
Eutrapela sp. (fig. 19).
fEletica rufa (figs. 20, 21); Mylabris
\_ palliata (fig. 22) ; Zonitis sp. (fig. 23).
Amphidesmus analis (fig. 25); Phila-
| gathes ltus (figs. a 27).
(fig. 28); N. similis (fig.

Nitocris ee
N. *nigricornis (fig. 29).

| 30) ;
Blepisanis haroldi (fig. 24).

Phanomeris sp. (figs. 44, 45) ; Iphiau-
( lax bicolor (fig. 46) ; Bracon Iuetuo-
ae ae (fig. 47) ; Bracon ?luctuosus (fig.

Pomnilus are (fig. 36); P. capensis
(fig. 37); P. diversus (fig. 38); P.
vindex (fe. 39).

Cerceris ee var. (fig. 40),
Notogonia croesus (fig. 41).
fRhynehium radiale (fig. 42) ;

L_ (fig. 43).

Serinetha mutilata Wee 33).
p Lygieus fureatus (fig. 34) ; Oncopeltus
\ famelicus, var. jucundus (fig. 35).

y Vitumnus miniatus (fig. 82); V. cin-

\ nabarinus (fig. 31).

Ilema elegans (fig. 50).
sf Neur osymploca ochreipennis (fig.
\_ Zygenid (genus *) (fig. 51).

Xiphocerus cruciger (fig. 49).

R. rubens

52) ;

In the above group the beetles of the genus Lycus un-
doubtedly constitute the dominant fete towards which
the other insects have converged with more or less exact-

Ness.

The members of this genus are very numerous

throughout S.E. Africa both in species and in individuals.

They are

most conspicuous insects, and* the majority of

them occur on flowers, though a few species are more often

518 Mr. G. A. K. Marshall on

to be found on the heads of grasses. In habits they are
very sluggish and have a slow heavy flight; when handled

they usually emit from their limbs a very ‘strong-smelling
white liquid, and they frequently feign death. The
Melynid Prionocerus has somewhat similar characteristics,
but is found only feeding on grass seeds. Among the
Phytophaga it is possible that the Peecilomorpha is a
Batesian mimic, for the species of this genus are nearly all
more or less scarce, and moreover exhibit marked mimetic
tendencies in very different directions. The Diacantha, to
which D. dimidiata might also have been added, is a very
plentiful insect with a nauseous smell; it causes much
damage to garden plants such as cucumbers and pumpkins.
The Peploptera i is likewise abundant, frequenting acacias.
All the species of Lagriadz and Cantharidze mentioned
are flower-feeders, and are er protected by distasteful
qualities. ‘I'he significance of the colouring in the Longi-
corns is still a matter of doubt, as I have obtained no
experimental evidence with regard to them; probably
most of them are pseudaposematie, but Prilagathes may
be a Miillerian mimic. Of the Hymenoptera the quick-
flying Notogonia is the only scarce species with us; the
other Aculeates all visit flowers more or less commonly.
The Braconids are slow and very conspicuous fliers, being
evidently protected by their very strong smell. The
species of Hemiptera also occur commonly on low plants
and bushes; the position of the Reduviids is not quite
certain, as experimental proofs are lacking. The Asilid fly
appears to be a very rare species and is doubtless a Batesian
mimic.

D. Miillerian (Synaposematic) Groups in South African
Coleoptera. (G. A. K. M.)

a. Cuntharid Group (Represented on Plate XIX).

Mylabris dicincta (fig. 1); M. tettensis
(figs. 2-3); M. tricolor (fig. 4); M.
oculata (figs. 5, 6, 13); M. holose-

Cantharide ; ‘
ricea ae 9); Actenodia chrysomelina

x pet (figs. 7, 8, 11); Decatoma lunata
COLEOPTERA (figs. 10, 12).
oy matura bifasciata (fig. 14); Cerople-
Longicorinia | sis caffer (fig. 15); Anubis mellyi
(fig. 16).
Phytophaga Gites wahlberai (fig. 20).

HEMIPTERA Lygevide Oncopeltus famelicus (fig. 17).

The Bionomics of South African Insects, 519

The same species is sometimes repeated two or even
three times in the Plate in order to show common varia-
tions in the pattern.

All the species of Cantharidie in this group are abundant
in Salisbury, some of them oceurring in such numbers as
to form a serious pest in the local flower- gardens. All
those mentioned feed on flowers exclusively, except JZ.
holosericea which seems to be more attached to grasses.
In every case they emit a quantity of vesicating yellow
juice from the antenne and joints of the legs when
handled, and also eject a liquid from their mouths; their
flight ig heavy and noisy, and they are most conspicuous
insects in every way, Ceroplesis caffer 1s probably the
commonest Longicorn in South Africa, attacking dead
wood of almost any description, but particularly frequent-
ing acacias. It has a slow conspicuous flight, and has been
proved by experiment to possess distasteful qualities,
though I have not noticed that it gives off any smell.

The Longicorn Cymatura bifasciata is specially attached
to a species of Lantana, which grows to a fair-sized bush,
and the insects occasionally may be found on it in some
numbers. It is certainly a Miillerian mimic, as it has a
strong smell, is very sluggish, and feigns death persistently
when captured. It was refused with evident dislike
by baboons. Anubis mellyi does not occur at Salisbury,
but is fairly common at Umtali, further east, where it
frequents the flowers of low plants. It is a brightly-
coloured insect, and emits the strong characteristic smell
of the diurnal Cerambycidee.

B. Intermediate Group connecting the Cantharid and
Coccinelloid Groups (Represented on Plate XIX).

Cantharidx Actenodia chrysomelina (figs. 7, 8, 11),

Melitonoma sp. nov. (fig. 18) ; M. trun-

catifrons (fig. 19); M. litigiosa (fig.

21); M. epistomalis (fig. 23); Anti-

CoLEorrTERA \ Phytophaga pus rufus (figs. 22, 25) ; Clythra lac-

ordairei (fig. 24) ; Aulacophora festiva

(fig. 26); Cryptocephalus 5-plagiatus

(fig. 27) ; C. varioplagiatus (fig. 28) ;
Syagrus marshalli (fig. 29).

This group merges very gradually into the Mylabroid

group on one hand, especially in those species where the
IV Pp

rows of spots coalesce into transverse bands, and at the

520 Mr. G. A. K. Marshall on

other end of the series it converges to the Coccinelloid type,
particularly in the Cryptocephali, The association is prob-
ably of a purely Miillerian character so far as the species
mentioned are concerned. They all have very similar
habits, occurring on low plants and flowers, and making no
attempt at concealment. A. festiva is much more plen-
tiful than the others, and is probably the dominant member
of the group.

y. Coccinelloid Group (Represented on Plate XIX).

fEpilachna dregei (figs. 40, 41); Chilo-

COLEOPTERA Coccinellidx 2
hs : \. menes lunata (fig. 42).
HEMIPTERA Pentatomidx Steganocerus multipunctatus (fig. 39).

I have frequently found all these three species occurring
together on the flowers of a Lantana, and as they are all
common, conspicuous, and strong-smelling insects, the
group is undoubtedly synaposematic.

6. Group of small pale yellow and red Phytophaga with
their Melyrid and Curculionid Mimics, (EK. B. P.)

With respect to a series of nine small brightly-coloured
Coleoptera of about the same size (with the exception of
Vrodactylus, ? sp. 2, which is conspicuously smaller than any
of the others), Mr. Marshall wrote that he should be unwill-
ing to hazard an opinion. It appears tolerably clear how-
ever that they form a beautiful Miillerian group, including
perhaps a single Batesian mimic. All were captured at
Salisbury, and, with the two exceptions noted below, in
January 1899. The species are arranged below as they
are on Plate XIX, where each is represented twice the
natural size.

PHYTOPHAGA.

Platyxantha Gynandrophthalma Monolepta
bicincta posticalis vincta
fig. 30. [or closely allied, M. [or closely allied, M.
Jacoby | Jacoby]
(Feb. 1899) fig. 32.
fig. 31.
Crioceris Paralepta Asbecesta
coronata, Baly ornata ornata
=balyi, Harold) fig. 34, fig. 35.

fig. 33.

The Bionomics of South African Insects. bal

MELyYRID®. CURCULIONID A.

Urodactylus, sp. Urodactylus, sp. 9 Apoderus

fig. 36. fig. 37. gentilis
(Nov. 1898)

fig. 38.

The group consists of pale yellow insects with bright
red anterior section, made up by thorax and head, of which
the eyes only are black in some of the species, although in
others black markings extend over a variable proportion
of the cephalic surface. The elytra are crossed trans-
versely by a black band at the base, and another at the
junction of the third and posterior fourth of their length.
In some species the anterior band, in others the posterior
is reduced to two more or less widely-separated spots or
patches. In the species of Crrioceris, the anterior band is
represented by four small black spots, two on each elytron.
In the Urodactyli, the posterior band is represented by a
semi-circular black mark with the concavity directed
posteriorly on each elytron.

In spite of the variation in detail the species would
produce the same effect at a little distance, and there can
be no doubt about the interpretation of the whole as a
synaposematic combination with the Curculio as a doubt-
ful pseudaposematic member. Mr. Marshall states that
there aré many other species of about the same _ size
which adopt the same warning pattern, including at least
four additional species of MJelyride.

Certain members of this group are very abundant and
conspicuous in the spring months upon the Mosasa-tree
(Brachystegia sp.).

E. Comparison between certain Coleopterous Groups in Borneo
and South Africa, with respect to Mimicry, Common

Warning Colowrs, ete. (G. A. K. M.)

[The following interesting comparison between the
phenomena of mimicry and common warning colours in
certain Coleopterous groups in Borneo and South Africa
respectively, was made upon the receipt of a set of photo-
graphs of the material of Mr. Shelford’s paper now being
published by the Zoological Society.—E. B. P.]

Salisbury, Jan. 11, 1901.—I should be interested to
know whether Shelford has proved the Anthribidx to be

529 Mr. G. A. K. Marshall on

distasteful; from my knowledge of our fairly numerous
South African species I should be much inclined to doubt
it, as their colouring appears to be, without exception,
protective, generally resembling bark or lichen; and
although there are somewhat similarly-coloured Longicorns
which frequent such surroundings, I should certainly class
their colours as syneryptic. A somewhat similar criticism
suggests itself with regard to the Brenthidex, although I
feel difttident in stating it, as the family is so very limited
here. I know of only four species in Salisbury, all of
which are sub-cortical and nocturnal in their habits, of
comparatively rare occurrence, and of dull colouring. On
the other hand, I have observed that the great majority of
our smaller Lamiids adopt the forward position of the
antenne, which I have always regarded as_procryptic,
as there can be no doubt that it renders them much less
conspicuous than if the antennz were held out at an
angle to the twig on which the insect sits. The procryptic
nature of the position is well illustrated in the small and
very elongate Longicorns Hyllisia and Hippopsicon, of
which we have a few species, all of which frequent grass-
stems in marshy places; they also have the elytra bifur-
cated, and this seems to be a common occurrence in all
very elongate beetles. The Endomychid groups are
very interesting. Unfortunately this family is extremely
poorly represented here—only some three or four species,
though the Erotylide are fairly numerous. In this latter
the pattern with four yellow or reddish blotches on a
black ground occurs also with us, and the insects are
probably distasteful, judging by the extremely pungent
smell emitted by the large Hncaustes. Curiously enough,
just after getting your photographs I found under bark a
large Endomychid (new to me) of this pattern, and with
it occurred an admirable mimic, a Carabid Thyreopterus
flavosignatus (Dej.). There is another Carabid <Arsinoé
Sraterna (Pér.), also sub-cortical, which mimics it closely,
but unfortunately I have no specimens now; I caught only
two here six years ago.

EF. Note on Rhynchophora with Procr ae Colouring as
Models for Minmicry. (i. B. P.)

Dr. A. R. Wallace has always thought that the extreme
hardness of the mimicked Curculionide and Anthribide

The Bionomics of South African Insects. 523

is the character which protects them (“Essays on Natural
Selection,” 1875, p. 94). In answer to a letter in which
I drew his attention to Mr. Marshall’s record of a large
Curculio found in the crop of a guinea-fowl (see p. 350), he
wrote, Feb. 5, 1901, “The large Malayan Anthribide are
intensely hard. The guinea-fowl proves nothing, as these
beetles are almost all arboreal, and their chief enemies are
smaller birds. Their protective colours may save them
from the larger insectivorous birds, their hardness from
the smaller.’ The mimicry of Malayan Curculionide,
Anthribide, and Brenthide by Longicorns cannot be
doubted. The cases are too numerous and the details of
the resemblance too precise to admit of any other explana-
tion. In South Africa, on the other hand, Mr. Marshall
shows that only the first group is mimicked, and of this
he has sent me a very beautiful example. Experiments
are greatly wanted, especially in Borneo, where all three
groups abound. In addition to their hardness Mr. Shel-
ford shows that the larger Bornean Curculios are defended
by their great strength; they can even cause intense pain
to man by elasping the fingers with their legs and digging
the proboscis into the flesh. Such defences as hardness
and strength depend for their success on the size of
enemies; for even hardness could not avail against an
enemy large enough to swallow the beetle whole, so that
it could be ground down in the gizzard, or the interior
slowly extracted by digestive fluids gaining access by the
joints and other apertures. Defence by a sting, a nauseous
taste or smell, or unwholesome qualities, is effective against
enemies of all sizes and all degrees of strength, although fail-
ing against occasional specially-adapted foes. It is possible
that these considerations may enable us to understand why
it is that certain Rhynchophora are remarkable among Cole-
optera for combining a cryptic colouring with sufficient
immunity to render them feasible models for mimicry.
The ordimary methods of active defence among verte-
brates—the power of biting or pecking, of kicking or
tearing with hoofs or claws*-together with the passive
resistance of a spiny or hard external covering, are
almost invariably associated with cryptic colourmg and
modes of life favouring concealment. The probable ex-
planation is that all such methods of defence must fail before
large and important classes of still stronger enemies or
foes with cunning sufficient to circumvent the passive

524 Mr. G. A. K. Marshall on

defence. Such enemies are numerous enough and
deadly enough to make the increased danger of a con-
spicuous appearance far greater than any advantage
gained by the warning off of smaller and weaker animals.
Such a defence as that of the skunk, on the other hand,
appeals to enemies of many classes, and is quite inde-
pendent of strength or size. Here and there special
animals, probably powerful birds with deficient sense of
smell, can endure the defensive odour, and to these the
skunk would be an easy prey; but on the whole the
increased danger of a conspicuous appearance and slow
movements is far more than compensated by the warning
off of an immense number of would-be enemies. Cases
like that of the skunk are very common among insects,
while those of active defence are very rare. Even the
passive defence of a spiny or hairy covering is very
different from that ordinarily adopted among vertebrates,
because in the insect the hairs and spines are themselves
a cause of unpalatability, and often of intense irritation, so
that they tend to be associated with an aposematic
appearance. It is, however, probable that the intensely
hard Rhynchophora with a cryptic appearance, and especi-
ally the largest and most powerful Curculionidae, are
strictly comparable with the large number of vertebrates
which also unite the methods of concealment with very
efficient modes of active or passive resistance. Certain of
the largest Curculionide possessing red marks on a black
ground appear to possess an aposematic appearance, and
these may be distasteful, although the conspicuous appear-
ance may only indicate an excessive hardness and thick-
ness of chitin which, coupled with the great size, may be
a most efficient defence against a majority of enemies.
Mr. Marshall tells me that the largest South African
Curculios of the genus Brachycerus, such as 6. apterus,
are purely terrestrial, move slowly, and freely expose
themselves, like our European distasteful species of
Phytophagous Zimarcha. Under these circumstances the
intense black ground-colour and red spots of B. apterus
must render ‘it remarkably conspicnous, and it would be
of great interest to ascertain, by a number of experiments
on many insect-eaters, whether so pronounced an aposeme
may indicate hardness alone or hardness combined with
some other special protection.

The small size of the Brenthide renders it improbable

The Bionomics of South African Insects, 525

that hardness alone can be sufficient protection to the
mimicked species, such as the Bornean Diurus furcillatus,
and we are led to suspect the existence of unpalatability.
In the cabinet the specimens seem to be markedly cryptic,
but Mr. Shelford assures me that they are very commonly
found on flowers, where their dull dark colours would
be most conspicuous. Above all things experiments with
insect-eating animals are greatly needed to throw light on
this most puzzling and exceptional occurrence, viz. the
existence of large numbers of models for mimicry among
Rhynchophora with an apparent, and certainly in many
cases an actual cryptic appearance.

32. ComMMON WARNING CoLoURS IN SouTH AFRICAN
HYMENOPTERA AND THE MIMICRY OF THEM BY
INSECTS OF OTHER ORDERS. (G. A. K. M.)

A. Group with Black Bodies and Dark Blue Wings, chiefly
Fossores.

Apide f Xylocopa hottentota (fig. 18); X. cari-
| (Pl XXI) \ anata (fig. 19).
Eumenide { Eumenes tinctor (figs. 14, 15); E. dys-
(Pl. XXII) -\ chera (figs. 16, 17).
Sphex bohemani (figs. 1, 2); S. cyani-
ventris (fig. 3).
S. pelopeiformis (figs. 4, 5); S. xantho-
cerus (fig. 6).
Sphegidx S. umbrosus (fig. 7) ; Sceliphron chaly-
GPL, 210) beum (fig. 8).
Tachytes natalensis (figs. 9, 10); Ammo-
phila ludovicus (figs. 11, 12).
A. beniniensis (fig. 13).
( Salius atropos (fig. 14); S. vindex (fig.
il).
| S. dedjax (fig. 16); S. regina (fig. 17);
) S. obscurus (fig. 18).
Pompilus sepulchralis (fig. 19); P. frus-
tratus (fig. 20).
Elis lachesis (fig. 3); E. fasciatipennis
(figs. 4, 5).
Scolia alaris (figs. 6, 7, 8); S. fraterna
7 (figs. 9, 10).
S. cyanea (figs. 11, 12); S. affinis (fig.
13).
Tiphiide Tiphia rugosa (Pl. XX, fig. 2).
Mutillides Mutilla atropos (Pl. XX, fig. 1).

HYMENOPTERA

Pompilidex.
(Oral C0)

Scoliadx
(Pl. XX)

4

526 Mr. G. A. K. Marshall on

Epicauta pectoralis (fig. 23); E. sub-
coriacea (fig, 24).

ccelestina (fig. 25); Lytta meesta
(fig. 27).
Asilidx Laparus sp. (fig. 20); L. ? tabidus (fig.
(Pl. X XI) Pl)
Bombylidx Exoprosopa 1mbrosa a XX, fig. 22).
Tabanidex Tabanus ? biguttatus (Pl. XX, 24).
Tac vinidx Orectocera (Paraphania) ae (BE

XX, figs. 21, 23).

COLEOPTERA

(PI. XX1) Cantharidex E.

DIPTERA

Cetoniidex: Trymodera aterrima (fig. 22).
f
L

HEMIPTERA ; Ferber ase Reduvius sp. (tig. 26) ; Harpactor tristis
(PI. XX) re; Gee 7):
LEPIDOPTERA Zygenidx Tascia homochroa (Pl. XX, fig. 25).

Among the Mashonaland Hymenoptera the most
dominant and conspicuous group is certainly that which
comprises the species having a generally black coloration
with more or less metallic purple wings; for not only is
the number of component species a large one, but the
individuals are likewise very numerous, since the group
contains many of our commonest large Aculeates, such as
Humenes tinctor, Salius vindex, Scolia cyanea, ete. All of
the species of this large group, which are figured in Pilates
XX and XXI, commonly visit flowers, and owing to the
numbers in which they often occur, and their close inter-
resemblance, it 1s very difficult for one who is not an expert
to distinguish the ee without examining them in the
net. The Scolias are the heaviest and slowest. flying species
of the group, though e largest Pompilids run them close
in this respect, being very conspicuous both on the wing and
when settled, and it seems probable that these two types
constitute the nucleus of the mimetic group towards which
the other genera have converged. Moreover, several of
the large species of Salius (Mygnimia) advertise their
offensive qualities by their loud, rattling flight, which can
be heard at a considerable distance. This is specially
noticeable in S. vindex, which sounds as though it were
going by clockwork, the noise often attracting one’s atten-
tion when the insect is flying high above the trees.
Among the other species the Hwmenes also fly somewhat
leisurely, and are not readily disturbed when feeding.
But the species of Sphex and Tachytes are very active,
restless insects with a swift, dodging flight. Of the Cole-
optera coming into this group, Trymoden ‘a aterrima is the
only flower- feeding species, occurring chiefly on Protea, the
flowers of which are much frequented by Hymenoptera.

The Bionomics of South African Insects. 527

Lytta moesta and all the species of Lpicauta are purely
terrestrial in their habits, all beimg very common insects
and evidently protected by their vesicating properties.
The blue-winged Pompilids when running on the ground
bear a distinct general resemblance to these insects. Of
the five species of dark-winged flies, Hxoprosopa wmbrosa
is the only one which visits flowers, and this has a rapid
tight. The two ZLapari and Orectocera (Paraphania)
diabolus are to be found settling conspicuously on low
plants, and have a comparatively slow flight. Zabanws
biguttatus is a rapid flier, with the usual habits of the
genus. The Zygzenid moth, Tascia homochroa, is a very
common species ; it 1s occasionally seen on flowers, but
is more frequently observed settling in a conspicuous
manner on shrubs and plants. It is very sluggish in
habits and emits a strong-smelling liquid when crushed.

B. Mimicry of Diploptera by Fossores and Insects of other
Orders. (H. B. P.)

The dominant members of the group described above
are evidently the Fossores, the Diploptera and Antho-
phila being represented by only two species each.

It is therefore somewhat surprising to note that in one
of the Scolias (Figs. 6, 7, 8, Plate XX), evident mimicry of
Diploptera has been brought about in a very interesting
way. The outer parts of both wings, except the costal
area of the fore-wing, have become extremely transparent
and invisible, possessing a peculiarly illusive quality pro-
bably due to approximation of the refractive index to that
of air. The effect is that both in flight (Figs. 6, 8) and
at rest (Fig. 7) the resemblance to the much narrower
wings of Diploptera is most striking. The resting resem-
blance is pecuharly interesting, for the dark areas of the
wings exactly resemble the narrow longitudinally-folded
organs of the Diploptera, as may be seen by comparing
Fig. 7 on Plate XX with 17 on Plate XXI, or 11 and 13
on Plate XXIJ, while the transparent portions of the Scolia,
although in two layers (compare Figs. 1, 4, 10, ete., on
Plate XX), are not seen. The transparency is well shown
in Fig. 8, Plate XX, where the dark apex of the wing of
Scolia fraterna was made to underlie the transparent part of
that of Scolia alaris, when the photograph was taken for
the preparation of the plate. The underlying wing loses

528 Mr. G. A. K. Marshall on

neither its sharpness of outline nor, in a print prepared
from the negative, any of the detail of the surface. A
tendency in the same direction is manifest in Elis fasci-
atipennis (Fig. 5), but it is hardly noticeable in the repre-
sentation of the resting position in Fig. 4. Mr. Marshall
informs me that the Diplopterous appearance of Scolia
alaris is very marked during life, both at rest and in
flight.

‘It is interesting to inquire for the possible reason of
this mimicry. Fabre (“Insect Life,” translation, London,
1901, pp. 99, 100) states that the Fossores, using their
sting for hunting and providing food for the larvee, are
much less prone to employ it in defence than the
Diploptera, in which it has this latter meaning alone, and
not only so, but when used it is far less painful in the
former than the latter. The Pompilidx, he considers,
produce the most painful effect, but far less than those
which follow from the sting of a bee. On the other hand,
my assistant, Mr. A. H. Hamm, who has had great experi-
ence of our British Aculeates, and always takes them out
of the net with his fingers, thus gaining very exceptional
knowledge of their relative powers in this respect, does
not altogether share Fabre’s opinion. He states that
while the common wasp, and of course the hornet, for his
experience includes even this insect, produce more pain
than any other British Aculeate (I leave the hive-bee out
of account), many of the Fossores produce more pain and
use their sting more readily than other Aculeates. At the
same time Mr. Hamm’s treatment is one that the mildest
Aculeate may be expected to resent if its sting can pierce the
skin of the fingers, and the question is rather whether the
Diploptera are “not recognized as more formidable than the
Fossores by the natural enemies of insects. It is very
probable that this is the case, the combined attack made
upon enemies incautiously disturbing a society being
one element in producing an increased respect for single
individuals of the same kind or with the same general
appearance.

When an illustration of Scolia alaris was shown to the
Entomological Society on March 5, 1902, and this interpret-
ation suggested, Colonel Yerbury and Mr. Verrall pointed
out, at the close of the meeting, that similar opacity of the
costal area of the wing and transparency in other parts was
characteristic of many Diptera mimetic of Hymenoptera,

a OOOO ee

The Bionomics of South African Insects. 529
and might have a similar meaning. Mr. A. J. Chitty
also suggested that the same interpretation applies to the
Sesiid moth shown on Plate XXII, figs. 15,16. In this
latter case the shape of the opaque area would present
during flight a very close approximation to the outline of
the narrow fore-wing and small hind-wing of the Diploptera,
many of which are figured on the same plate as the Sesiid
(compare Figs. 7, 8, 10, 12, etc., with Fig. 16).

It seems probable that the case of Scolia alaris will
throw much light on the manner in which the mimicry
of Aculeates is brought about in insects of many orders.

C. Group with Black Bodies and Yellow Tails, chiefly
Diploptera (Represented on Plate XXII).

HYMENOPTERA. LEPIDOPTERA.

Scoliadx.

Humenidx.

Sesiadex.

Scolia erythropyga (figs. Rhynchium synagro- Trochilium sp. (figs.
1, 2). ides (fig. 6). 15, 16).
Pompilide. Synagris abyssinica
Salius tamisieri (figs. 3,4). (fig. 7). Drrrera,
5, spectrum (fig. 5). Synagris mirabilis (fig. Bombylius sp. nov.
8). (fig. 17).
Synagris emarginata Silvius pertusus (fig.

(figs. 9, 13).
Synagris analis (figs.
1@)> alal)),
Synagris xanthura (fig.
12

18).

Eumenes
(fig. 14).

dyschera

All the Hymenoptera in this group are common insects
occurring in some numbers on suitable flowers. Their
metallic blue wings and yellow tails make them very notice-
able objects, and their flight is leisurely. Of the Zrochiliwm
only two specimens were taken—both of them on the wing,
when the resemblance to a large Synagris was most remark-
able, being much heightened by the transparent edging of
the hind-wing, which is thus made to appear of about the
same size and shape as that of the wasp. Of the two
Diptera, one, the Bombylius, is very common, especially
about flowers; the other, Si/viws pertusus, is apparently
scarce. There is not sufficient evidence to judge on their
true relation to the group.

[The conspicuous yellow apex of the abdomen probably
acts as an easily seen directive mark indicating the sting.
Compare pages 510 to 512 —E. B. P.]

TRANS. ENT. SOC. LOND. 1902.—PART III. (NOV.) 35

530 Mr. G. A. K. Marshall on

D. Group with Dark Bodies, Central White Patch and
Red-Brown Tails : Megachile the Models (Represented
on Plate X XI).

HYMENOPTERA. DIPTERA.

Megachile chrysorrheea (fig. 29). Laphria nr. flavipes (fig. 31).
nasalis (fig. 28).
Coelioxys pusilla (fig. 30).

The Megachile bees mentioned above are both common,
not only visiting flowers, but also, during the drier months,
occurring in some numbers on damp sand or mud near
water. The parasitic Caliorys may be seen with them in
both stations, but is a much scarcer insect. The Asilid is
also an uncommon species, having the usual habits of its
congeners and settling about on low plants.

E. Group with Black Thorax and Yellow Abdomen :
all Hymenoptera (Represented on Plate XXIII).
Tenthredinide Athalia bicolor (fig. 11).

Chalcididx Chalcis bicolor (fig. 10).
Scoliade Elis aureola (fig. 1).

Grabroniies Jeepers fuscipennis (fig. 2); P. diadema (figs. 3, 4);
P. bucephalus (fig. 5).

Humenide Rhynchium radiale (fig. 6); R. rubens (figs. 7, 8).

Andrenidz Sphecodes rufiventris (fig. 9).

This is a beautifully compact and uniform little group,
and is specially interesting owing to its comprising species
of no less than six families of Hymenoptera, which exhibit
great differences in their food and general habits—especi-
ally in their earlier stages. Yet the imagines may be
frequently observed all flying together about the same
patch of flowers, and the uniformity of their coloration is
then very striking. All the species are common and
efficiently protected, so that the association is probably
Miillerian.

F. Group with Black Bodies, Blue Wings, and Yellow
or Ted Thorax (Represented on Plate XXIII).

Xylocopa lateritia (fig. 12); X. flavo-
rufa (fig. 13).
Xylocopa olivacea (fig. 16) ; X. modesta
Apidx (fig. Oy
Podalirius aeraénsis (fig. 14); Antho-
phora ?basalis (fig. 15).
Scoliadx Elis celebs (fig. 18).
LEPIDOPTERA Sesiadee Melittia sp. (fig. 19).

HYMENOPTERA «

The Bionomics of South African Insects. 531

This group forms another interesting illustration of the
way in which the larger mimetic groups tend to merge
one into the other. Such species as the Podalirius and
X, olivacea in which the anterior yellow is largely de-
veloped clearly belong to the outlying portions of the
Lycoid group (Plate XVIII), whereas Elis cvlels has more
affinity with the succeeding group; and this latter again
merges away into the great black group with blue wings
(Plates XX and XX1). The two largest Xylocopas CY.
lateritia and X. flavorufa) only belong to the outskirts of
the association, as the red of the thorax is in them of a
much darker tint and does not show up well in the Plate.
All the Hymenoptera of the group are found plentifully
on flowers; but as regards the J/elittia, the specimen
figured is, I believe, the only one known.

G. Group with Black Bodies, Blue Wings, and Red or
Yellow Heads (Represented on Plate XXIII).

Pompilus marshalli (fig. 20); P. dichrous
P (figs. 21, 22); P. anticus (fig. 24).

Hvmxopruna | 2 ommilide | Pompilus lascivus (fig. 23); Salius ta-
misieri (fig. 25).
Sphegidx Tachysphex fluctuatus (fig. 26).
COLEOPTERA Longicornia Jonthodes sculptilis (fig. 28).
DiprERA 3romophila caffra (fig. 27).

Converging towards these but with black wings instead
of blue are the Braconid Jphiaulax ruber (Fig. 31) and
the Cantharid Eletica rufa, var. (Fig. 29).

This is only a subdivision of the group of black-bodied
and blue-winged insects. The Hymenoptera have all
much the same habits as the species contained in that
group, though they are mostly of much smaller size. The
Longicorn Jonthodes bears a very good general resemblance
to the blue-winged, yellow-legged Salius dedjax, owing to
its blue elytra and yellow legs; it is not a particularly
common species, being diurnal and arboreal in its habits.
It possesses a strong scent-like smell, and the mimicry is
probably Miillerian. The Lromophila fly is very plentiful ;
it is the most sluggish fly known to me, and settles about
on trees and bushes in a very conspicuous manner. It
ejects a yellow liquid from the mouth when handled, and
was refused when offered to my baboons and Cercopithecus
monkey.

yay Mr. G. A. K. Marshall on

H. Group with Black and Yellow-Banded Bodies: all
Hymenoptera (Represented on Plate XXIII).

Scoliade. Pompilide. Ichneumonide.
Myzine capitata Pompilus festivus Metopius discolor
(fig. 35). (fig. 34). (fig. 36).

In Europe, owing to the predominance of the genus
Vespa, black and yellow bands constitute a very dominant
type of coloration among the Hymenoptera; but in
Mashonaland (where Vespa is entirely absent) this pattern
is of comparatively rare occurrence, and, except in the case
of one large Scolia and some Bembex, is confined to small
insects. The two Aculeates in the above group are
common frequenters of flowers, and to them might have
been added several small species of Hilis, etc. ; the Ichneu-
mon is common in woods.

I. Group with Dark Wings and Black-and- Yellow Legs:
Ichneumonid Models (Represented on Plate XXIII).

HYMENOPTERA. COLEOPTERA. HEMIPTERA.
Ichneumonide. Longicornia. Reduviide.
Osprynchotus flavipes Litopus dispar Pirates wneicollis
(fig. 32). (fig. 30). (fig. 33).

The Litopus is evidently one of the protected Ceram-
bycids, as shown by its diurnal habits, blue elytra, brightly-
banded legs, and strong smell. The Pirates, with its dark
wings and black-and-yellow legs, shows a distinct approxi-
mation to the former insect, which is probably of a
Miillerian nature, as it is capable of emitting a strong
and unpleasant smell, can pierce very effectively with its
strong rostrum, and has the power of stridulation. Ospryn-
chotus is also a conspicuous and very common insect. In
South Africa it seems to be principally parasitic upon
Pelopeus spirifex, and it is a curious thing that although
these two insects are so very different in coloration (except
the hind-legs), yet there is an undoubted similarity
between them when seen together on the wing.

J. Black and VYellow-Barred Braconid Group and Mimies
(Represented on Plate X VIII).

Braconidex Phanomeris dubius (fig. 59).

a 2 A
HYMENOPTERA Ichneumonide Pimpla tuberata (figs. 60, 61).
HEMIPTERA Redwviide Callilestes stigmatellus (fig. 62).

The Bionomics of South African Insects. 533

This is a beautiful little group, presenting very striking
resemblances. The Phanomeris is doubtless the chief
model, being a common species with a strong smell and a
slow, conspicuous flight. The colouring of the Pimpla is
probably Miillerian, while that of the Reduviid is cer tainly
Batesian. he latter is evidently a very scarce insect, the
only example which I have met with having been captured
accidentally in mistake for the Braconid, to which it bears
a wonderful resemblance on the wing.

K. Black and Red Braconid Group and Mimics (Repre-
sented on Plate XVIII).

f Bracon coccineum (figs. 53, 54); Iphian-

HYMENOPTER. raconidx

NOPTERA Braconade \ lax pictus (fig. 55); I. flagrator (fig. 56).
COLEOPTERA Longicornia Oberea scutellaris (fig. 57).
HEMIPTERA Rediuviide. Callilestes bicolor (fig. 58),

In this group the pattern is certainly set by the
Braconidx, which are common, conspicuous, slow-fying
insects, protected by their strong smell. The Reduviid is
an admirable mimic of them (Batesian, as I believe) both
at rest and on the wing; it is a scarce species, and
frequents the same stations as the Braconids. The Longi-
corn agrees also in the latter respect, but its exact relation-
ship to the group is doubtful; normally it is not a very
common species in Salisbury, but in one or two seasons it
has appeared quite plentifully, settling on low plants on
wooded kopjes.

L. Diptera Mimicking Single Species of Hymenoptera rather
than the General Type of a Group.

a. Asilid Fly Mimiching Xylocopid Bee (Represented on
Plate XXII).

HYMENOPTERA. DIPTERA.
Xylocopa flavorufa (fig. 19). Hyperechia marshalli (fig. 20).
[With reference to these insects Mr. Marshall wrote :]

Salisbury, Feb. 12, 1899.—I was immensely delighted on
catching the large Asilid tly, and I have been delaying the

534 Mr. G. A. K. Marshall on

box on purpose to include the large Yy/ocopa bee which it
mimics so admirably; but though usually they are common
enough J have not seen a single example for the last three
weeks. The fly completely deceived me when on the
wing, but I happened to notice it settle on the trunk of a
tree, and it struck me there was something curious about
the way it alighted for a Yylocopa. It is the only
example I have seen.

Salisbury, April, 25, 1899.—I have already got a couple
of the Xylocopas for you, and I have seen another
specimen of the Zaphria that mimics them, but unfortun-
ately failed to catel it.

B. Syrplid Fly Mimicking a Wasp (Represented on
Plate XXIII).

HYMENOPTERA. DIPTERA.
Polistes marginalis (fig, 40). Ceria gambiana (fig. 41).

This is an excellent and typical case of Batesian
mimicry. The wasp is a very common species, building
a small hanging nest on bushes or grass-stems. Like most
of its congeners it is a somewhat sluggish insect, and in-
stead of flying away when approached, adopts a bold and
defiant attitude, stinging sharply when disturbed. It
visits flowers freely, and may there be seen in company
with the Ceria, which resembles it so closely (especially
on the wing) that I have been deceived by it over and
over again,

y. Bee-like Group (Represented on Plate XXIII).

HYMENOPTERA. DIPTERA.
Apide. Syrphide. Asilide.
Megachile apiformis Eumerus, sp. nov. ? l.axenecera inollis
(fiz. 37). (fig. 38). (tig. 39).

The resemblance of the two flies to the Megachile
is very marked in the field, especially in the case of the
Laxenecera ; although the plate does not do justice to the
Humerus. I have on several occasions seen all three
species flying together in the vicinity of flowers.

The Bionomics of South African Insects, 535

M. Group of Ant-like Insects captured together (Repre-
sented 14 times the natural size on Plate XIX).

( Camponotus sericeus (fig. 54)3 C.

HYMENOPTERA Form? cidx cosinicus (figs. 55, 56); C. sp.
\ (fig. 53).

HEMIPTERA Pyrrhocoride Megapetus atratus (figs. 57, 58).

ORTHOPTERA Locustid Myrmee ophana ? fallax (fig. 59).

The above insects were all caught on the same day
(Feb. 17, 1901) on a single plant—a small bushy vetch.
The Pyrrhocorid habitua ally frequents this plant, being
fond of sucking the young pods; the ants are more or
less ubiquitous, but are principally attracted to this vetch
by the juice which exudes from the injuries made by the
bug; the Myrmecophana, which is apparently a very rare
insect, was probably only a chance visitor. The latter, in
spite of its long antenne, bears a very strong resem-
blanee to an ant, and might very readily be passed over
for one of these insects ; it is probably a Batesian mimic.
The bug is not nearly so ant-like in its mature form
(which is shown in Plate XX) as in its earlier stages, at
which period the similarity is most striking both in shape
and movements. The insect is a comparatively common
one, and the mimicry has probably a Miillerian character.

[The following extracts from letters refer to this in-
teresting group. Mr. Malcolm Burr, to whom I have
shown the Locustid, thinks that it may be the same
species as Mf. fallax, inasmuch as the habits of the latter
are not certainly known, and the green marking which
obliterates the unant-like parts of the body-form had
faded to a pale yellowish tint much lke that of the
corresponding parts of the specimen described by Brunner
von Wattenwyl (verhandl d. K. K. Zool.-botan. Ges. in
Wien, Bd. xxxiii, 1883, Pl. XV, tigs. la and 14).—E. B. P.]

Salisbury, April 25, 1899.—The Locustid ant-mimicker
Myrmecophana occurs both here (Mashonaland) and in
Natal, though very rarely, but it is perhaps a different
species from JM. fallax. It does not live on the ground
but on low plants, which are also frequented by the ant it
mimics, and the light parts are pale green; we have also
a bug which mimics the same ant.

Salisbury, April 19, 1901.—I expect you will be glad to
have an example of Myrnccophana,.it makes a grand
series with the ants and bugs; the two latter can often be

536 Mr, G. A. K. Marshall on

found together on a certain vetch, but it was a bit of luck
getting the Locustid there too.

33. MIMETIC RESEMBLANCE OF MANTISPIDH TO HyMEN-
OPTERA. (HK. B. P.)

Prof. W. M. Wheeler, of the University of Texas, was,
so far as I am aware, the first to observe the mimicry of
Hymenoptera by Mantispidex. “While studying the prairie
insect fauna of south-eastern Nebraska early in the
summer of 1888,” he observed that Mantispa brunnea
(Say.) closely resembled Polistes variatus (Cress), resting
half-concealed like the wasp “on the petioles in the
terminal leaf-clusters of the golden rods.” He was at
first quite deceived, and took care to avoid being stung.
“The colouring of the Polistes is carefully copied ; the body
is banded with yellow, brown, and black, the wings are
smoky brown, and the legs yellow. While lying in wait
the Mantispx closely appose their large raptorial fore-legs
to the lateral faces of the prothorax, which, when these
appendages are extended, is so narrow as to resemble but
slightly the wasp’s thorax. The wings are carried in the
same manner by both insects. Several times during the
course of a week I found these two insects... resting
in the same position, both intent upon the slaughter of
the many insects... which swarmed about the rank
vegetation” (Proc. Nat. Hist. Soc. Wisconsin, U.S.A.,
April 1889, p. 217). Professor Wheeler considers it to
be an example of protective mimicry.

Mr. R. Shelford has recently observed that at least four
species of Mantispa from Borneo and Singapore are beauti-
fully mimetic of Ichneumons, Bracons, or Aculeates. His
observations are now being published by the Zoological
Society. 1 sent a photograph of some of his examples to
Mr. Marshall, who replied with the observation printed
below. These interesting records constitute, so far as I
am aware, a distinct addition to the list of insect mimics
of the Hymenoptera. Mr. McLachlan, whom I have con-
sulted, writes that he cannot find anything further recorded
about such resemblance on the part of Mantispidy. There
can be little doubt, after these observations from three such
widely-different regions, that mimicry of the Hymenoptera
will prove to be prevalent in the group. Mr. Shelford and

The Bionomics of South African Insects. 537

Mr. Marshall both call attention to the cause which has
doubtless prevented the fact from being generally recog-
nized at an earlier date, viz. the changes which take place
in dried specimens of JMantispa. On this account, and
because of the important part played by movement, the
appreciation of the mimetic resemblance required the
study of the living insect.

“ Salisbury, Sept. 21, 1900.—The large South African
Mantispa grandis is an excellent mimic, on the wing, of the
Belenogaster wasps. I caught one at Malvern, on my way
home in 1896, which I gave to McLachlan. This insect
completely took me in; it flew out of a loquat-tree which
I was beating, and I at once took to my heels thinking I
had struck a nest of these vicious wasps. Fortunately I
kept an eye on the insect, and, as it seemed to be a species
of Belenogaster new to me, I followed it up and caught it,
when to my surprise and delight it proved to be only a
Mantispa. Unfortunately in a dried specimen the rcsem-
blance is much spoilt by the shrivelling and discoloration
of the abdomen.”

34. CONVERGENT GROUPS OF SoUTH AFRICAN HEMI-
PRBRAN (Goan, Key MM)

A. Black and Red Lygevoid Group (Represented on
Plate XIX).

: f Lygeus rivularis (fig. 44); L. elegans (fig. 46); L. crudelis
Lygeide \ (ig. 47); Graphostethus servus (fig. 45).
Ivduviide Reduvius sp. (fig. 43).

In this group I consider that the Lygzids form a
Miillerian association, of which the Reduwvius is probably a
Batesian mimic. The former insects are very abundant,
occurring on many different plants, but the Lygexi are
especially fond of the balloon-like seed-vessels of Gompho-
carpus. The Reduvius inhabits much the same stations,
though I have never seen it (to my remembrance) actually
in company with the Lygzeids, and it is a decidedly rarer
insect,

538 ' Mr. G. A. K. Marshall on

B. Growp of Yellow Hemiptera with Black Apex and one
or two Black Bars (Represented on Piate XIX).

At Malvern, Natal.

Pyrrhocoride. Reduviide.
Dysdercus nigrofasciatus Phonoctonus nigrofasciatus

(fig. 49). (fig. 48),
At Salisbury, Mashonaland.

} E

Pyrrhocoridex. Redurviide.
Dysdereus superstitiosus (fig. 50). Phonoctonus formosus (fig. 52).
an intermedius (fig. 51).

The significance of the mimicry in this group has not
yet been tested by experiment, and the exact relationship
of the Reduviids to the commonand undoubtedly distaste-
ful Dysderct is not quite clear. Dr. Dimock Brown, who
observed Phonoctonus in company with myself at Malvern,
suggested that its colouring may be pseudepisematic, and
that it may feed upon the Dysdercus which 1t mimics so
marvellously well. Personally I incline rather to the
belief that both this species and the northern P. formosus
are Batesian mimics. Both species occur but rarely (in-
deed, of the latter, I know only two specimens), they
do not possess the strong smell which characterizes
some of the Reduviids, and their Jointed rostrum is a
very inefficient weapon for protective purposes. I am
not aware that they have been observed feeding on Dys-
derct or even in company with them (cf. G. Breddin,
Zeitsch. f. Naturw. 1896, pp. 36-38).

[Breddin considers the resemblance of the Reduviid to
be a case of aggressive (pseudepisematic) mimicry, as
he thinks with Dr. Dimock Brown it would prey on the
Dysdercus. I believe that all such groups in the Hemiptera
are synaposematic.—E. B. P.]

35. MISCELLANEOUS OBSERVATIONS ON SOUTH AFRICAN
Insects. (G. A. K. M.)
A. Note on the Courtship of Limnas chrysippus.

Salisbury, June 26, 1900.—In some old notes I find
the following observation on the courtship of chrysippus,
When first observed the female was settled on the

The Bionomics of South African Insects. 539

ground and was sharply fluttering her wings to keep off
the male which was hovering above her. Whenever she
rested for a moment with open wings the male would
drop down on her, trying to settle on the costa of her fore-
wing in such a position that the badge on his hind-wing
came directly down on her head ; and while hovering over
her, his position was usually at right angles to hers, which
renders it probable that the badge is some sort of scent-
gland used for attractive purposes. The female however
kept on fluttering pretty incessantly, and the male kept
bumping down on her. Then another male came round
and the first one went off and had a skirmish with him
and drove him away. The female then took flight, the
male usually keeping above her and trying to beat her
down to the ground again. The female, on settling,
renewed her defensive fluttering, and the male, apparently
getting tired, flew off. The whole observation occupied
five minutes. I never saw any use made by the male of
his curious terminal tufts,

B. The possible meaning of the Sac of Female
Acrvine.

Malvern, May 14, 1897.—The species in which the sac
is best developed are Acr@a neobule and A. horta. With
regard to the use of the organ, I remember making some
observations at Salisbury in 1894 on A. caldarena and
A. nohara-halali while ovipositing, and I then came to
the conclusion that the sac was of no use during laying,
being apparently rather an obstruction than otherwise. I
therefore rather incline to your second suggestion, that it
is probably to prevent copulation a second time. This
view moreover seems to be borne out by what I have
noticed in the courtship of the insects. So far as I have
at present observed, Acrzeas appear to be the only butter-
flies which indulge in the system of ‘‘ marriage by capture.”
In such of the Nymphaline as I have watched, the males
have in no case attempted to seize the females, which,
when anxious to escape their addresses, did so either by
dodging among the vegetation or soaring. The females of
some Piering (notably Belenois, Pinacopters w) have a very
noticeable method of refusing the males; they settle with
wings outspread but with fore-wings directed backwards
so as almost to cover the hind-wings, and the abdomen is

540 Mr. G. A. K. Marshall on

raised in the air. This position is probably to prevent the
male running along the side, for copulation is effected
from the side. It might however be done in order to
allow the male to see by her abdomen that she was gravid,
for I have a case in my note-book (P. pigea) in which the
male ran up and felt the abdomen with his palpi and then
flew off. In the Acrzeas however I have observed several
cases of copulation taking place in A. petrva and A. horta,
and in all of them the male seized the female on the wing,
erasping her with his intermediate legs about the thorax
or base of the fore-wings, and they would fall struggling to
the ground, where coition would take place. If this is
the normal method of copulation, and unfortunately my
observations have been too few to enable me to feel sure
of it, then any organ which would protect the female from
the attentions of an unlimited number of males would not
only be useful but absolutely necessary.

Malvern, July 15, 1897.—The other day I saw a pair
of Acrva encedon struggling together on the ground, the
male clasping the female round the thorax from below.
Unluckily a second or two after I noticed them they
separated, so that I had not time to see whether it was
really the sac which prevented coition. However I caught
the female and found she had the sac fully developed and
hard.

C. A Rhodesian Muscid Fly Parasitic on Man.

[Mr. E. E. Austen informs me that the fly sent by
Mr. Marshall belongs to the J/uscidx, and is certainly a
near ally of the genus Lengalia. A closely similar or
possibly identical species with precisely similar habits
occurs on the West Coast of Africa. Mr. Austen’s observ-
ations upon this latter entirely confirm Mr. Marshall's
conclusions as to the method by which the larva enters
the skin. Mr. Austen proposes to publish a full description
of the species—E. B. P.]

Salisbury, April 19, 1901.—I should be glad to know
the name of the parasitic fly I send. It has been a great
scourge this year in Salisbury, especially among young
babies, the maggots forming a painful boil-like swelling
under the skin. One baby had no less than sixty maggots
extracted from it, and there have been several cases in
which they have had a dozen or more,

The Bionomics of South African Insects. 541

Salisbury, Sept. 27, 1901.—The fly which lays eggs in
man is very common here, but I have no specimens by
me; I will catch you a series as soon as they appear
again. The one I sent you was a male, the female is
very much larger. I am much puzzled to understand how
the larva obtains an entrance into the skin. It certainly
eannot be through the stomach as in the case of some
other bots. I fancy the egg or living larva must be laid
on the clothing, and the latter being very minute might
wander about and eventually enter the skin through a
pore without being felt. The position of the bots in many
cases renders it impossible for the egg to have been placed
under the skin by the mother.

AE EENDEX.

Description of a new species of Hyperechia, Schin.
(Family Asttip2), from Mashonaland. By ERNEST
K. AUSTEN.

HYPERECHIA, Schiner:

Verhandlungen der kaiserlich-koniglichen zoologisch-
botanischen Gesellschaft in Wien, xvi. Band, p. 673

(1866).

Hyperechia marshalli, sp. nov. (Pl. XXII, f. 20.)

3. Length 28 millim.

Black, abdomen steely; cheeks, posterior margin of thorax in front
of scutellum, outer side of front tibix, wnder side of thorax between
bases of legs and in front of front coxx, and outer side of middle
femora, except apical fourth, clothed with orange-rufous * hair: fringe
on posterior margin of thorax very conspicuous, and more ferruginous F
than orange-rufous.

Front and face clothed with ochraceous hair ; mysfax ochraceous
above and black below, with two or three black hairs in the middle

* Ridgway, “A Nomenclature of Colors” (Boston ; Little, Brown
and Coy., 1886), Pl. IV, 13.
+ Ridgway, op. cit., Pl. IV, 10.

542 Appendix.

line above. Tips of front femora on outside clothed with orange-
rufous hair; a large tuft of similar hair on the pleura below the
humeral angles. Thorax duller than abdomen, and, except on
posterior margin and also in front, where there is some ochraceous
pile, clothed with very short black hair. Leys thickly fringed with
black hair, except where otherwise stated. Wings of the usual
blackish-brown tint, with a slightly purplish sheen, which, however,
is not so conspicuous as in the case of H. xylocopiformis, Walker.

Salisbury, Mashonaland; Feb. 1899 (G. A. K. Marshall) :
one specimen. Type in the Hope Museum, Oxford.

I have much pleasure in associating this fine new
species with the name of its discoverer.

It may be of interest to note that the genus Hyperechia
occurs in the Oriental as well as in the Ethiopian Region :
Hyperechia (Laphria) xylocopifornis, Walk., the type of
the genus, was described from a specimen from Madras,
while HZ. fera, v. d. Wulp, occurs in Borneo.

Descriptions of new species of South African Rhynchota.
By W. L. DIsTanr.

Family PYRRHOCORID.

Megapetus, gen. nov.

Head elongately subquadrate, narrowed in front of insertion of
antennze, apex truncately rounded, outer margins of eyes not so wide
as posterior angles of pronotum, antennz four-jointed, simple, second
and third joints subequal in length, each shorter than first, fourth much
the longest ; pronotum with the lateral margins convex, narrowed
anteriorly to base of head, lateral angles spinously produced, posterior
margin truncate ; hemelytra rudimentary ; scutellum triangular ;
abdomen inflated ; rostrum reaching the intermediate coxie, basal
joint very robust, subequal to or very slightly shorter than second
joint ; legs simple, posterior legs much the longest, posterior femora
obsoletely spinous before apex.

Allied to Myrmoplasta, Gerst.
Megapetus atratus, sp. nov. (Pl. XIX, f. 57-58 x 14.)

Black ; antennz pale fuscous, basal half of first joint greyish,
fourth joint dark fuscous ; head anteriorly somewhat longly greyishly
pilose ; in ¢, base of abdomen black with a small whitish spot near
each anterior lateral angle, in Q base of abdomen brownish ;

Appendiw. 543

posterior pronotal angles distinctly spinously produced, the spines
directed a little backwardly ; the upper surface is very finely and
somewhat sparingly pilose ; base of anterior tarsi pale fuscous ; in ?
the intermediate tibize somewhat brownish.

Long. 8 millim. ; exp. pronot. angl. 25 millim.

Hab. MASHONALAND ; Salisbury (@. 4. AK. Marshall).
Type in the Hope Museum, Oxford, and in Coll. Distant.

Dysdercus intermedius, sp. nov. (Pl. XIX, f. 51.)

Above pale ochraceous ; head, anterior area and lateral margins of
pronotum, basal area of lateral pronotal margins, scutellum, body
beneath rostrum, and legs reddish-ochraceous or pale sanguineous ;
antenna, discal narrow transverse fascia to corium, membrane, tibia,
tarsi, and anterior margins of sternal and abdominal segments, black ;
anterior margin of pronotum broadly, posterior margin narrowly,
anterior and posterior margins of prosternum, lateral and posterior
margins of pro- and mesosterna, lateral and posterior margins of
abdominal segments and the cox, pale luteous; anterior femora
spined beneath at apex; second joint of antennz slightly shorter
than the first the base of which is reddish.

Long. 16 to 21 millim.

Hab. GERMAN Easr AFrica (Capt. Atkinson—Brit.
Mus.). MASHONALAND; Salisbury (G. A. A. Marshall).

Type, from German East Africa, in the British Museum,
Specimens from Mashonaland in the Hope Museum,
Oxford, and in Coll. Distant.

Intermediate between D. superstitiosus, Fabr., and D.
cardinalis, Gerst. From the first it differs by the absence
of the black fascia to the posterior margin of the pronotum
and the transverse fascia to the corilum; from J. cardinalis
it differs by the black tibize and tarsi.

Family REDUVIID.
Phonoctonus formosus, sp. nov. (Pl. XIX, f. 52.)

Pale luteous ; head, anterior lobe of pronotum, rostrum, anterior
and intermediate femora, base of posterior femora, and coxe
sanguineous ; antennz, scutellum (excluding apex), an indistinct
transverse fascia to corium at and somewhat confused with base of
membrane, tibia and tarsi, posterior femora, central area of inter-
mediate femora, membrane, and apex of rostrum black ; sternum

544, Appendix.

piceous, lateral and posterior segmental margins, luteous ; abdomen
beneath luteous, lateral and posterior segmental margins, sanguine-
ous, apex fuscous.

Anterior Jobe of pronotum posteriorly suleate, posterior lobe
entire, the last with its anterior lateral margins narrowly black ;
body beneath, sparingly and finely pilose.

Long. 19 millim.

Hab. ANGOLA (Brit. Mus.). MASHONALAND; Salisbury
(G. A. K. Marshall),

Type, from Angola, in the British Museum.
Specimens from Mashonaland in the Hope Museum,
Oxford.

Callilestes stigmatellus, sp. nov. (Pl. XVIII, f. 62.)

Ochraceous ; anterior lobe of pronotum, lateral margins of pro-
and mesosterna, the metasternum, and abdomen beneath reddish-
ochraceous ; antennae, apex of head, eyes, and a transverse fascia
between eyes, a postmedian transverse fascia and the apical angles to
corium, membrane, tarsi, and posterior tibize black ; bases of posterior
tibize, basal angle and a central spot to membrane, ochraceous,

Anterior and posterior pronotal lobes with a distinct central
suleation, scutellum with a strong, circular, basal foveate impres-
sion; posterior lobe of pronotum, scutellum and corium thickly
and finely punctate ; femora moderately nodulose and with the tibiae
distinctly pilose ; hemelytra extending considerably beyond the apex
of the abdomen.

Long. incl. membr, 14 millim.

Hab. MASHONALAND; Salisbury (G. A. XK. Marshall).
Type in the Hope Museum, Oxford.

Descriptions of new species of South African Hymenoptera.
By CoLoneEL C. T. BINGHAM.

Chaleis bicolor, sp. nov. (Pl. XXIII, f. 10.)

¢. Head, thorax, the cox and trochanters of the anterior, and
intermediate, and the upper side of the coxw, the trochanters, tibie
and tarsi of the posterior legs black, the femora and tibiw of the
anterior and intermediate legs, the front of the posterior coxx, the
posterior femora and the abdomen orange-yellow, the apex of the
mandibles castaneous, the tibial calearia of the hind-legs yellow.

Appendia, 545

Head and thorax coarsely closely punctured, cribrate, the wide deep
antennal groove on the face, and the mesopleure transversely striate,
the scutellum broad and convex above, posteriorly bidentate, the
postscutellum and median segment shining, the latter areolated and
bearing a strong tooth on each side; legs finely punctured opaque,
abdomen smooth and shining. The sides of the broad vertical
furrow on the front of the head, the cheeks and the metapleure
covered densely with long white hairs, the legs and the rest of the
head and thorax with sparse shining white pubescence. Wings dark
fuscous with a purple iridescence in certain lights,
Length § 9m.m.; exp. 17 m.m.

Hab.Salisbury,5000 feet, MASHONALAND, SOUTH AFRICA.

Bears a superficial resemblance to C. semirufa, Walker,
from the Oriental Region, but in sculpture and in the
distribution of colour widely ditterent.

Type in the Hope Museum, Oxford.

Iphiaulax ruber, sp. nov. (Pl. XXIII, f. 31.)

Q. Red, the front below the antenne, the head above, the cheeks,
except a narrow line behind the eyes, the antennx, the apex of the
mandibles and the claws black ; wings fusco-violaceous, the basal
three-fourths of the stigma and two obscure spots beneath it on the
fore-wing reddish hyaline. Head smooth and shining, a semicircular
transverse depression in front of the ocelli. Thorax glabrous, shining ;
mesonotum gibbous, the parapsidal grooves indistinct ; the scutellum
triangular somewhat laterally compressed, the pos‘scutellum trans-
verse ; the median segment elongate obliquely sloping to the apex
not areolated, the pear-shaped stigmata placed in a depression on
each side. Head, thorax and legs covered with a fine, short, erect,
brownish pubescence. Abdomen somewhat elliptical, as long as the
head and thorax united ; basal segment with a raised longitudinal
rectangular centre portion smooth and shining, the lateral margins
foveate and bounded by a carina, the apex of the raised part depressed,
longitudinally striate and margined by a trausverse broad carina;
2nd segment with a medial subtriangular plate smooth and
shining from which coarse divergent striz radiate in the deep
depressions on each side, these depressions bounded laterally and
posteriorly by raised carinz ; remaining segments coarsely punctured,
rugose, the sutures between the segments 2-5, and oblique lateral
grooves on segments 3-5 crenulate.

Length @ to apex of abdomen 20; of ovipositor 22 m.m. ; exp.
38 m.m.

TRANS. ENT. SOC, LOND. 1902,—PART Il. (NOV.) 36

546 Appendix.

Hab. Salisbury, 5000 feet, MASHONALAND (G. A. K.
Marshall).

Type in the Hope Museum, Oxford.

Phanomeris dubius, sp. nov. (Pl. XVIII, f. 59.)

?. Reddish-yellow, the apex of the mandibles, the eyes, the
antenne, a triangular spot enclosing the ocelli, and the ovipositor
black, the posterior tibizw shaded with fuscous black ; wings hyaline
yellow shaded as follows, fore-wing the stigma jet black, aspot at the
apex of the median cell, spreading across the nervure into the Ist
discoidal cell, a spot at the base of the 2nd discoidal cell, a bar
interrupted below crossing the dise of the wing below the stigma,
and the apical margin of the wing fuscous ; hind-wing, a spot beyond
the transverse nervure closing the median cell, and the apical margin
of the wing broadly, fuscous, Head cubical, the front above the
antenne, the vertex, occiput and cheeks smooth and shining, head
in front below the antenne closely and somewhat coarsely punctured
rugose. Thorax not broader than the head coarsely but sparingly
punctured, the mesonotum gibbous, the parapsidal grooves deep, the
scutellum compressed smooth, legs moderately long with the femora
and tibiew incrassate ; median segment finely and closely punctured
rounded above, and bearing a delicate median longitudinal carina.
Abdomen longer than the head and thorax united, elongate oval, the
basal twe segments finely striate above, the disc of the 2nd segment
raised, the raised portion semicircular, the depression on either side
of the raised portion above smooth, the suturiform articulation
distinct, crenate, the apical segments smooth and shining with
transverse impressions at their bases, these latter crenulate. Ovi-
positor longer than the head and body, the sheath densely pubescent.

Length @, to apex of abdomen 17 m.m.; ovipositor 26 m.m. ;
exp. 32 m.m.

Hab. Salisbury, 5000 feet, MASHONALAND, SouTH AFRICA
(G. A. K,. Marshall).

It is with much doubt that I record this species under
Forster’s genus Phanomeris. It has the appearance of a
Vipio, but there are no tufts of hair at the base of the
clypeus, the submedian cell in the fore-wing is longer than
the median cell, and in the hind-wing the submedian cell
is about equal to half the length of the median cell.

Type in the Hope Museum, Oxford.

Appendix. 547

Pompilus marshalli, sp. nov. (Pl. XXIII, f. 20.)

?. Resembles P. collaris, Saussure, from Madagascar, but the thorax
is comparatively longer, the scutellum not so raised and prominent,
and the median segment is almost cylindrical very convex above,
roughly transversely striate, and posteriorly truncate but not concave.
In P. collaris the median segment is smooth almost flat above, while
the posterior face is concave with the sides distinctly produced
backwards.

Black with a beautiful purplish bloom on the abdomen above ;
the head with the mandibles and the scape of the antenne, the pro-
notum, the tibiz and tarsi of the anterior, and the tibia and base of
the first joint of the tarsi of the intermediate and posterior legs red ;
wings fusco-violaceous the posterior scarcely lighter in colour than
the anterior wings. In P. collaris the fore-wing is markedly darker
than the hind-wing. Abdomen massive as long as the head and
thorax united. Base of the 2nd ventral segment with a distinct
transverse groove ; pygidial area densely pubescent.

Length 2 20; exp, 28 m.m.

Hab. Salisbury, 5000 feet, MASHONALAND (G. 4A. K,
Marshall).

This species belongs to Kohl’s ferreola group of
Pompilus.

Type in the Hope Museum, Oxford.

Description of a new species of Carabidee from South
Africa. By Guy A. K. MarsHatt.

Polyhirma bennettii, Mshl., sp. nov.

Long. 15 m.m. Length of elytra 8 m.m. ; width at base 1°75
before middle 3, at apex 1°75 m.m.

Body depressed and very elongate. Colour black with a broad
line of thin greyish pubescence from labrum to basal part of elytra ;
the fovex of the elytra filled with ferruginous pubescence, and at the
apex an elongate sutural white patch.

Head broadly depressed in middle, indistinctly punctured and
with a short central carina just behind the labrum, which is bare
and impunctate; eyes prominent; the band of pubescence very
broad in front, narrow posteriorly. Antenne strongly compressed,
black ; the three basal joints shiny and with sparse white pubescence

548 Appendi.

exteriorly ; the remaining points appear dull owing to their being
strongly aciculate, except for a narrow smooth central line. Pro-
thorax elongate, broadest at apex, sides subparallel nearly to middle
and thence strongly narrowed to base. A broad central furrow
throughout, containing the pubescent stripe, and a deep short stria
on either side of it at base ; punctuation strong, close and even. In
the basal portion the lateral part of the thorax is produced so as to
extend a good deal beyond the dorsal edge from which it is separated
by a very deep incision. The mesonotum is broadly exposed and
bears a part of the central pubescent stripe. Elytra very narrow and
elongate, scarcely broader than the head at their widest part, which
is before middle. Sides distinctly rounded, apex broadly truncate.
Dorsal surface very much flattened, with six sharp, narrow carine
(the sutural one not reaching the base) and a very short apical one
between the 5th and 6th ; all the carimex disappear before reaching
the apex; the 6th carine form the dorsal edges of the elytra, the
portions between them and the true edges being folded over under-
neath so as to form false epipleura. The spaces between the carine
are occupied by large reticulate fovew, filled with ferruginous
pubescence, which are largest near the suture and diminish in size
laterally. The basal sutural stripe is short, being about as long as
the apical white patch ; the latter is narrow and elongate, bifid
posteriorly and reaches the extreme apex. Legs black with fine
white pubescence ; the posterior pairs of femora are more strongly
compressed than in any other species of the genus.

This remarkable species was discovered at Somerset
West, Cape Colony, in January 1900, by Mr. E. N. Bennett,
M.A., Fellow of Hertford College, Oxford, to whom I have
dedicated it. Its depressed and narrow form is more
suggestive of a subcortical insect than of a terrestrial
Polyhirma, and the folding under of the elytral mar-
gins is a unique character in the genus. The species
to which it is most nearly allied is P. macilenta, OL,
from which it may at once be distinguished by its very
different facies; and it also differs in the following
points: (1) the cost of the elytra are extended further
towards the apex (as in P. semisutwrata, Chd.); (2)
the costa are sharper and straight, not undulating ; (3)
the prothorax is not nearly so heart-shaped, owing to
the sides not being rounded in front. The last two points,
as well as the shortness of the basal stripe on the elytra,
distinguish it from semisuturata.

The type is in the Hope Museum, Oxford,

A ppendix, 549

Description of a new species of TELEPHORIDA from
Mashonaland. By J. BouRGEOISs.

Lycocerus mimicus, sp. nov. (Pl XVIII, f. 11 $,f 12 2.)

Oblongus, subparallelus, niger, pubescens ; capite prothoraceque
dense punctulatis, nitidiusculis, hoe antice leviter reflexo-marginato,
ad latera incrassato et vage rufo-limbato ; elytris opacis, tomentosis,
ruguloso-punctatis, flavo-testaceis, apice nigris, costis duabus obsoletis
singulatim notatis; corpore subtus nitidiusculo, nigro, abdomine
lateraliter flavo-marginato ; unguiculis simplicibus, rufis.

¢. Prothorace subelongato, antice rotundato, lateribus parallelis,
subrectis, haud marginatis, disco longitudinaliter suleato ; abdominis
segmento ultimo bivalvato, valva inferiori cupuliformi.

?. Prothorace transverso, antice arcuato, lateribus minus parallelis,
anguste submarginatis, suleo dorsali obsoleto; abdominis segmento
ultimo integro, semilunato,

Long. 9-11 null.

Hab. MASHONALAND; Salisbury (G. A. K. Marshall).
Sex Ge, i)

Type in Hope Museum, Oxford.

C’est la premiére espece africaine connue du genre
Lycoverus, Gorham (Proc. Zool. Soc. Lond., 1889, p. 108).
De méme que ses congéneres asiatiques, dont elle différe
surtout par la coloration, elle présente une certaine analo-
gie de facies avec les Lycides du groupe des Calochromides.

t=)

Chez le fle prothorax est plus etroit et plus allongé que
chez la 9. En outre, dans l’exemplaire que j’ai sous les
yeux, le pronotum présente, de chaque cdté du milieu et
contre le liseré roussitre qui existe dans les deux sexes, une
autre petite tache rousse qui manque dans la 9. Mais je
ne saurais dire si cette particularité de coloration est

constante.

550 Explanation of Plates.

EXPLANATION OF PLATE IX.

INJURIES TO WINGS OF SoutH AFRICAN BUTTERFLIES.

Injuries inflicted at the apex or hind margin of one or both fore-
wings, or near the overlap of fore- and hind-wings, or at two or more
points in the total wing margin.

All the figures are about 2 of the natural size.

All the specimens were captured, on the dates mentioned, at
Salisbury, Mashonaland, 5000 feet, by Guy A. K. Marshall.

Fie. 1. Limnas chrysippus 2, Sept. 22, 1900. Very rarely settles
on the ground, so that the attacks of lizards are not likely
to be common.

2. Parosmodes icteria, March 9, 1898. Found in the bush and
woodland districts, settling on flowers. Probably injured
when flying, as it rests with its wings closed.

3. Junonia cebrene g, Feb. 15, 1899. Settles on the ground,
injuries were very probably inflicted by a lizard.

4, Acrea anemosa ¢, March 11, 1899. Flies high for an
Acrea, and never settles on ground : feeds on tree flowers,
and usually at some height. The injury was probably
caused by a bird.

5. L. chrysippus 9, March 11, 1898.

6. Catochrysops mashuna ¢, Sept. 29,1900.. Flies very rapidly,
settles on low flowers and the ground, rests at night on
grass-stems. The injury was probably caused by a
lizard.

A. anemosa 2, Jan. 2, 1899.

8. Teracolus omphale g, March 31, 1901. Flies rapidly,
settles on low flowers and ground, so that lizards are
probable enemies, but the narrow symmetrical notch
rather suggests a bird’s beak.

9. Atella phalantha &, March 22,1899. Flies rapidly, settling
on bushes and flowers and not on the ground. Birds are
the probable enemies.

10. L. chrysippus 9, July 14, 1901.

1l. L. chrysippus 9, Jan. 2, 1898. The evidence of crumpling,
the scratching of the surface, as well as the extensive
injuries point to the probable attack of a mantis.

12. A. phalantha 9, March 5, 1899. Tips of both fore-wings
snipped off, probably by a bird.

=

Fia. 13.

14.

16.

18.

HY)

20.

al.

23.

24.

Explanation of Plates. 551

A, nohara, form halali, March 9, 1898. Flies low and
settles on low flowers and the ground. A lizard is a
probable enemy except that the injury is unilateral and
the insect closes its wings in all but the short rests, when
it opens and shuts them.

Catopsilia florella 9, Dec. 18, 1898. Flies very rapidly,
rests on trees under leaves, visits flowers and bushes. It
only settles on the ground to drink in damp places. It
is unlikely to be attacked by a lizard, and the character of
the injury probably indicates a bird.

Precis sesamus 2, April 8, 1991. Probably injured by a
bird: the notch is too narrow for a lizard. The specimen
was evidently freshly emerged.

Terias brigitta 9, December 18, 1898. Flies slowly and
settles on the ground and low flowers. Probably attacked
by lizard.

C. florella $, Jan. 21,1899. The injury strongly suggests
the beak of a bird. Both wings are symmetrically
snipped.

C. florella g, Jan. 14, 1899. The habits imply that birds
are the usual enemies.

P. sesamus g, April 8, 1901. The shape of the tear is
such as might have been made by a lizard, and the habits
of the butterfly render it quite probable that the injury
was thus caused. A very fresh specimen.

Pseudonympha extensa g, Jan. 2, 1899. A woodland
species with feeble flight, settling on low flowers and the
ground. It was very probably attacked by a lizard.

Terias brigitta ¢, March 9, 1898. Similar to description
of Fig. 16.

Belenois severina ¢, Jan. 25, 1899. Flight like that of
Teracolus omphale, see Fig. 8 description. Attack of
lizard probable.

Precis archesia g, April 8, 1901. The character of the
injury suggests the attack of a bird. Very fresh
specimen.

Precis antilope, form simia 2, Dec. 31, 1898. All Precis
in woodland and open country settle frequently on rocks
and ground. The injury probably caused by a lizard.

B. severina 9, Feb. 1, 1899. Probably injured by a lizard.

552

Explanation of Plates.

EXPLANATION OF PLATE X.

INJURIES TO HIND-WINGS OF MASHONALAND AND Honarcric

BUurrEeRFLIES.

Injuries inflicted at the anal angle or hind margin of one or both
hind-wings, suggesting that the insect was being pursued or, if
settled, approached from behind.

All the figures are 3 of the natural size.

All the specimens not otherwise described were captured on the
dates mentioned by Guy A. K. Marshall at Salisbury, Mashonaland,
5000 feet.

Tey Me

9

“-

3.

“I

10.
Wile

Linnas chrysippus 2, July 14, 1901.

Atella phalantha 9, Jan. 18, 1899.

Hypolimnas misippus 9, April 10, 1898. Flies like its
model chrysippus until disturbed, when it is swift. Settles
on low flowers. Lizards probable enemies. Very fresh
specimen.

A. phalantha , March 26, 1899. Character of injury
strongly suggests a bird’s beak.

A, phalantha 2, Dee. 1900.

Byblia acheloia g, Feb. 25,1899. The species of Byblic
fly low at a medium pace, and settle on low plants and
occasionally on the ground, they rest at night on grass-
stems, Lizards are probable enemies, but the character
of the injury rather suggests a bird.

Acrea rahira 2, Dec. 31, 1898. Marsh insects, settling on
flowers in low marshy places where lizards are not
often seen, hence birds are more probable enemies. Mr.
Marshall noted concerning the specimen here represented,
when it was in the fresh state, “judging from the state
of the abdomen this insect had been caught and rejected,
presumably by a bird.”

Nyctemera leuconoé, April 8, 1901. Never settles on ground.
It almost invariably covers its hind-wings direetly it
settles, assuming a A shape, although it occasionally
walks a short distance with its wings erect. The injury
was almost certainly inflicted during flight by a bird.

Pyrameis cardui, var. 9, Jan. 1900. Often settles on
ground. Lizards probable enemies.

Byblia ilithyia g, Jan. 27, 1899.

B. ilithyia g, Feb. 1, 1899.

Fig. 12: .
LS:

if).

29.

30.

Explanation of Plates. 558

B. acheloia 3, Feb. 22, 1899.

Cyclopides willemi ¢, March 1, 1899. A woodland insect
settling on flowers and never on ground. Rests at night
on grass-stems. Probably attacked on the wing, as the
injury is unilateral and the butterfly closes its wings
when settled.

Psewdonympha vigilans &, March 17, 1901. A rock butter-
fly, lizards the probable enemies.

Junonia cebrene ¢, Jan. 14, 1899.

* Peach. 1899)

C. willemi g, March 11, 1899. If this butterfly was
attacked, a bird is the probable enemy.

Herpxiia eriphia 9, March 9, 1898. Flies slowly and
rather low, settling on flowers and the ground like a
Teracolus. Lizards the probable enemies.

Mylothris riippelii g, Feb. 1900. Slow flapping flight,
settling on flowers and twigs of bushes in exposed
positions. The wings are generally open during a short
rest. The hind-wing was probably shorn through by a
bird.

Terias senegalensis ¢, Jan. 21, 1899.

T. regularis 9, May 24, 1901.

Belenois severina 9, Jan. 25, 1899.

55 - g, Feb. 15, 1899.

Catopsilia florella 3, Dec. 18, 1898.

Teracolus achine ¢, April 16, 1899.

Argywnis cybele, Aug. 2,1897. Captured by E. B. Poulton,
near North Lake, Hartland, Wis, U.S.A. <A flower-
haunting fritillary very probably injured by a bird.

B. severina g, Feb. 15, 1899.

Epinephele janira 9, Aug. 15, 1900. Captured by A. H.
Hamm, near Dawlish, 8. Devon. Both hind-wings
probably shorn through by a bird ; lizards are not im-
portant eneinies in this country. The straight line of the
injury also suggests a bird’s beak.

Melanargia galathea, July 21, 1898. Captured by E. B.
Poulton on the Stalden Rd., near Visp, Valais, about
2300 feet. The injury might well be by a lizard, but in
Europe they are not nearly such important enemies as
they become further south.

Lycena icarus, July 26,1898. Captured by E. B. Poulton
at Creux de Genthod, N.W. shore of the Lake of Geneva,
1230 feet. The character of the injury clearly suggests a

bird.

554 Kuplanation of Plates.

Fic. 31. Vanessa atalanta g, Aug. 23, 1897. Captured by F. A.
Dixey at Morthoe, N. Devon. The injury can hardly
have been produced otherwise than by a bird snipping a
notch in all four wings when they come together in the
attitude of repose as shown in the figure.

32. Hpinephele lycaon, July 22, 1898. Captured by E. B.
Poulton between Visp and Stalden, Valais, about 2450
feet. The same conclusion as in the description of
Fig, 29,

33. Vanessa atalanta ¢, Sept. 23,1901. Captured by A. H.
Hamm in the University Parks, Oxford. The character
of the injury implies an attack by a bird, probably made
in one of the short rests when the insect assumed the
attitude shown in the figure.

EXPLANATION OF PLATE XI.

INJURIES TO DIRECTIVE MARKS AND STRUCTURES ON THE WINGS
oF South AFRICAN BUTTERFLIES.

Injuries inflicted in the neighbourhood of special marks or
structures near the anal angle of hind-wing or apex of the fore-wing.

All the figures are about 3 of the natural size.

All the specimens not otherwise described were captured, on the
dates mentioned, at Salisbury, Mashonaland, 5000 feet, by Guy
A. K. Marshall.

Fie. 1. Precis antilope g, May 19, 1898. Rests on leaves of low
herbage and rarely on the ground. Birds are the most
likely enemies, and the specimen was probably at rest
when both “tails” were shorn off.

2. Precis cuama 9, May 1900. Same habit and conclusions
as in the last figure.

3. Spindasis natalensis, June 13, 1900. Captured by
Champion B. Russell near Eshowe, Zululand. Flight
very rapid, settles on outer leaves of trees and never
on ground. Almost certainly injured by a bird.

4. Precis archesia g, March 9,1901. This unilateral injury

was probably inflicted on the wing.

Charaxes guderiana %, Dee. 31,1898. All Charaxes here

represented fly and settle like S. natalensis. Birds by

or

Fig. 6.

~I

8.

10.

1d:

EHxplanation of Plates. 555

far the most probable enemies unless there is evidence of
the attacks of a mantis (possible but not likely in the
example represented in this figure).

Tarucus plinius g, March 11, 1899. Flies at medium pace,
settles on bushes and low trees, birds the probable
enemies. The injury moreover suggests a bird.

Hypolycena philippus g, March 11, 1899. Habits and
conclusions as in the last figure. These two Lycanids
and many other butterflies will settle on the ground in
damp spots to drink, but Mr. Marshall has not seen
lizards in such. places.

Papilio demodocus ¢, March 2,1901. Flies rapidly, settling
on flowering bushes ; probable enemies mantides or birds ;
lizards improbable. Character of injury suggests posterior
part of both hind-wings shorn off while at rest by a
bird.

P. demodocus &, Jan. 24, 1901. Same conclusion as in
last figure.

Charaxes achemenes 9, March 6, 1898. See description of
Fig. 5.

Catochrysops parsimon 2, Jan. 25, 1899. Very rapid
flight, settling on ground and low veldt flowers and at
night resting on grass-stems. Lizards probable enemies,
but the character of the injury suggests a bird.

Axiocerces harpax ¢, March 3, 1899. Habits and con-
clusions as in Figs. 6 and 7.

Charaxes saturnus g, March 6, 1898.

ss % », March 12, 1898.

C. guderiana 9, March 6, 1898. In this and the two
preceding figures, see description of Fig. 5.

Uranothauma poggei g, Jan. 1901. A woodland species
flying at medium pace and settling on low flowers. Birds
probable enemies.

T. plinius 9, March 11, 1899. Injury suggests bird.
Probably seized on the wing.

Axiocerces amanga ¢, Dec. 27, 1900. Habits and con-
clusions as in Figs.6 and 7. Probably seized at rest with
wings closed.

H. philippus $, March 11, 1899. Injury suggests bird.
Probably attacked at rest.

C. saturnus g, Jan. 2, 1899. Symmetry of injury suggests
conclusions as in Figs. 8 and 9.

Teracolus achine g, April 9, 1899. “

“f yf 3) April 2, 899.

556 Explanation of Plates.

Fia. 23. Teracolus achine 2, March 11, 1899. In this and the two
preceding figures the unilateral injury suggests an attack
on the wing.

24. Pyrameis cardwi ¢, Dec. 31, 1898.

EXPLANATION OF PLATE XII.

SEASONAL PHASES OF BUTTERFLIES OF THE GENUS PRECIS.

Representation of parent and offspring in Precis sesamus and P.
antilope.

Demonstration of the seasonal phases of South African Butterflies
of the genus Precis.

All the figures are about ,; of the natural size.

All the specimens represented were captured or bred by Guy A.
K. Marshall.

Fria. 1. Precis sesamus, form natalensis 9, Salisbury, 5000 feet,
captured Feb. 27, 1898, after it had laid three eggs.
Parent of butterflies represented in Figs. la and 1b.

la. Precis sesamus , offspring No.1 of butterfly represented in
Fig. 1. Egg laid Feb. 27, hatched March 5, larva pupated
March 31, imago emerged April 15.

1b. Precis sesamus, form natalensis 2 , offspring No, 2 of butter-
fly represented in Fig. 1. Egg laid Feb. 27, hatched
March 5, larva pupated April 5, imago emerged April 20.
A distinctly dark individual showing some slight tend-
ency towards sesamus, especially in the width of the
black margin of the hind-wings and the size of the blue
spots in this margin.

These two offspring show the overlap of summer and
winter phases remarkably well. The summer form,
Fig. 1b, even appeared a few days later in the beginning
of winter than the winter form, Fig. la. At the same
time the former is unusually dark.

2. Precis sesamus, form natalensis 2, Salisbury, 5000 feet,
captured March 6, 1898, after it had laid one egg. Parent
of butterfly represented in Fig. 2a.

2a. Precis sesamus g , offspring of butterfly represented in Fig. 2.
Egg laid March 6, hatched March 12, larva pupated
April 7, imago emerged April 30. The last part of larval

Explanation of Plates. 557

life and the first part of pupal were passed in a damp jar
(March 30 to April 5). The imago is nevertheless a
characteristic example of the dry phase.

Fic. 3. Precis antilope 2 , form simia, Salisbury, 5000 feet, captured
Feb. 23, 1902, after it had laid eleven eggs. Parent of
butterflies represented in Figs ms and 3b. The under-
side is shown on Plate XITI, fig.

3a. Precis antilope 9, offspring No. 1 of yen: represented in
Fig. 3. Eyg laid Feb. 23, hatched March 1, larva pupated
April 10, imago emerged April 27. The under-side is
shown on Plate XIII, fig. 4a.

3b. Precis antilope ¢, offspring No, 2 of butterfly represented
in Fig. 3. Evg laid Feb. 23, hatched March 1, larva
pupated April 14, imago emerged April 29. The under-
side is shown on Plate XIII, fig. 4b.

4. Precis archesia 9 , captured Oct.7, 1897, at Malvern, 800 feet,
near Durban, Natal. This insect was in coitw with the
form represented in Fig. 5

5. Precis archesia ¢, a form varying somewhat distinctly in
the direction of the wet phase (pelasqis), captured in coitu
with the insect represented in Fig. 4. The tendency
towards pelasgis is better shown upon the under-side,
Plate XIII, fig. &

EXPLANATION OF PLATE XIII.

SEASONAL PHASES OF BUTTERFLIES OF THE GENUS PRECIS.

Under-sides of seasonal phases of Precis sesamus, P. antilope, and
P. archesia, including those of P. antilope and its two offspring
shown on Plate XIT.

All the figures are about ,°, of the natural size.

All the specimens not otherwise described were cap{ured by Guy
A. K. Marshall.

Fra. 1. Precis sesamus, form natalensis 9, under-side of wings ;
Salisbury, 5900 feet, captured Feb. 20, 1898. Comparing
this with Figs. 1, 1b, and 2 on Plate XII, it is seen that
the under-side, although very similar to the upper, is even
more conspicuous than it. The difference is brought
about by the pale spots on the black basal patch of the

558 Explanation of Plates.

hind-wings and the more distinct white markings in the
black marginal band, on the under-side; also by the
paler tint of the ground-colour, making a stronger
contrast with the black markings.

Fria. 2. Precis sesamus 9, under-side of wings; Salisbury, 5000 feet,
captured April 2, 1898. Even in the absence of colour
the astounding difference between this and the wet-season
phase (see Fig. 1) is clearly shown.

3. Precis sesamus, form natalensis ¢, strongly tending towards
the dry phase, under-side of wings; Machakos, British
East Africa, captured June 6, 1900, by Mr. and Mrs. 8,
L. Hinde. This beautiful variety stands well on the wet,
or red, side of intermediate.

4. Precis antilope, form simia 9, under-side of wings ; Salis-
bury, Feb. 23, 1902. Parent of butterflies represented in
Figs. 4a and 4b. This figure shows the under-side of the
specimen represented in Fig. 3, Plate XII. It is seen
that the under-side is much more conspicuous than the
upper in the points mentioned above in the description
of P. sesamus, form natalensis (Fig. 1) ; especially in the
far greater contrast between the lighter ground-colour
and the black markings and in the amount of white in
the black border.

4a. Precis antilope 2, under-side of wings. Offspring No. 1 of
butterfly represented in Fig. 4. This figure shows the
under-side of the butterfly represented in Fig. 3a, Plate
XII. The difference between the under-side in offspring
and parent is as astonishing as in the two phases of
sesamus, While the difference in shape of the wings is
even more remarkable.

4b. Precis antilope ¢, under-side of wings. Offspring No. 2 of
butterfly represented in Fig. 4. This figure shows the
under-side of the butterfly represented in Fig. 3b, Plate
XII. The great difference in the character of the under-
side in the two offspring (compare this figure with the
preceding) is well shown, although both represent dead
leaves equally well.

5. Precis archesia, form pelasgis ¢, wunder-side of wings ; Salis-
bury, 5000 feet, captured Dec. 11, 1897. The figure
shows well that the difference between the two phases of
this species is far more important upon the under- than
the upper-side. It is also seen that the pale, conspicuous,
sharply-outlined, black-spotted band of the wet phase
represents the intensely cryptic mid-rib-like streak of the

Explanation of Plates. 559

dry phase (compare this figure with Figs, 6, 7, and the
somewhat intermediate Fig. 8). .

Fic.6. Precis archesia ¢, under-side of wings ; Salisbury, 5000
feet, captured April 30, 1899. The mottled browns and
greys of the under-side are very markedly different from
the uniform dark brown of the wet phase. Great varia-
tion in these mottled tints occurs in different individuals,
harmonizing with the varied appearance of rocks, being
at the same time dead-leaf-like.

7. Precis archesia?, under-side of wings; Berea, near Durban,
Natal, captured April 5, 1896, by Mr. D, Chaplin. In
this variety of archesia the ground-colour is comparatively
unmottled, and to this extent transitional towards the
pelasgis form. This difference upon the under-side corre-
sponds with a slight approach in the same direction on
the upper-side, where the deep red-brown band is broader
and more continuous than is usual in typical archesia.

8. Precis archesia ¢, under-side of wings ; variety distinctly
transitional towards the pelasgis (wet) phase ; Malvern,
800 feet, near Durban, Natal. This specimen was cap-
tured in coitu with a 2 of typical pelasgis, Oct. 7, 1897.
The intermediate character is seen in the conspicuous
broad light band as well as in the unmottled ground-
colour. Comparing this with Figs, 5 and 6, it is seen that
in spite of this distinct tendency towards pelusgis, the speci-
men represented in Fig. 8 is still well on the archesia side of
intermediate, and would doubtless be well concealed with
its wings closed. The character of the surface of both
wings outside the broad band is especially unlike pelasgis,
as is the gradual transition of the outer border of the
band itself into the ground-colour. The upper-side of
the specimen here represented is shown on Plate es
fig. 5, and that of the Q pelasgis with which it was in
coitw on Plate XII, fig. 4.

560 Kxplanation of Plates,

EXPLANATION OF PLATE XIV.

Mimicry IN MASHONALAND BUTTERFLIES.

Acreine and Lycznid mimics of Limnas chrysippus. Incipient
mimicry of Acrweas by Lycenidxe and Hesperide.

All the figures are about 8, of the natural size.

All the specimens figured on this plate were captured by Guy A.
K. Marshall in Mashonaland.

Fie. 1. Limnas chrysippus 9, April 9, 1899. Captured together
with the specimens represented in Figs. la, 2, and 2a at
Salisbury (5000 feet). The specimen is rather small ;
the ground-colour is of the dark-brown shade char-
acteristic of the Ethiopian Region.
la. L. chrysippus, under-side @. The peculiar shade of
the apex of the fore-wing beyond the sub-apical white bar
much resembles the ground-colour of the hind-wing, but
is distinguished from it by a faint greenish-orange tinge.
It is seen to be mimicked in the under-side of Mimacrea
marshalli (Fig. 2a), and less perfectly in that of Acrea
encedon (Fig. 3a).

2. Mimacrea marshalli ¢. A beautiful mimic of chrusippus
(compare Fig. 1). The shape of the sub-apical white bar
however more strongly suggests that of Acrxa encedon
(Fig. 3).

2a. M. marshalli, under-side ¢. The spots at the base
of the hind-wing as well as the shape of the sub-apical
har suggest encedon rather than chrysippus (compare
Fig. 3a), although the mimicry of the latter is strong
and evident.

3. Acrea encedon ¢, Dee. 1900. Captured together with the
specimens represented in Figs. 3a, 4, 6, and 7 at Umtali
(3700 feet). This, the typical form of the species, is seen
to be synaposematic with chrysippus.

3a. A. encedon, under-side ¢. The under-side is seen to be
synaposematic as well as the upper.

4. Catochrysops peculiaris, under-side ¢. The figure wWell
shows incipient mimicry of the black-spotted under-side
of an Acrexa.

5. C. mashuna, under-side 9, Nov. 7, 1897. Captured
at the Umcheki River (4200 feet). The resemblance
to an Acres is closer in this species than in the

Explanation of Plates. 561

last, because of the pale tawny ground-colour of the
under-side.
Fia. 6. Abantis tettensis, under-side. The under-side of the hind-
wings shows strong mimicry of an Acreeoid type.
7. Kedestes macomo, var., under-side. The under-side generally
shows the incipient mimicry of an Acreoid type.

EXPLANATION OF PLATE XV.

British East AFRICAN BurrerRFures Mimickine L. CHRYSIPPUS.

Acreine and Lyczenid mimies of Limnas chrysippus form klugit in
British East Africa, Varieties of Acrea encedon chiefly transitional
towards the form daira.

All the figures are about ,°; of the natural size.

Fig. 1. Limnas chrysippus, form klugit g, May 22, 1900. Captured
by Mr. and Mrs. 8. L. Hinde, at Machakos Road, British
East Africa (about 5400 feet). Very faint traces of the
sub-apical white band of chrysippus can be detected.

la. L. chrysippus, form klugii, under-side 2, Oct., Nov. 1900.
Collected by W. Doherty on the Kikuyu Escarpment,
British East Africa (6500 to 9000 feet), Distinct traces
of the sub-apical white band of chrysippus can be seen.
Compare Fig. la, Plate XIV.

2. Mimacrexa dohertyi 2, Oct., Nov. 1900. Collected in the
same locality as Fig. la, by W. Doherty. The position
and outline of the costal part of the sub-apical white bar
of M. marshalli can be distinctly traced in the contour of
the black markings, although the bar itself is only very
faintly paler than the fulvous ground-colour of the wing.
Compare Fig. 2, Plate XIV.

2a. M. dohertyi, under-side ¢, Oct., Nov. 1900. Collected in
the same locality as Fig. la, by W. Doherty. Traces of
position of white bar as in upper-side ; compare Fig 2a,
Plate XIV. The black spots on the under-side of the
under-wing resemble Acrea encedon (Fig. 3a) rather than
the Danaine model of both insects (Fig. 1a).

3. Acrea encedon, form daira ¢, June 6, 1900. Captured by
Mr, and Mrs. 8. L. Hinde, at Machakos, British East
Africa (about 5400 feet). Faint traces of the sub-apical

TRANS. ENT. SOC. LOND, 1902.—PART III. (NOV.) 37

562 Explanation of Plates.

white bar of fore-wing although the black ground-colour
of the apex is almost absent. Compare Figs. 4-7 and
Fig. 3, Plate XTV.

Fic. 3a. A. encedon, form daira, under-side ¢, Oct., Nov. 1900.
Collected in the same locality as Fig. la, by W. Doherty.
The under-side shows the same faint traces of the sub-
apical white bar as the upper-side. Compare Fig. 3.

4. A. encedon, form daira ¢, Jan. 28, 1909. Captured by
Mr. and Mrs. 8. L. Hinde at Kitui, British East Africa
(about 4000 feet). The sub-apical bar is rather more
distinct and the surrounding ground-colour rather darker
than in Fig. 3.

5. A. encedon, intermediate between form daira and encedon
¢, May 18,1900. Captured by Mrs. Leakey at Ndeje,
Bulemezi, near Mengo, Uganda. In this interesting
variety the sub-apical bar is very clearly defined by the
greatly increased darkness of the adjacent ground-coiour,
and thus becomes very distinct, although it is itself of a
darker shade than any of the specimens represented in
the three previous figures (3, 3a, and 4).

6. A. encedon tending towards the form daira in the darkness
of the sub-apical bar, and towards the form aleippina in
the whiteness of the inner part of the hind-wing ¢, May
19, 1900. Captured by Mrs. Leakey in the locality
described in Fig. 5. The apex of the fore-wing would be
that of typical encedon but for the darkness of the bar.
It is noteworthy that the costal spot of the bar is much
lighter than any of the other four component spots. On
the under-side of this specimen the bar is much lighter,
although not so white as in typical encedon.

7. A. encedon, form alcippina g, Feb. 27 to March 2, 1900.
Captured by Mrs. Leakey in the locality described in
Fig. 5. With white, conspicuous hind-wings this variety
closely resembles the form alcippus of Limnas chrysippus.
The fore-wings are as in typical encedon except that they
are somewhat darker than usual, suggesting, in the inner
contour of the black patch, an approach towards the form

lycia.

Explanation of Plates. 563

EXPLANATION OF PLATE XVI.

ComMMON WARNING CoLouRS oF MASHONALAND ACR#INE

BUTTERFLIES, ETC.

All the figures are about 4% of the natural size,
All the specimens represented on this Plate were captured by Guy
A. K. Marshall at Salisbury, Mashonaland, 5000 feet.

Fia.

6.

10.
Male

12.
13.

1
2.
3.
4
5

Acrxa doubledayi, form axina ¢, Dec. 31, 1898.

” ” ” ” ? ” ” ”
» caldarena 6, Dec. 31, 1898.
” ”» 2 3” ” bb}

5 nohara, form halali &, Dee. 31, 1898.

” ” 9 ” 2 bb) oh) ”
» rahira g, Dec. 31, 1898.
” ” 2 ” ” ”

» violarum, form asema 2, Dee. 31, 1898.

The above five species were therefore captured on the same
day in the same place together with other examples of
all species except rahira, as described on p, 492. The
group is a beautiful example of synaposematic (Miillerian)
resemblance between the species of a specially-protected
group inhabiting the same locality.

A, anemosa 9 , Jan. 7, 1899.

A. natalica,? ,,

A male of anemosa and a second female of natalica were
captured on the same day, Another beautiful example
of synaposematic resemblance between two fine species.

A. doubledayi, form axina ¢, April 6, 1898.

Baoris netopha 2, April 6, 1898.

The Hesperid at rest with its wings closed is a mimic of the
smaller Acreas such as that shown in Fig. 12. There
are several white spots on both upper- and under-side of
the fore-wing of the skipper, but these are concealed by
the hind-wing in the position of rest with the fore-wings
pressed well back between the hind. In the specimen
figured the fore-wings are not quite sufficiently far back, so
that the border of the dark shade which should be entirely
concealed is exposed together with a part of the most
anteriorly-placed white spot.

564

Explanation of Plates.

EXPLANATION OF PLATE XVII.

MIMIcRY AND WARNING COLOURS IN COLEOPTERA.

Mutilloid Coleoptera: Cleridx, Carabide and Cicindelide. Warning
patterns in the Carabid genus Anthia.

All the figures are about 5°, of the natural size.

All the specimens, not otherwise described, were captured at the
times mentioned, at Salisbury, Mashonaland, 5000 feet, by Guy A. K.
Marshall.

rGe le

Mutilla purpurata &, Nov. 1898.
55 tettensis ? Fe $3
Ry cepheus ° 53 5
ss leucopygat 2, FS
- sycorax os “
s horrida 93 » to Jan. 1899.
Atractonota nidsanti _,, » Mimic of ant.
” ” ”” ” ” ”
Polyhirma xnigma 5 - 5 » and yet
Mutilloid also.
Polyhirma xnigma 55 » to Jan. 1899. Mimic of

large ant, and yet Mutilloid also.

Eccoptoptera cwpricollis, Nov. 1898. A very perfect mimic
of a Mutilla.

Graptoclerus sp., Nov. 1898. Mutilloid mimic.

Polyhirma boucardi, Nov. 1898 to Jan. 1899.

Graphipterus antiokanus, Nov. 1898 to Jan. 1899. Resembles
the much larger Fig. 15, and, like it, Mutilloid.

Piexia selousi, Nov. 1898.

Polyhirma bilunata, Noy. 1898 to Jan. 1899. This and the
last species, when running, bear a general resemblance
to Mutillids.

Myrmecoptera marshalli, Nov. 1898.

4 anvicta 93 -
y bilunata ,, 5 This and the two
previous species are mimics of Figs. 15, 16, ete.

Myrmecoptera polyhirmoides, var. mashuna, Noy. 1898 to
Jan. 1899. A mimie of Fig. 13.

Anthia massilicata, Nov. 1898 to Jan. 1899.

” petersi ” ” ” ”
5 thoracica cs x 5; A

Explanation of Plates. 565

Fic. 24. Anthia sexguttata 9. Collected by Maj.-Gen. Hearsey,

India.
25, Anthia nimrod 2. West Africa.
26. » omoplata, var. mellyi, Nov. 1898 to Jan. 1899.

EXPLANATION OF PLATE XVIII.

Miuicry IN MASHONALAND INSECTS.

Mashonaland insects of many Orders with Lycoid pattern and
colouring: small groups probably mimetic of Ichneumonids.

All the figures are of the natural size.

All the specimens were captured by Guy A. K. Marshall, at the
times mentioned, and unless otherwise described, at Salisbury,
Mashonaland, 5000 feet.

Figs. 1 to 52 show a large and complex group, including species
of the genus Lyews and insects of various Orders and numerous
families mimetic of, or converging towards, the orange and black
coloration of this abundant and distasteful genus.

Fig. 1. Lyeus (Merolycus) rostratus ¢, Jan. 1899.
2. ” ” ” 2 ” ”
3. » (Acantholyerus) constrictus g, Jan. 1899.
4, » ampliatus ¢, June 1900.
5 * - 9, Jan. 1899.
6. » (Lopholycus) zonatus 2, Jan. 1899.
G » (Lopholycus) huagi ¢ ,, 7
8. ” ” ” 9 ” ”
9. » (Chlamydolycus) subtrabeatus g, Jan. 1899.
10. ” ” ” 2 ” 9
ll. Lycocerus mimicus g, Nov. 1899.
W2e Fe - 9, Jan. 1899.
13, Prionocerus dimidiatus, Jan. 1899.
14. Pecilomorpha fasciaticollis, Nov. 1900.
15. Peploptera zambesiana, Jan. 1899.
16. Diacantha conifera 9, Nov. 1898.
<5 5 ¢, Jan. 1899.
18. Aphodius holubi, Feb. 1899.
19. Hutrapela sp., Dec. 1901. Mt. Chirinda, Gazaland.
20. Hletica rufa 3, Nov. 1898.
21. a pate Jan. 1899.
22. Mylabris palliata Ps Fe

566 Explanation of Plates,

Fic.23. Zonitis sp. Jan. 1899.
24. Blepisanis haroldi s —
25. Amphidesmus analis ,, PS
26. Philagathes letus ¢ ,, .
27. » 9, Jan. 1902.

28. ieee sp., Feb. 1900.
29. 5 nigricornis 1 Dee. 1897.

30. 55 similis, Jan. 1901.

31. Vitwmnus scenicus, var. cinnabarinus, Stal., Jan, 1899.
32. 5, miniatus, Stal., Dec. 1900.
33. Ser etka aatilatgs April 1900.

34. Lygeus furcatus a

35. Oncopeltus famelicus, var. juc site the Sept. 1900.
36. Pompilus morosus 2, Jan. 1900.
Asclepiad pollen masses are seen on the fore tarsi of both
sides.
37. Pompilus capensis 2, Jan. 1899.
38. » Gwersus 2 ,, F
39. », vindex, April 1900.
40, Cerceris orientalis, var. 9, Jan. 1899.
41. Notogonia cresus 9, April 1900.
42. Rhynchium radiale 9, Jan. 1899.

43. x rubens 9, Jan. 1900.
44, Phanomeris sp.? 9, Jan. 1901.
45, he 1) dy Feb. 1900.

46. Iphiaulax bicolor 9, Sept. 1900.

47. Bracon luctuosus 2, Aug. 1900.

48, 3 Yuctuosus ¢, Jan. 1902.

49. NXiphocerus cruciger, Jan. 1899.

50. Ilema elegans, April 1900.

51. Zygenid (gen. indet.). Mt. Chirinda, 4500 feet, Mashona-
land, Dee. 1901.

52. Neurosymploca ochreipennis, April 1900.

Figs. 53 to 58 show the mimicry of three convergent and

common Braconids by a Reduviid and a Longicorn.

53. Bracon coccineum 2, Aug. 1900.

54, 5; eS dg, Nov. 1900.
55. Iphiaulax pictus 9, Aug. 1900.
56. ss flagrator 2, Jan. 1899.

57. Oberea scutellaris, Jan. 1899.
58. Callilestes bicolor ,, 5
The remaining figures exhibit a beautiful case of mimicry
by a Reduviid of Synaposematic conspicuously marked
parasitic Hymenoptera.

Explanation of Plates. 567

Fie.59. Phanomeris dubius 2, Nov. 1900.
60. Pimpla tuberata &, April 1900.
61. A » 9, Nov. 1900.
62. Callilestes stigmatellus, Noy. 1900,

EXPLANATION OF PLATE XIX.

MULLERIAN Mimicry IN GROUPS OF SouTtH AFRICAN COLEOP-
TERA AND HEMIPTERA, ETC.

Figs, 30 to 38 twice the natural size; Figs. 53 to 59 one and a half
times natural size ; all other figures of the natural size.

All the specimens, except that shown in Fig. 16, were captured by
Guy A. K. Marshall, and all, unless otherwise stated, are from
Salisbury, Mashonaland, 5000 feet.

Figs. 1 to 16 represent a powerful group of Cantharide and the
insects convergent towards them, and having conspicuous cream,
orange or red bands on a black ground. The group gradually merges
into that shown in Figs. 17 to 29, in which the ground-colour is
orange or yellow with black bands, sometimes broken up into spots.

Fic. 1. Mylabris dicincta, Dec. 1898.

2, 9 tettensis, Nov. 1898 to Jan, 1899.
Bh i 5 Dee. 1898.

A. - tricolor, Nov. 1898 to Jan. 1899,
5. 9 oculata 9 59 rr
6. a "5 Dec. 1898.

7. Actenodia chrysomelina, Dec. 1898.

8. ie a Nov. 1898.

9, Mylabris holosericea, Dec. 1898.

10. Decatoma lunata, Nov. 1898 to Jan, 1899.
1l. Actenodia chrysomelina, Dec. 1900.

12. Decatoma lunata, var., Nov. 1898 to Jan, 1899.
13. Mylabris oculata — ,, - x i
14. Cymatura bifasciata, Dee. 1898.

15. Ceroplesis caffer, Dec. 1898.

16. Anubis mellyi, Colenso, Natal.

17. Oncopeltus famelicus, Sept. 1900.

18. Melitonoma n. sp., Dec. 1898.

19: a truncatifrons, Oct, 1901.

20. Clythra wahlbergi, Dec. 1898.

21. Melitonoma litigiosa, Jan. 1901,

Explanation of Plates,

Antipus rufus 9, Oct. 1901.

Melitonoma epistomalis, Oct. 1901.

Clythra lacordairet, Oct. 1901.

Antipus rufus 2, Feb. 1899.

Aulacophora festiva, Oct. 1901.

Cryptocephalus quinqueplagiatus, Dec. 1900.

5 varioplagiatus, Oct. 1901.

Syagrus marshalli, Sept. 1900.

Figs. 30 to 38 show a very well-defined group of small
Coleoptera in which the head and thorax are bright red
and elytra creamy-white with transverse black bands or
spots.

Platyxantha bicineta, x 2, Jan. 1899.

Gynandrophthalma posticalis ?, x 2, Feb. 1899.

Monolepta vincta?, x 2, Jan. 1899.

Crioceris coronata 4, of 9
Paralepta ornata ,, ” ”
Asbecesta ,, ” ” ”

Urodactylus sp. gy, “ 5
” ” & ” ” ”

Apoderus gentilis ,, Noy. 1898.

Figs. 39 to 42 show a convergent group of an Hemipteron
and two Coccinellids with the characteristic red or yellow
and black spotted coloration of the latter family.

Steganocerus multipunctatus, Feb. 1899.

Epilachna dregei, Nov. 1898.

re Feb. 1899.

Chilomenes lunata, Feb. 1899.

Figs. 43 to 47 represent a convergent group of red and black
coloured Hemiptera—a Reduviid mimicking Lygeids.

Reduvius sp., Jan. 1900.

Lygeus rivularis, Nov. 1899.

Graphostethus servus, Feb. 1900.

Iygexus elegans, Feb. 1900.

55 crudelis, Jan. 1900.

Another Hemipterous group showing Reduviids mimicking
Pyrrhocorids is shown in Figs. 48 to 52.

Phonoctonus nigro-fasciatus, Oct. 1897, Malvern, near Dur-
ban, Natal.

Dysdereus nigro-fasciatus, Oct. 1897, Malvern, near Durban,
Natal.

Dysdereus superstitiosus, April 1900.

* intermedius, Feb. 1900.

Phonoctonus formosus, Feb, 1900,

Kxplanation of Plates. 569

The remaining figures represent a group containing a
Loeustid and an Hemipteron mimetic of ants.
Fie. 53. Camponotus sp.? 8 x 14, Feb. 17, 1901.

54. 95 sericeus, % x 14, Feb. 17, 1901.

Boe _ cosmicus, § min. x 14, Feb. 17, 1901.
56. 53 3 | we MAS 5 ” ”
57. Megapetus atratus, x 14, Feb. 17, 1901.

58. ” ” b) ’9

oJ») ’
59. Myrmecophana ? fallax, x 14, Feb. 17, 1901.

EXPLANATION OF PLATE XX.

GROUP OF BLACK, DARK-WINGED MASHONALAND ACULEATES AND
THEIR Mrmics.

First Part oF GROUP.

The Aculeates with one exception all Fossores.

All the figures are about + of the natural size.

All the specimens were captured by Guy A. K. Marshall, at
Salisbury, Mashonaland, 5000 feet.

Fie. 1. Mutilla atropos ¢, Nov. 1899.
2. Tiphia rugosa 9, Feb. 1899.
3. Lis lachesis 2, April 1900.
4. ,, fasciatipennis 2, April 1900.
5. ” ” 2, Feb. ”
6. Scolia alaris $, April 1900.
ile r 3 Mars
8. ” eed April ”
9. » Jraterna g, Jan. ,;
10. ” ” Q, April ,,
a » cyanea g,Jan. ,,
12. 9 45 @5 Mars Ay;
13}, » affinis 6, June,
14. Salius atropos 2, Mar. ,,
15), » vindex 9, Nov. 1899.
16. » dedjax ¢, Feb. 1899 to Jan, 1900.
lie » regina 6, Feb. 1899.
18. » obscurus 9, Bec. 1899.
19. Pompilus sepulchralis 2, April 1900. ©
20. - frustratus 2, Nov. 1899,

26.

Explanation of Plates.

Orectocera (Paraphania) diabolus, Feb. 1900.
Exoprosopa umbrosa, Feb. 1900.

Orectocera (Paraphania) diabolus, Jan. 1900,
Tabanus, very near biguttatus, Dec. 1900.
Tascia homochroa, Dec. 1900.

Reduvinus, sp., April 1900.

Harpactor tristis, April 1900.

EXPLANATION OF PLATE XXI.

GROUP OF BLACK, DARK-WINGED MASHONALAND ACULEATES

CHIEFLY FossoRES, AND THEIR MIMICs, ETC,

Sreconp Part oF GROUP, CHIEFLY FOSSOREs.

All the figures are about 4 of the natural size.
All the specimens were captured by Guy A. K. Marshall, at Salis-
bury, Mashonaland, 5000 feet.

Fia.

1k
2.
3
4

18.

Sphex bohemani 9, Dec. 1899.
55 _ 3, Jan. 1900.
cyaniventris g, Nov. 1899.
pelopeiformis 3, Jan. 1900.
” ” 2 ) June ”
» wanthocerus 2, Dec. 1899. An Asclepiad pollen-
mass is seen on distal end of the left anterior leg.

Sphex umbrosus 9, May 1900.

Sceliphron chalybeum ¢, Feb. 1900.

Tachytes natalensis 3, Jan. 1900.

5 3 2, Dec. 1899.

Ammophila ludovicus 9, Feb. 1899 to Jan. 1900. Ascle-
piad pollen-masses are thickly crowded on the tarsi of the
two anterior pairs of legs.

Ammophila ludovicus g, June 1900.

A beniniensis, 9, Jan. 1900.

Ewmenes tinctor $, Dec. 1899.

” ” 22 b) ” oh)
dyschera ¢, Feb. 1900.
i » 9, Nov. 1899.
Xylocopa hottentotta 2, Jan. 1900.
carinata 2, Feb. 1899 to Jan. 1900.

”

”

Fia. 20.
SA.
22.

Kxplanation of Plates. 571

Laparus sp. , Jan. 1900.

35 ? tabidus, Feb. 1900.
Trymodera aterrima, Nov. 1898 to Jan. 1899.
Epicauta pectoralis, Nov. 1898 to Jan. 1899.

5 subcoriacea ,, - . 5
$ celestina, Feb. 1900.
53 5 or subcoriacea, Nov. 1898 to Jan. 1899.

Lytta moesta, Nov. 1898 to Jan. 1899.

The following figures represent a beautiful little group
of species of Megachile and their mimics.

Megachile nasalis 9, Nov. 1899.

3 chrysorrhea 9, Jan. 1900.

Nelioxys pusilla ¢, Dec. 1899. Probably parasitic on one
of the two former species.

Laphria sp., near flavipes, Feb. 1900,

EXPLANATION OF PLATE XXII.

GROUP OF YELLOW-TAILED, BLACK MASHONALAND ACULEATES,
ALMOST ENTIRELY DIPLOPTERA, AND THEIR Mimics. A XyYLo-
COPID AND ITs AsILID MIMIc.

All the figures are about 4 of the natural size.

All the specimens were captured by Guy A. K. Marshall, and all,
except those otherwise described, are from Salisbury, Mashonaland,
5000 feet.

real:

Scolia erythropyga &, Dec. 1899.
3 “ 9, April 1900.
Salius tamisieri ¢, Mar. 1900.
%9 ps ?, Nov. 1899.
» spectrum fg, Jan. 1900.
Rhynchiwm synagroides ¢, Feb. 1900.
Synagris abyssinica, var. 2, Umtali, 3700 feet, Dec. 1900,
% mirabilis 9, April 1899.
ss emarginata 9, Mar. 1900.
93 analis 2, Umtali, 3700 feet, Dec. 1900,
a » &, Dec. 1899.
- xanthura 2, Jan. 1900.
7 emarginate ¢, April 1900,

Or
~I
bo

Fia. 14.

Explanation of Plates.

Euwmenes dyschera &, Jan. 1900.
Trochilium sp. , Umtali, 3700 feet, Dec. 1900.
” ” ”
Bombs ylius so, nov., Feb. 1900.
Silvius pertusus, Nov. 1898.
The two remaining figures illustrate the mimicry of a large
Asilid fly for a common type of African Xylocopid bee.
Xylocopa flavo-rufa 2, Oct. 1899.
Hyperechia marshalli ¢, Feb. 1899. Mimic of the above.

EXPLANATION OF PLATE XXIII.

MASHONALAND ACULEATES AND THEIR MIMIcs.

All the figures are about ,%, of the natural size.

All the specimens were captured by Guy A. K. Marshall at Salis-
bury, Mashonaland, 5000 feet.

Figs. 1 to 11 are a group of small insects wath black head and
thorax and yellow or fulvous abdomen.

aes A

1)

Elis aureola 2, April 1900.
Philanthus fuscipennis 2, Feb. 1900.
a diadema 92, Oct. 1899.

‘5 Bs 3, Jan. 1900.

4 bucephalus 3, Feb. 1900.
Rhynchiwm radiale 2, Jan. 1899,
rubens, var. 9, Jan. 1899.

» _ var. QO, Feb. 1900,

Spheredes rufiventris 9, April 1899.

Chalcis bicolor ¢, April 1900.

Athalia bicolor, Jan. 1900.

The two following figures represent outlying members of
the group represented in Figs. 14 to 19, the colour of the
head and thorax being of a darker tint.

Xylocopa lateritia 9, Jan. 1900.

» flavo-rufa 9, Oct. 1899.

Figs. 14 to 19 are black, dark-winged insects with yellowish
head and thorax.

Podalirius acraénsis 9, May 1899.

Anthophora basalis? 9, Oct. 1899.

Xylocopa olivacea 2, June 1900.

55 modesta 9 ,,

”

Fia. 18.
19.

34,

40.
41.

Huplanation of Plates. 573

Elis cxlebs 2, Dee. 16,°1900.

Melittia sp. 5 » Mimic of the above.

Figs. 20 to 29 and 31 are insects with dark-blue or blackish
wings and yellow heads. Where the anterior yellow is
most developed (as in Fig. 25) the group merges into the
preceding one; where it is least, the convergence is
towards the blue-winged group shown on Plates XX
and XXI.

Pompilus marshalli 2, Oct. 1899.

s dichrous ¢, Jan. 1900.
. » , Oct. 1899.
55 lasciwus 9, June 1900.
s anticus 2, Jan. 1900.

Salius tamisiert 9, Feb. 1899.

Tachysphex fluctuatus 2, June 1900,

Bromophila caffra, Dec. 1899.

Jonthodes sculptilis, Nov. 1898 to Jan. 1899.

EHletica rufa g, Nov. 1898 to Jan. 1899.

Figs. 30, 32, and 33 are blue- or black-winged insects with
yellow-banded legs which are very conspicuous during
flight.

Litopus dispar, Feb. 1899.

Iphiaulax ruber 2, Oct. 1900.

Osprynchotus flavipes 9, Jan. 1900.

Pirates xneicollis, Dec. 1900.

Figs. 34 to 36 are black insects with narrow yellow bands
and clear wings.

Pompilus festivus 9, Jan. 1899.

Myzine capitata § - -s

Metopius discolor 3 ,, 3

Two flies with their model a Megachile bee.

Megachile apiformis ¢, Jan. 1899.

Eumerus sp. nov. ? - -

Laxenecera mollis, Mar. 1900. .

The following figures represent a Syrphid fly which closely
mimics a common social wasp.

Polistes marginalis 9 , Dec. 1898.

Ceria gambiana 7 ae,

(Sii

=I

INDEX
TO THE PRECEDING PAPER.

MAMMALIA.

Cebus, sp., 406

Cercopithecus callitrichus, F, Cuv., 406

pygerythrus, F, Cuv., 379,
386, 387, 402, 531

Crossarchus fasciatus, Desm., 378

Herpestes galera, Erxl. (mungoose), 376,
378

Papio porcarius, Bodd, (baboon), 346,
380-91, 397, 398

AVES.
Aquila wahlbergi, Sund., 352
Artamus fuscus, Vieill., 360
Asio capensis, Sm., 378
Astur polyzonoides, Sin., 352
Asturinula monogramnrica, Tem., 349,
352
Bradyornis mariquensis, Sm., 351, 357
Bubo maculosus, Vieill., 350, 352
Buchanga assimilis, Sharpe, 351, 357,
358, 378
Bucorax caffer, Verr., 346, 347, 378
Buteo jakal, Daud., 352
Campothera bennetti, Sm., 351
Cauprimulgus rufigena, Sm., 351
Carpococcyx radiatus, Tem., 406
Cerchneis amurensis, Rad., 350, 351
naumanni, Fleisch , 340-47,
350
rupicoloides, Sm., 340-42,
345-47, 350, 352, 378
Ciconia abdimii, Licht., 350
Circaétus pectoralis, Sm., 352
Coceystes caffer, Licht., 357
A glandarius, i., 851
Coracias caudata, L., 349, 351, 356
garrula, L., 349
olivaceiceps, Sharpe, 349, 352
,, spataulata, Trim., 349, 354
Cosmetorius vexillarius, Gould., 378
Crateropus kirkti, Sharpe, 351
Crecopsis egregia, Pet., 351
Cuculus gularis, Steph., 349
Dicrurus ater, Herm. (king-crow), 359,
362, 363
Eurystomus afer, Lath., 354

29

»)

99

be)

2)

Falco subbuteo, L., 349, 351, 357, 378
Fringillaria tahapisi, Sm., 351
Gallinula chloropus, L., 378
Geocichla litsitsirupa, Sm., 351
Glaucidium perlatum, Vieill., 352
Graucalus pectoralis, Jard., 351
Hailcyon chelicutensis, Stanl., 357
,, pallidiventris, Cab., 351
Herodias lucidus, Raf., 350, 378
Trrisor erythrorrhynchus, Lath., 349,
351, 352, 378
Laniarius guttatus, Hart, 351
Lophoceros lewevmelas, Licht,, 351
Macronyx capensis, L., 349, 351, 352
Melierax polyzonus, Riipp., 352
Melittophagus pusillus, P. Miill., 349
Merops upiaster, L., 354
natalensis, Reichenb., 349, 350,
352
philippinus, L., 360, 363
», swinhoci, Hume, 361
Microhierax cerulescens, L. (M. cutol-
mus, Hodgs. ), 363-65
Sringillarius, Drap., 364
Monticola angolensis, Sousa, 351
Nettapus auritus, Bodd., 378
Numida coronata, G. R. Gray, 350
Oriolus larvatus, Licht., 352
», notatus, Pet., 351
Pachyprora molitor, H. and K., 357
Pratincola torquata, L., 351, 358
Prionops talacoma, Sm., 351
Rhinopomastus cyanomelas, Vieill., 351,
352, 378
Saxicola pileata, Gmel., 351, 378
Terpsiphone perspicillata, Swains., 354,
357
Thamnolxa cinnamomeiventris, Lafr.,
351
Turdus libonyanus, Sm., 352
Turtur capicola, Sund., 378
Upupa africana, Bech., 351
Zosterops, sp., 407

2?

29

REPTILIA.

Amphibolurus, sp., 407
Bitis arietans, L., 3970

Index.

Cistudo ewropea, Schn., 407
Coluber xsculapii, Host, 407
Lacerta muralis, Laur., 407

» viridis, Laur., 399, 407
Zonurus, sp., 407

AMPHIBIA.

Triton alpestris, Laur., 408
punctatus, Daud., 407

29

PISCES.

Chetodon plebeius, Gmel., 375

ARACHNIDA.,

Agelena labyrinthica, L., 326, 327, 408 |

Amaurobius ferox, Walck., 326, 408

Epeira diadema, L., 325, 327, 408

Gasteracantha ornata, Thor., 384

Lycosa, sp., 325, 326

Nephila, sp., 325

Nephilengys malabarensis, Walck., 319

Prosthesimaalbomaculata,O. P.Cambr.,
511

Solpuga marshalli, Poc., 350, 353

Tegenaria domestica, Walck., 326, 527,
408

Titus lugens, O. P. Cambr., 511

Thomisus, sp., 328

INSECTA.
RHOPALOCERA.
TIthomiine.
Melinxa, sp., 468

Danwine.

Amauris dominicanus, Trim., 486, 503,
507, 508
echeria, Boisd.

Offered to Mantide and
spiders, 297, 298, 313,
321, 323, 324

Effect upon human taste,
414

As model for mimicry,
469, 484-87, 503-5, 507

echerid, var. albimaculata,
Butl., 485, 487, 490
niavius, L., 486
» ochiea, Boisd., 466, 490, 503
Crastia core, Cr., 360
Hamadryas, sp., 467, 468
Linnas bataviana, Moore, 474

22

be)

57

13)

Limnas chrysippus, L.

Offered to Mantide, 298, 300,
301, 306, 311-13, 315, 317

Offered to spiders, 319, 320,
323

Insect foes of larve and
imagines, 329, 338

Offered to captive
341-48

Probably injured by wild
birds, 366, 367

birds,

Offered to mungoose and
baboons, 3877, 380, 382,
388

Etfect upon human taste,
413, 414

As model for mimicry, ete.,
468-84, 490, 494, 496,
499, 501

Courtship of, 538, 539

chrysippus, var. alcippoides,
Moore, 468, 470, 473, 474,
477, 478

chrysippus, var. alcippus, Cr.,
474, 476, 478-80, 484

chrysippus, var. dorippus,
Klug, 473, 474

chrysippus, var. klugii, Butl.,
442, 470, 473-80, 482-84,
494, 499

Salatura, sp., 476

Tirumala limniace, Cr., 332

petiverana., Doubl. Hew., 507

be)

Satyrine.

Cenonympha pamphilus, L., 440
Epinephele janira, L., 372

5 lycaon, Kithn., 554
E. tithonus, L., 372
Melanarge galathea, L., 558
Melanitis leda, L., 416.

a zitenius, Hbst., 363
Mycalesis perseus, F., 365

>», safitza, Hew., 302, 313
Psewdonympha extensa, Butl., 551

3% vigilans, Trim., 553

Satyrus semele, L., 372

Elymniine.

Elymiias, sp., 489

Nynphaline.

Anxa, sp., 373

Argynnis cybele, F., 553

Atella phalanthi, Dr., 340, 343, 344,
357, 367, 384

576

Athyma, sp., 468
Byblia goetzius-acheloia, Wallg., 322,
379
», tithyia, Dr., 337, 340, 342-45,
377, 380-85, 388
Cethosia, sp., 502
Charaxes achwmenes, Feld., 505
», athamas, Dr., 362
» guderiana, Dew., 505
+ psaphon, Westw., 360
ie saturnus, Butl., 505
,, sehretbert, Godt., 365
., waranes, Cr., 299, 375
», whytet, Butl., 505
is xviphares, Cr., 505
Cyrestis thyodamas, Boisd., 362
Ewphexdra, sp., 499
Euralia anthedon, Doubl., 486, 492
», deceptor, Trim., 503
», dubia, Beauv., 492
., mima, Trim., 485, 487, 490-
92, 503, 504

», wahlbergi, Wallgr., 486, 491, |

492, 503, 508
Hurytela hiarbas, Dr., 299, 300, 320
Hananumida dedalus, ¥., 311, 343
Hypolimnas bolina, L., 502
var. nerina, F., 502

9 9

es misippus, L.
Offered to insect-eaters,
299-301, 313, 347,

348, 384, 385
Mimetic resemblance of,

468, 470, 479-84,
490, 501
50 misippus, var. inaria, L.,

384, 468, 470, 477,
479-81, 483, 484
Junonia cebrene, Trim., 310, 311, 340,
342-44, 357, 380
,, clelia, Cr., 300, 301, 348
», hierta, F., 336
», orithyia, L., 365
Neptidopsis, sp., 468
Neptis agatha, Cr.
Offered to insect-eaters, 299,
300, 3802, 313, 382, 384,
386, 388
As Miillerian mimic, etc., 466,
467, 496, 497, 505
5, tmitans, Obthr., 468
», lactaria, Butl., 467
5, venilia, Cr., 467
Precis actia, Dist., 416, 423
, antilope, Feisth. (= simia,
Wallg.), 379, 414, 416, 418-
21, 423, 424, 427-31, 438,
456, 458

Index.

Precis archesia, Cr. (= pelasgis, Godt.’
Offered to insect-eaters, 340,
341, 344, 379, 384
Seasonal phases of, 415, 416,
418-20, 423, 424, 427-29,
431, 438, 450, 453-55, 458
» artaxia, Hew. (= nachtigalii,
Dew.), 416, 422, 4238, 430,
439-41, 455, 456, 457
» ceryne, Boisd.(=tukuoa, Waller.),
416, 424, 425, 430, 435
», cuama, Hew., 418-20
,, elgiva, Hew., 338, 421, 423
», natalica, Feld., 421, 423
,, octavia, Cr. (= amestris, Dr.),
415, 452
» sesamus, Trim.
Staud. )
Offered to Mantide and
spiders, 311, 321
Offered to kestrels, 340-43
Offered to captive mam-
mals, 377, 379, 380, 382-
85, 387-89
Seasonal phases of, 415-31,
435-39, 443-58, 455, 456,
458
Gregarious instinct of, 460,
462
», btriment, Butl., 419
», tugela, Trim., 416, 421, 423
Protogonius, sp., 373
Protogoniomorpha, sp., 502
Pseudacrxa expansa, Butl., 503
5 tarquinia, Trim., 487
a4 trimenit, Butl., 504
Symphedra dirtea, F., 365
Vanessa atalanta, L., 368
a cardut, L., 300, 311,
380
5, 40, L., 442, 452, 460, 461
x kaschmirensis, Kolb., 359
5, polychloros, L., 461
5 urtice, L., 831, 442, 452, 461

(= natalensis,

377,

Heliconine.

Heliconius eucrate, Hiibn., 366
6 sp., 468

Acreine,
Acrea acara, Hew., 418, 4383, 434,

504

,, admatha, Hew., 330

;, anacreon, Trim., 337, 413

», a@nemosa, Hew., 324, 342, 382,
388, 413, 433, 493

», atolmis, Westw., 433

Index.

Acrexa bonasia, F., 328

be)

”

9

cabira, Hopff.

Offered to insect-eaters, 299,
301, 302, 308, 313, 314,
316, 324, 348

Ichneumonid parasites of,
337

Effect upon human _ taste,
413

Summer and winter forms
compared, 433

caldarena, Hew.

Offered to Mantide, 308-11,
314, 315, 319

Attacked by Asilid, 333

Offered to kestrels, 340, 342-
44

Offered to mammals, 376,
377, 379, 382, 388

As member of Miillerian
group and as model, 492—
94

Observations on sac of ? ,539

doubledayi, Guér., 468, 469

9 Saal CUGUILO.
Westw.

Offered to Mantidx, 311, 312,
315

Offered to kestrels, 340, 344

Offered to mammals, 376,377,
380, 382, 383, 388

Summer and winter forms
compared, 433, 434

As member of Miillerian
group and as model, 492,
491, 496

encedon, Li.

Offered to Mantide, 299, 300,
302, 303, 305-8, 313, 314,
316

Offered to hornbill, 347

Effect upon human taste, 413

Summer and winter forms
compared, 433

As Miillerian mimie and as
model, 468-72, 479, 480,
483, 484, 490, 495

Courtship of, 540

encedon, var. alcippina, Aur.,

480, 484

encedon, var. daira, Godm.,

Salv., 479, 480, 483, 484

encedon, var. lycta, F., 413, 479,

horia, L.

Offered to Mantide and
spiders, 297, 298, 313, 316,
318, 320-24

Dual

Acrea horta, L.

>

29

Dipterous parasites of, 337
Effeet upon human senses,
414
Summer and winter forms of,
compared, 433
Observations on sie of 9 and
courtship, 539, 540
induna, Trim., 308, 314, 315,
494
natalica, Boisd., 300, 309, 313-
15, 339, 384, 413, 493
neobule, Doubl., 306-8, 314,
$15, 330, 433, 539
nohara, Boisd., 324, 413, 494
» var. halali, Mshl.
Offered to Mantide, 308-10,
314, 315
Offered to kestrels, 340, 342,
344
Offered to mammals, 379,
380, 382, 384, 388
Summer and winter forms of,
compared, 433, 434
As member of Miillerian
group, 492
Observations on sac of ? , 539
petrea, Boisd.
Offered to insect-eaters, 299,
313, 347, 348
Rarity in Natal in 1897, 325
Summer and winter forms of,
compared, 433-35
As Miillerian mimic, 470,
490
Courtship of, 540
rahira, Boisd., 341, 344, 383,
388, 492, 493
serena, F., var. buxtoni, Hew.
Offered to Mantide, 299, 300,
302, 303, 805, 307, 313,
314, 316
Offered to spiders, 321-24
Offered to hornbill, 347
Effect upon human taste,
413
Summer and winter forms of,
compared, 453
As Miillerian mimic, 468-70
viole, F., 316, 489
violarum, Boisd., 321, 324
PAVIA NCLSCTIIG

"Hew., 433, 434, 492-94

Actinote thalia, L., 366

Pardopsis punctatissima, Boisd., 305,
307, 308, 314, 315

Planema aganice, Hew., 324, 348,
413, 466, 490

TRANS. ENT. SOC. LOND. 1902.—PART II. (NOV.) 38

578

Planema esebria, Hew.
Offered to insect-eaters,
392, 348
Effect upon human taste,
413
As Miillerian mimic and
as model, 466, 468-70,
472, 479
gea, ¥., 488

Ie)
Lycxenide.
Alxna amazoula, Boisd., 800, 319,
315, 414, 497-99
», wyasse, Hew., 496, 497, 499
Axiocerces amanga, Westw., 555
x harpax, F., 311, 341
Castalius calice, Hopff., 497
a melena, Trim., 497
Catochrysops contracta, Butl., 335
gigantea, Trim., 494
hypoleuca, Butl., 494
mashuna, Trim.,
499
osiris, Hopff., 310
parsimon, F., 555
peculiaris, Rghfr.,
495, 499
Citrinophila vas Hew.,
Deloneura, sp., 497
Deudorix antalus, Hopff., 374
Durbania, sp., 824
Ewumexus, sp., 500
Hypolycena philippus, F., 555
Tolaus, sp., 497
Lachnocnema, sp., 498
Lampides betica, L., 333
Larinopoda lircea, Hew., 499
tera, Hew., 499
Liptena libyssa, "Hew., "499
» wndularis, Hew., 499
Lycena icarus, Rott., 553
Lycenesthes, sp., 497
Mimacrexa dohertyi, Roths., 480, 483,
484, 493, 499
a marshalli, Trim., 470-72,
480, 482-84, 493, 499
Myrina ficedula, Trim., 310, 375
Pentila abraxas, Doubl., 499
» phidia, Hew., 499
,, tropicalis, Boisd.,

494,
499

Spindasis natalensis, Doubl., 310, 374
Talicada, sp., 500
Tarucus plinius, L., 357, 358, 497
Telipna bimaculata, Plotz, 499

5 sanguinea, Plotz, 499
Teriomima, sp., 324, 472
Thecla iarbas, ¥., 374

494, 497, |

Index.

Thecla W-album, Knoch, 374
Uranothauma pogget, Dew., 555
Zeritis taikosama, Trim., 389
Zizra gaika, Trim., 333

Pierine.

Belenois gidica, Godt., 435
», mesentina, Cr.
Offered to Mantide, 311
Eaten by wild Mantis and
spider, 316, 328
Offered to mammals, 376,
381, 385, 386, 389
Discussion of mode of pro-
tection, 435
severina, Cr.
Offered to Mantide, 298,
305, 310, 311, 313, 314
Offered to kestrel, 342
Offered to mammals, 376,
380-82, 384, 386
Effect upon human taste,
413
Discussion of mode of pro-
tection, 435
Ph thysa, Hopft., 485, 506
Catophaga, sp., 360
Catopsilia florella, F., 311, 341, 357,
380, 383
Colias edusa, L., 366
», electra, L., 882
Delias eucharis, Dr., 316, 489
5 pandemia, Wallace, 504
», descombest, Boisd., 316
Eronia cleodora, Hiibn., 357
Hebomoia, sp., 362
Herpenia eriphia, Godt.,
388, 460, 506
Mylothris agathina, Cr.
Offered to Mantide, 300
> 6g amoMals yee.
377, 382, 384, 386, 388
Effect on human taste,
414
Specially protected and
mimicked, 506
riippelli, Koch, 506
as trimenia, Butl., 506
Nepheronia argia, F., 321, 506
Pieris brassicx, L., 338
», rape, L., 464
Pinacopteryx pigea, Boisd., 539, 540
Pontia hellica, L., 321, 339, 433, 506
Prioneris, sp. 362
Synchloé glauconome, Klug., 333
Teracolus achine, Cr., 304, 305, 314,
344, 380, 382, 386, 388

9

385, 386,

99

Teracolus eris, Klug., 460, 506
aC omphale, Godt., 303, 318,
370, 376, 386

”

vt, Swinh., 328
Terias brigitta, Cr.
Offered to Mantidee and

39
29

2)

Papilio antheus, Cr., 507

9
99
bi)

29

Index.

phlegyas, Butl., 386

spiders, 303, 310, 321
Offered to lizard, 339
», kestrel, 341, 342
376,

2?
5 », Mammals,

877, 383
Cryptic larva of, 506
hecabe, L., 362
regularis, Butl., 553
senegalensis, Boisd., 310, 376,
377, 386

Papilionine.

arcturus, Westw., 359

aristolochix, F., 489

caunus, Westw., 365

cenea, Stoll, 469, 482, 485-87,
491, 507, 508

cenea, var. hippocoon, ¥., 486

constantinus, Ward, 324

coon, F., 482

corinneus, Bert., 842, 344, 346,
358, 380, 385, 388, 507

cynorta, F., 488, 489

cyprexofila, Butl., 488, 489

demodocus, Esp.

Offered to Mantide and
spiders, 298-301, 313, 320,
321

Discussion of results, 324, 325
Offered to captive birds, 341,
345, 348
Eaten by wild cuckoo, 357
Injuries probably caused by
birds, 373
Protection of larva of, 507
echerioides, Trim., 487, 488
ertthonius, Cr., 862, 365
euphranor, Trim., 320, 324
gallienus, Dist., 488
jacksoni, E. M. Sharpe, 487, 488
leontdas, F., 507
leonidas-brasidas,
301, 320, 348, 485
507
lyzxus, Doubl., 320, 247, 348
macareus, Godt., 362
merope, Cr., 486
var. hippocoon, F., 486,
note

Feld., 300,
, 486, 506,

2?

nomius, Esp., 360

579

Papilio ophidicephatus, Obthr., 320,
324

», policenes, Cr., 507

», polyctor, Boisd., 359

» sarpedon, L., 362, 364

», wenocles, Doubl., 362

», xenobia, F., 488, 489

Hesperidx.
Abantis tettensis, Hopff., 495
Baoris netopha, Hew., 495, 496 _
Cyclopides willemi, Wallgr., 496°
Hasora alexis, F., 316
Hesperia spio, L., 310
Kedestes macomo, Trim., 495
Parosmodes icteria, Mab., 310
Rhopalocampta forestan, Cr., 348, 376,
377, 380, 385, 388,
496
5 pisistratus,
388
Sarangesa eliminata, Holl.,

F., 385,

357, 422

HETEROCERA.

Abraxas grossulariata, L., 825-27, 330,
338, 405-11
Acherontia atropos, Li., 401, 402, 404
Aletis, sp., 482, 499
Alypia octomaculata, F., 411
Callioratis bellatriz, ’Dalm., 359
Cherocampa elpenor, Iie 399, 401
osiris, Dalm. .) 397
Cirina similis, Dist., 350
Diacrisia mae alosa, Cr, +) O08
Eqybolis vaillantina, Stoll, 300, 315,
316
Euchelia jacobex, L., 338, 407, 410
Halia wavaria, L., 325
Halias prasinana, L., 404
Hybernia aurantiaria, Esper., 465
as defoliaria, Glerck, 465
Llema elegans, Butl., 517
Lsharta pandemia, Roths., 504
Melittia, sp., 530, 531
Neurosymploca ochreipennis,
516, 517
Nyctemera leuconoé, Hopft., 370, 466,
468, 497
Pericopis, sp. 468
Letovia dichroaria, H. 8., 497,
Porthesia aurifiua, F., 338
Protoparce convolvuli, L., 380, 388
Pscudaphelia apollinaris, Boisd., 350
Pseudohazis, sp., 411
Tascia homochroa, Holl., 526, 527
Trochilium, sp., 529
Zygenid (2 gen. et sp.), 517

Butl.,

498

COLEOPTERA.
Cicindeclidex.

Mantichora herculeana, Klug., 510
Myrmecoptera bilunata, Dohrn., 512
invicta, Pér., 512

”

Ms marshalli, Pér., 512

Pe polyhirmoides, Bates, 511
a var, mash-

b]
Unt, ’Pér., Apibets sls)
Tricondyla, sp., 514

Carabide.

Anthia massilicata, Guer., 383, 392,
508

mazxillosa, Fab., 509

nimrod, Ol., 509

‘ omoplata, Lequ., var. mellyt, |
Bréme, 509
5, pachyoma, Chd., 349

petersi, Klug., 508, 509
serguttata, F., 509
4 thoracica, F., 380, 383, 392,
508, 509
Arsinoé fraterna, Pér., 522
Atractonota mulsanti, Perr., 512, 513,
515
Carabus awratus, L., 330, 408
Chlenius cylindricollis, Dej., 382, 390

Eccoptoptera cupricollis, Chd., 511,
512
Graphipterus antiokanus, Pér., 512,
514
ee bilineatus, Boh., 343
sa lineolatus, Boh., 343
ie mashunus, Pér., 380,
390

tibialis, Chd., 380, 390

a wahlbergi, Boh., 348

Piezia marshalli, Pér., 350, 380, 390,
511, 513

mashuna, Pér., 511

selowsi, Pér., 342, 380, 381,
390, 512-14

Polyhirma xnigma, Dohrn., 348, 380,
388, 390, 512, 513

bennettii, Mshl., 511, 547

bilunata, Boh., 512

oi)

9)

- boucardi, Chd., 380, 390,
514, 515

re mactlenta, Ol., 511, 548

5 notata, Perr., 511

‘ rutata, Per., 511

semisuturata, Chd.,
380, 390, 511, 513, 5

Scarites, sp., 350, 353

Thyreopierus filavesignatus, Dej.,

99

Index.

| Cetoniidex.

| Clinteria infuscata, G. and P., 343,
| 347, 380, 391
Celorrhina loricata, Jans., 381, 391
Nyassinius lugubris, Westw., 349
Oxythyrea dysenterica, Boh., 381, 391
Pachnoda flaviventris, G. and P., 380,
391
Fes rufa, de G., 380, 391
Protxtia amakosa, Boh., 380, 381, 391
» mandarinea, Web., 335
Trymodera aterrima, Gerst., 526

Copridx.
Aphodius holubi, Dohrn., 517
| Gymnopleurus fastiditus, Har., 349
smaragdinus, Fahr.,
380, 392
Oniticellus militaris, Castn., 380, 392
Onitis alexis, Klug., 331, 343, 346, 349,
380, 381, 392
innuus, F., 380, 391

2)

Onthophagus gazella, F., 344, 347, 351,
380, 392
an sp., 349

Scarabeus femoralis, Kirby, 395

Buprestidex,
Agrilus, sp., 349
Amblysterna vittipcnnis,
346, 379
Anthaxia, sp., 349
Buprestis flavomaculata, F., 335
Psiloptera chalcophoroides, Pér., 350,
379, 380, 390
a valens, Pér., MS., 340
Sphenoptera disjuncta, Boh., 349
Sternocera funebris, Boh., 380, 390

Boh., 342,

Malacodermata.
Lycocerus mimicus, Bourg., 517, 549
Lycus ampliatus, F., 344, 380, 517
constrictus, Fahr., 344, 380, 517

Le}

5, haagi, Bourg., 517

5 rostratus, L., 340, 844. 380, 517
,, subtrabeatus, Bourg., 380, 517
>, «zonatus, Fahr., 517

Cantharide.
Actenodia chrysomelina, Krichs., 518,
519
Decatoma lunata, Pall., 380, 391, 518
Eletica rufa, F., 344, 347, 380, 391,
516, 517, 531
Epicauta celestina, Haag., 526, 527
pectoralis, Gerst., 526, 527
subcoriacea, Makl., 526, 527

.?

2?

Index.

Lytta mesta, Pér., 526, 527
Mylabris dicincta, Bert., 382, 391, 518

i$ holosericca, Klug., 348, 347,

518, 519
si oculata, Thunb., 349, 352,
518
5 palliata, Mars., 344, 347,
380, 391, 517
an tettensis, Gerst., 380, 382,
391, 518
3 tricolor, Gerst., 518
Zonitis, sp., 844, 347, 380, 391, 516,
lal 7
Tenebrionide.
Anomalipus plebeius, Pér., 343, 346, |
380, 381, 391
Dichtha inflata, Gerst., 342, 346, 379

Eutrapela, sp., 517
Hoplonyx, sp., 349
Lagria, sp., 334, 342, 347
Micrantereus carinatus, Pér., 380, 391
Praogena festira, Mikl., 380, 392
i splendens, Mikl., 379
Psammodes scabratus, Gerst., 350, 382,
391

5 ventricosus, Faihr., 850

Zophosis, sp., 351

Longicornia.

Amphidesmus analis, O1., 517
Anubis mellyi, White, 518, 519
Blepisanis haroldi, Fahr., 340, 345,
383, 388, 391, 517
Ceroplesis caffer, \hunh., 518, 519
fallax, Pér., 342, 346, 381,
389, 392
Cymatura bifasciata, Gerst., 518, 519
Daphisia, sp., 395
Dyenmonus apicalis, Fihy., 616, 517
Hesperophanes amicus, White, 350
Hippopsicon, sp., 522
Hyllisia, sp., 522
Jonthodes sculptilis, White, 531
Litopus dispar, Thoms., 532
Nitocris ¢nigricornis, Ol., 517
similis, Gah., 517
50 Spe old
Oberea scutellaris, Gerst., 533
Phantasis giyantea, Guér., 350
Philagathes letus, Thoms., 516-18
Tragiscoschema wahlbergi, Fahy., 382,
391

”

”

Rhynchophora.

Alcides hemopterus, Boh., 351
Apoderus gentilis, Pér., 521

581

Brachycerus apterus, L,, 524

brevicostatus, Fahr., 380,
382, 390

Cleonus, sp., 349

Diurus furciliatus, Gyl., 525

Eremnus, sp., 344, 346, 349

Hipporrhinus bohemanii, Fahr., 350

Oosomus, sp., 344, 346, 351

Polycleis decorus, Pér., 351

equestris, Boh., var., 380, 390
ee longicornis, Fahr., 380, 390

Symprexiorrhynchus, sp., 849

?

9”

Phytophaga.

Antipus rufus, de G., 519

Ashecesta ornata, Jac., 520

Aspidomorpha punctata, F., 351, 352

Aulacophora festiva, Gerst., 519, 520

Clythra lacordairei, Jac., 519

wahlbergi, Lac., 842, 381, 891,
518

Crioceris coronata, Baly, 520, 521

Cryptocephalus quinqueplagiatus, Jac.,
519, 520

varioplagiatus,
519, 520

Diacantha conifera, Fairm., 344, 347,
380, 391, 516-18

Gynandrophthalmna Lef.,
520

Macrocoma aureovillosa, Marsh, 350,
352

Malacosoma discoidalis, Jac., 380, 391

Melitonoma epistomatis, Lac., 519

litigiosa, Lae., 519

truncatifrons, Lac., 519

ae sp. nov., 519

Monolepta ? vincta, Gerst., 520

Paralepta ornata, Jac., 520

Peploptera anchoralis, Jac., 380, 389,
391

zambesiana, Pér., 880, 389,
391, 517, 518

Plagiodera thoracica, F., 380, 391

Platypria mashuna, Pér., 349, 352

Platyxantha bicincta, Jac., 520

Pecilomorpha fasciaticollis, Jac., 517,
518

Syagrus marshalli, Jae., 519

,, puncticollis, Lef., 349, 252
Timarcha, syp., 524

93

5 Jac.,

? posticalis,

29

”

9

Other families.

Adoretus flaveolus, Gerst., 380, 390
Allochotes, sp., 394
Anomala, sp., 349, 380, 390

582

Chilomenes lunata, F., 380, 391, 520
Clerus, sp., 344, 347
Corymbites virens, Schr., 331
Dytiscus dimidiatus, Berg., 330
5 marginatus [marginalis], L.,
330
Enewustes, sp., 522
Epilachna dregei, Muls.,
380, 381, 391, 520
Graptoclerus, sp., 512, 515
Ffeteronychus licas, Klug.,
Hister caffer, Erichs., 331
Hydaticus, sp., 350
Lemidia, sp., 895
Lucanus cervus, L., 408
Pentodon nireus, Burm., 349
Prionocerus dimidiatus, Gerst.,
344, 345, 347, 380, 391, 516-18
Trochalus, sp.,351
Vrodactylus, sp., 520, 521

342, 347,

350

341,

HYMENOPTERA.

Ammophila beniniensis, P. de B., 525
ie hirsuta, Kirby, 464
ludovicus, Sm., 525
Anthophora ? basalis, Sm., 530
Apis florea, F., 335
Athalia bicolor, S auss.,
Belenogaster, sp., 477,
Bembex, sp., 532
Bracon coceinewm, Brul., 533
», luctwosus, Brul., 517
», ?luctuosus, Brul., 517
Camponotus cosmicus, Sm., 535
3 sericeus, F., 5385
Carebara, sp., 350
Cerceris orie nialis, Sm. var., 517
Chalcis albicrus, Klug., 338, note
», bicolor, Bing., 530, 544
», cuplea, Hope, 338, note
>, semirufa, Walk., 545
Celioxys pusilla, Gerst, 530
Elis aureola, Klag., 530
3, celebs, Sich., 530, 531
», fasciatipennis, Sm., 525, 528
», lachesis, Sauss., 525
Eumenes dyschera, Sauss., 525, 529
5 tinctor, Sauss., 525, 526
Halictus, sp., 3384
Ichneumon, sp., 338
Iphiaulax bicolor, Brul., 517
53 flagrator, Gerst., 533

530

537

95 pictus, Brul., 533

is ruber, Bing., 531, 545
Megachile apiformis, Sm., 534

5 chrysorrhea, Gerst., 530

ae nasalis, Sm., 530

| | Seeléphron chalybewm, Sm.,

Tndex.

Melipona apicalis, Sm., 334, 336
Metopius discolor, 'Tosq., 532
Microgaster, sp., 3388
Mutilla atropos, Sm., 525
>, cepheus, Sm., 512
Pe horrida, Sm., 512
», 2 lewcopyga, Klug., 512
», purpurata, Sm., 512
» sycorax, Sm., 512
,, tettensis, Gerst., 512
Myzine capitata, Sm., 532
Notogonia cresus, Sm., 517, 518
Osprynchotus flavipes, Brul., 532
Pelopeus spirifer, F., 532
Phanomeris dubius, Bing.,
546
Spy old
Philanthus bucephalus, Sm.,
5 diadema, ¥., 530
Suscipennis, Guér., 530
Pimpla tuberata, Tosq. , 932, 538
Podalirius acraénsis, F., 580, 531
Polistes gallica, L., 335. :
> marginalis, F., 534
> variatus, Cress., 536
Pompilus anticus, Klug., 531

532, 533,

530

Ae capensis, Dahl., 517
6 collaris, Sauss., 547
es dichrous, Brul., 531
ue diversus, Dahl., 517
a Jestivus, Klug., 532
or Srustratus, Sm., 525
ss lascivus, Cam., 531

marshalli, Bing., 531, 547

9
54 morosus, Sm., 517
55 sepulchralis, Sm., 525
Me vindex, Sm., 517
| | Rhynchium radiale, Sauss., 517, 530
55 rubens, Sauss., bilge 530
“ synagroides, Sauss., 529

Satis atropos, Sm., 525

», dedjax, Guér., 525, 531

;, obscurus, Sm., 525

» regina, Sauss., 525

> spectrum, Sm., 529

» tamisieri, Guér., 529, 531

vindex, Sm. sy Bs BAS

525
| Scolia affinis, Guér., 525

;, alaris, Sauss. , 525, 527-29

», cyanea, Lepel., 525, 526

>, erythropyga, Burm., 529

» Jraterna, Sm., 525, 527
Sphecodes rufiventris, Sm., 5380

| Sphex bohemani, Dahh:, 525

», cyaniventris, Guér., 525
5, pelopeiformis, Dahl., 525
», wmbrosus, Christ., 525

Index. 583

Sphex xanthocerus, Il., 525
Synagris abyssinica, Guér., 529

WA analis, Sauss., 529

33 emarginata, Sauss., 529

ee mirabilis, Guér., 529
aanthura, Sauss., 529

Tachysphex fluctuatus, Gerst., 531
Tachytes natalensis, Sauss., 525, 526
Tiphia rugosa, Sm., 525
Vespa maculata, L., All
Vipio, sp., 546
Xylocopa carinata, Sauss., 525

» jravorufa, de G., 530, 531,

533, 534

» hottentota, Sm., 525

» lateritia, Sm., 530, 531

», modesta, Sm., 530

», olivacea, F., 530, 531

HETEROPTERA.

Anoplocnemis curvipes, F., 345, 346,
382, 383, 413

Callilestes bicolor, Dist., 5383

$3 stigmatellus, Dist., 582, 544
Cyclopetia, sp. [Cyclopelta], 317
Dysdercus cardinalis, Gerst., 543
intermedius, Dist., 538, 543

50 nigrofasciatus, St&l, 538

Ss superstitiosus, F., 538, 543

Graphostethus servus, F., 537
Harpactor tristis, Stal, 526
Lygeus crudelis, F., 537

5p elegans, Wolff, 537
Surcatus, F., 517

», rivularis, Germ., 537
Megapetus atratus, Dist., 535, 542
Myrmoplasta, sp., 542
Oncopeltus famelicus, F., 518

var. jucundus,

”

9)

99
Dall., 517
Petascelis remipes, Sign., 356, 382
Phonoctonus formosus, Dist. ; 538, 545
- nigrofasciatus, Stal, 538

Physomerus, sp., 317
Pirates xneicollis, Schaum., 532
Reduvius, sp., 526, 537
Serinetha mutillata, Gerst., 517
Steganocerus multipunctatus, Thb., 520
Vitumnus cinnabarinus, Stal, 517

- miniatus, Stal, 517

HOMOPTERA,

Pyrops, sp., 349
Tibicen nubifurca, Walk., 332

ORTHOPTERA RAPTORIA.

Creobotra urbana, F., 316

Gongylus gongyloides, L., 316, 325

Hierodula bipapilla, Serv., 317

Idolum diabolicum, Sauss., 317

Phyllocrania insignis, Westw., 304, 314

Polyspilota caffra, Westw., 298, 304,
306, 313, 314

Pseudocreobotra wahlbergi, Stal,
3, 306, 313, 314, 399

301-

Sphodromantis lineola, Burm. ,308, 310,

314, 315

ORTHOPTERA SALTATORIA.

Clonia wahlbergi, Stal, 349

Condylodera tricondyloides,
514

Myrmecophana ? fallax, Brun., 535

Phymateus morbillosus, L., 349, 353,
356, 377

Podisina frigida, Boh., 335

Westw.,

DIPTERA.

Alcimus ? stenuwrus, Lw., 333
Apoclea femoralis, Wied., 333
Bengalia, sp., 540
Bombylius, sp. noy., 529
Bromophila caffra, Macq., 531
Ceria gambiana, Saund., 534
Chrysops cecutiens, L., 333
Damatlina, sp., 334
Dasypogon diadema, F., 334-36
Dioctria atricapilla, Mg., 333

; elandica, Ia., 332

» Tufipes, deG., 333
Epitriptus arthriticus, Zlv., 335
Humerus, sp., 332

», nov.?, 534
Eutolmus ? wpicatus, Lw., 334
Exoprosopa wmbrosa, Lw., 526, 527
Exorista vulgaris, Fin., 338
Hyperechia marshallt, Aust., 5338, 541
Ss fera, v.d. Wulp, 542

ar aylocopiformis, Walk.,
542
Laparus ?tabidus, Lw., 526, 527

sp., 526, 527

Laphiri ia nr. flavipes, Wied.,

3 gibbosa, L., 335
Laxenecera, sp., 335

- mollis, Liw., 534
Lophonotus ?suillus, F., 333
Lucilia, sp., 317 -
Machimus atricapitlus, Fln., 333
Muaira, sp., 332

584

Microstylum apicale, Wied., 332
a dux, Wied., 335

Musca, sp., 317

Mydexa, sp., 334

Neoitamus cyanurus, Lw., 333

2 griseus, Wied., 335

be)
», elongistylus, Wied., 335
Orectocera (Paraphania) diabolus,

Wied., 526, 527

Philodicus gracilis, v. d. W., 333

Philodicus? sp., 332

Proagonistes ? preceps, Walk., 334

Promachus equalis, Lw., 334

t flavibarbis, Macq., 334

3 maculatus, F., 332

sokotre, Ric., MS., 334
» 2vagator, Wied., 334

Sarcophaga, sp., 334

Index.

Scleropogon ambryon, Walk., 332
Silvius pertusus, Lw., 529
Spilomyia fusca, Lw., 411

Tabanus biguttatus, Wied., 526, 527
Volucella bombylans, L., 491
55 56 var. mystacea
L., 491
Niphocerus cruciger, Lw., 517

NEUROPTERA.

Bracvythemis contaminata, ¥., 332
Mantispa brunnea, Say., 536

| » grandis, Erichs., 537

| Rhyothemis phyllis, Sulz., 334

| Trithemis arteriosa, Burm., 333

? dorsalis, Ramb,, 334

9?

(Ete, Glo BG

Trans. Ent. Soc., Lond., rooz. PL. IX.

Marshall. André & Sleigh, Limited.

All the figures ave about 2 of the natural size.

Injuries to Wings of South African Butterflies.

Trans. Fint. Soc., Lond., r902. PL. X.

G. A. K. Marshall. - André & Sleigh, Limited.

All the figures ave about 2 of the natural size.

Injuries to anal angle and hind margin of hind wings of
South African and Holarctic Butterflies.

jojo, JPit,, AM,

Lond.,

Sac sit

Ent.

7) QHNs.

h, Limited.

Sle ig

lvé &

An

Marshall,

K.

All the figures ave about # of the natural size.

Injuries to directive marks and structures on wings of

South African Butterflies.

Trans, Ent. Soc., Lond., roo2. Pv. XII.

Y, Marshall.

André & Sleigh, Limited.

All the figures ave about 32 of the natural size.

10

Seasonal Phases of South African Butterflies of

the Genus Precis.

Parents and Offspring.

Trans. Ent. Soc., Lond., rooz. Pu. XIII.

G. A. K. Marshall. André & Sleigh, Limited.

All the figures are about 5°, of the natural size.

Under sides of Seasonal Phases of South African Butterflies
of the Genus Precis.

CHW.

Soc., Lond., 1902. PL.

Ent.

Trans.

‘seoeioy pue sidgis\iyo “7 BUTYSIUIIUL SaTy19}]Ng pPUeTeEUOYsSeW

‘azas qounjou ay, fo 9 ynogv atv sainsy ay) 1K

‘paywny ‘4.31915 9 24puy

‘]VYSAVIN MF DO

XV.

Fees

Soc., Lond., 1902.

Ent.

Trans.

“para y ‘YsIa1S Y 24pUVy

‘snddisi1yo “7 SULYSTIUIW SaTpssINg UvOLIpY Jseq Ysa

azas qoungou ayy fo %* gnoqv aav saansy ay? 11K

“1171840 JY

M

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79)

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{
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Pca:
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7 “

er
.

Trans. Ent. Soc., Lond., 7902.

ie, SOW

André & Sleigh, Limited.

A, K. Marshall.

G.

Ze.

of the natural si

9

10

All the figures are about

Common Warning Colours of Mashonaland Acraeine Butterflies, &c.

Lond., r9g02. PL. XVII.

nt. SOC.,

E

Trans.

Sleigh, Limited.

André &

A. K. Marshall.

G,

2 ees :
All the figures ave about =o of the natural size.

Warning patterns and Mimicry of Mutillide in Carabide,

and Cicindelide, &c.

LAGS ETL SOG VEOHAs LOO 2h ben XO SII ee

G. A. K. Marshall. André & Sleigh, Limited.

All the figures are of the natural size.

Mashonaland Insects of many Orders with Lycoid pattern
and colouring, &c.

eo aD av = ie nary
: 0 v
ia et a ' an yt] : y 7
y - 7 : } a ' i: 7
|
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Trans. Ent. Soc., Lond., rg0o2. Pt. XIX.

52 x14
G. A. K. Marshall. André & Sleigh, Limited.
Figs. 30-38 ave twice the natural size.

Figs. 53-59 ave 1% times the natural size.

All other figures ave the natural size.

Mullerian Mimicry in South African Beetles, &c.

Ent. Soc., Lond., roo2. Pr, XX.

Tyvans.

G, A. K. Marshall. André & Sleigh, Limited.

All the figures are about # of the natural size.

Group of black, dark-winged, Mashonaland Aculeates and

their Mimics. First part of Group.

on

Trans. Ent. Soc., Lond., rgo2. PL, XXI.

G, A. K. Marshall. André & Sleigh, Limited.
All the figures ave about £ of the natural size.

Group of black, dark-winged Mashonaland Aculeates and their
Mimics. Second part of Group.

”
“85

aoe S
Sor eer

Trans. Fit. Soc., Lond., roo2z. PL. XXII.

G. A. K. Marshall. André & Sleigh, Limited.

All the figures ave about * of the natural size.
LS 5

Group of yellow-tailed, black, South African Aculeates and
their mimics.

Trans. Ent. Soc., Lond., rooz. PL. XNIII.

G. A. K. Marshall. André & Sleigh, Limited.

2 ‘ :
All the figures ave about 75 of the natural size.

South African Aculeates and their Mimics.

March 5th, 1902.

Papers, ete., read.

Prof. E. B, Poutton, F.R.S., gave an account of a paper by Mr.
Guy A.K. Marshall, entitled “Five years’ (1896-1901) Observa-
tions and Experiments on the Bionomics of South African
insects, chiefly directed to the Investigation of Mimicry and
Warning Colours, with an Appendix containing Descriptions of
New Species by Colonel ©. T. Bingham and Mr. W. L. Distant.”
The paper not only contained Mr. Marshall’s account of his
researches, but the unrecorded notes of many other naturalists
bearing on the same subjects, together with numerous quota-
tions from Mr. Marshall’s letters to Prof. Poulton and the
full description of a large number of specimens illustrating
insect bionomics sent by him tothe Hope Collection at Oxford.
These latter will be permanently retained in the bionomic
section of the Collection, where they can be studied by any
naturalist visiting Oxford. Perhaps the most important part
of the memoir consists of a full description of the long series
of observations and experiments, conducted, with great care,
upon South African insect-eating animals. The insect enemies
made use of were spiders, Mantidx, lizards, frogs, kestrels,
the Ground Horn-bill, the Mongoose, and the Baboon. Exact
observations of birds in the wild state capturing butterflies
are also added, together with a statement of the insects, etc.,
found in the digestive canal of birds, and numerous specimens
of butterflies which, from the character of the injuries received,
appear to have been attacked by enemies. The experiments
on the Mantide are of the highest importance, several genera
being made use of, and the researches continued for many
weeks or even months in the case of certain species. These
predaceous insects freely devoured such forms as Mylothris and
often even Limnas chrysippus, while the Acreinx were evi-
dently distasteful to them. Some of the series of experiments
rendered it probable that Acrxinx are poisonous, or at least
unwholesome food to the Mantidx. The experiments on
kestrels and baboons were also very numerous and important.
As regards the Lepidoptera the support afforded to the theories

Ca)

of warning colours and mimicry was very striking. Another
feature was the great use made of Coleoptera and the strong
witness to the distastefulness of the conspicuous groups, such
as the Lycidx, Melyridex, Cantharidex, etc. As a result of
these experiments and the work of other naturalists, the
attempt is made to draw up a list of the Coleopterous groups
which correspond to the Rhopalocerous Ithomunx, Danaine,
Heliconine and Acrxine. The chief of these are*the con-
spicuous, convergent, frequently mimicked and presumably
distasteful Krotylidx, Endomychidx, Malacodermidx, Melyride,
Coccinellidx, Cantharidx, and Chrysomelide.

The memoir also contains the evidence produced by Mr.
Marshall in refutation of Haase’s conclusion that conspicuous
specially defended butterflies are not attacked by insect
parasites. The conclusion of Profs. Plateau and Wheeler
that human experience of the taste of insects affords trust-
worthy evidence of their effect upon the senses of insectivorous
animals is examined in the light of Mr. Marshall’s experiments
and found to be unsound.

The remarkable mimicry of Curculios with a_ cryptic
colouring by Longicorns is discussed, and the conclusion reached
that the defence provided by sucha quality as hardness does not
appeal like distastefulness to many classes of enemies irrespec-
tive of size, and that the concealment afforded by protective
resemblance is necessary against the considerable number of
foes which are strong enough or large enough to attack
successfully.

The mimetic resemblance of Mantispide to Hymenoptera,
already recognized in North America (Wheeler) and Borneo
(Shelford), is further illustrated by a fine example from South
Africa. The use of insect stridulation as a means of warning
or intimidation is discussed. The experimental evidence of the
value of the terrifying markings and attitudes of Cherocampa
larvee is strengthened by the remarkable impression produced
by an African species upon a pair of baboons. As bearing on
this section it is shown that Cherocampa elpenor is still an
object of superstitious dread in Ireland. The shorter notes
deal with such subjects as ‘“‘The courtship of Limnas chrysip-
pus,’ ‘‘The meaning of the sac of female Acrexine,” “ A

(ta 5)
Rhodesian Muscid fly parasitic on man,” “ Evidence of terror
caused by the squeak of A. atropos,’’ ete.

The material described in the paper illustrates many varied
aspects of mimicry, warning colours, and common warning
colours from the points of view provided by several very
diverse orders of insects. The most striking contribution to
the subject is probably the remarkable group of synaposematic
insects with a Lycoid colouring (fulvous and black), and
the varied assemblages which are made up by Hymenoptera
Aculeata with well-marked types of colouring, each being
attended by insects of other orders, many of which are un-
doubtedly Miillerian components, while others may be Batesian,
although the latter interpretation can only be accepted as
probable in a very small proportion of the examples. Among
these groups perhaps the most important consists of insects
which are entirely black with iridescent blue-black wings.
No less than twenty-eight convergent species of Aculeates form
the centre of an assemblage, round the periphery of which are
scattered Hemiptera, Diptera, Coleoptera (Cantharidx), with
a single Zygenid moth. In another group of almost equal
importance the colouring is similar to the last, except that
the posterior end of the abdomen is bright yellow or orange.
In another the whole abdomen is yellow, and the wings trans-
parent: another is similar to the first described Aculeate-centred
group, except that the head is red : another differs in having a
yellowish head, thorax, ete. Not only the Aculeates but
other specially protected South African insects sent by Mr.
Marshall form beautiful synaposematic groups, sometimes in-
cluded within the limits of a single order, sometimes attracting
insects of other orders. Thus one group of conspicuous little
beetles consisted of six species of Phytophaga, belonging to
six genera, at least one Melyrid, anda Curculionid. The black-
and-orange banded Cantharid type comprises many species,
and a few different genera of these Coleoptera, together with
two Longicorns, two Phytophagous beetles and a Hemipteron,
make up a strong and characteristically marked combination.
The Hemiptera form well-marked and apparently self-contained
groups, one with a conspicuous pattern of black and red with
a black membrane, or a white membrane through which

Calves)

the black body is more or less clearly seen; another pale
yellow with black transverse bars. The first of these groups
is composed of Lygid and Reduviid species, the second of
Pyrrhocorid and Reduviid. To return to the Aculeate-centred
groups, the Mutillide are resembled by Carabidx and
Cicindelide, and yet there is also a secondary resemblance
between these two latter, which becomes primary in the case
of species which do not resemble the Mutilide. In other
cases small slender Carabide of the genus Atractonota primarily
resemble ants in movement and appearance, and yet secondarily
resemble other species of Carabidx in the markings by which
these latter resemble the Mutillide. These complex inter-
relationships suggest proto-, deutero-, and perhaps trito-
synaposematic resemblances for the Miillerian associations,
proto-, deutero-, and perhaps, tritopseudaposematic resem-
blances for the Batesian.

Another important group has for its centre three species
of ants, resembled by a Pyrrhocorid bug of a‘new genus,
Megapetus, described by Mr. Distant in the Appendix, and a
little Locustid of the genus Myrmecophana, with the parts of
the body which would interfere with the likeness to an ant
obliterated, upon the plant on which the insect occurs, by
their green colour. Examples of all these were taken on one
plant in a single day.

Nearly all groups here shortly described were illustrated by
photographs projected on the screen. A brief account of
some of the chief results of Mr. Marshall’s work was read
before Section D of the British Association at Bradford (1900),
and published in abstract in The Report (p. 793).

The number of new facts is so large, the experiments so
numerous and complete, and the range of observation ex-
tended over so many orders in addition to the usually-studied
Lepidoptera, that this memoir places South Africa in the
first rank as the country from which the chief evidence in
support of existing theories of Mimicry, Warning Colours, etc.,
has been supplied.

A discussion ensued in which Mr. F. Merrifield, Dr. F. A.
Dixey, Prof. Hudson Beare, Colonel Yerbury, Mr. J. W. Tutt,
and Prof. Poulton took part.

( 695 )

XX. The Protective Resemblance to flowers borne by an
African Homopterous Insect, Flata nigrocincta,
Walker. By Srpney Lancrorp HINDE. Com-
municated by Prof. EDwARD B. Pou.ton, M.A.,
D.Sc., F.R.S.

[Read June 4th, 1902.]
PLATES XXVI anp XXVII.

THE “cluster of insects grouped to resemble a flower
spike” which forms the frontispiece of Professor J. W.
Gregory's “ Great Rift Valley” (London, 1896) has attracted
much attention and interest, as well as a certain amount
of criticism. As I have had many opportunities of seeing
the insect, and still oftener its larvee, in the wild state, in
British East Africa, and have drawings of both i sitw
made upon the spot by my wife, it seems desirable to
publish the evidence.

Professor Gregory’s plate was apparently drawn in
England from his description and the dried specimens.
In the insects grouped on the vertical stem the green
individuals occupying the uppermost position (Fig. 1) are
represented as considerably smaller than the red ones
below, like the unopened green buds towards the top of a
flowering spike as compared with the expanded blossoms
below. On the other hand, the separate representations
of the green (Fig. 3) and red forms (Fig. 2) of the insect,
as well as the description on pages 273-275 of the work,
indicate that there is no difference in size between the two.
My own experience entirely confirms this latter conclusion,
and there is no doubt that the impression conveyed by
Fig. 1 is in this respect erroneous. Furthermore, the
uniform deep pink colour of the exposed parts of the
insects represented in Figs. 1 and 2 of the frontispiece is
incorrect. The colours of the red forms of the living insect
are as shown on the accompanying Plate X XVI, being of a
bright orange-red anteriorly passing into a reddish-orange
over the remainder of the surface exposed in the attitude
of rest.

Furthermore, I have never seen the insects grouped
according to their colours, but invariably mixed; I have

TRANS. ENT. SOC. LOND. 1902.—PART IV. (DEC.) 46

696 Mr.S. L. Hinde on the Protective Resemblance to

never seen the larvee and imagines on the same stem or
even together on the same tree or bush. I have never
seen the imagines on vertical stems, but always on those
which are actually or approximately horizontal.

It does not by any means follow that Professor Gregory
was mistaken in his impressions, but it is certain that
conditions other than those which he records are common.
The discrepancy is not, however, to be explained by the
hypothesis that I have been observing one species and
Professor Gregory another. My material has been com-
pared with that of Professor Gregory in the British Museum
of Natural History by Professor Poulton, and he states
both sets of specimens certainly belong to the same
species, viz. a form slightly different from Mata nigro-
cincta (Walker), but evidently closely allied and perhaps
specifically identical with it.

One criticism of Professor Gregory’s plate and description
we cannot sustain. I understand that the experienced
African naturalist, Mr. W. L. Distant, holds that the
position shown in Professor Gregory’s Fig. 1 was merely
due to the heavy rain which is stated to have occurred at
the time (loc. cit., p. 273), the insects having crept up the
vertical stem to as great a height as possible in order to
escape the wet. Mr. Distant accordingly believes that
their grouping is unconnected with any protective re-
semblance to an inflorescence. My wife and I, on the
other hand, recognize a strong superficial likeness between
the mixed groups of imsects and the flowers and buds of a
leguminous plant with which we are perfectly familiar.
We have mistaken the groups of insects for the flowers,
and conversely the flowers for the insects. We unfortun-
ately omitted to bring a piece of the plant to England in
order that it may be identified, but this omission can easily
be rectified on our return to Hast Africa.

Although, as I have said, we have never seen the
imagines on vertical stems, the groups of larvae were
generally, although not always, in this position, as may be
seen on Plate XX VII, reproduced about two-thirds of the
natural size from a drawing made and finished upon the
spot by Mrs. Hinde (Jan. 20, 1901). The locality was
an island in the Athi River near the “ Falls,” about twenty-
three miles from Kitui Station. There were dozens of
groups on the shrubs and small trees under the shade of
large trees on the island, and the group painted—a small

Flowers borne by an African Homopterous Insect. 697

one—was that which was most convenient in position,
about four feet from the ground. The long wax filaments
so easily break that it was impossible to obtain satisfactory
results by painting the captured larvee.

The drawing of the imagines was made Jan. 23, 1901,
at Kitui Station, from a branch of a bush which was
covered with groups and single insects, although in both
larvee and imagines these latter are rare as compared with
groups. The bush, which was small, was about fifteen feet
high and ten feet in diameter.

When disturbed the imagines fly and the larve hop a
short distance in any direction, but they soon begin to
collect in groups again: the larva will have reformed into
small groups in half-an-hour. The larve are often seen
on rotten wood and dead leaves, but this is probably after
they have been disturbed. Frequently too, [ have seen
the waxy secretion left adhering to branches where they
have been. The larvae seem to prefer a moist atmosphere
and shade, although I have seen them in the broadest
sunshine at Kibwezi, the locality where the insects were
seen by Professor Gregory. The imagines I have observed
in numbers on three or four occasions and in single groups
several times. The groups of larve are usually about three
or four inches in length, but I have seen a group as much
as two feet long.

The larve towards the growing end of a branch are the
smallest of the group (see Plate XXVIJI), and Professor
Poulton suggests that this observation may perhaps
reconcile Professor Gregory’s account with ours. Professor
Gregory, indeed, considers that the eggs of the Mata are
laid from below upwards so that the insects towards the
top of the stem would be the younger, and he thinks
possibly immature (Joe. cit., p. 275). But the difference in
colour cannot be due to immaturity, for we have found old,
worn specimens of the green form. The first to emerge of
any group may, however, be green, and those that emerge
later red; and Professor Gregory may have come across
undisturbed groups which therefore were green above and
red below. Our groups, on the other hand, may have
reassembled, and thus have lost the arrangement which it
is possible they may have possessed on emergence from
the pupal state. Specimens of larve and imagines
captured at the time when the sketches-were made were
sent by us to the Hope Collection at Oxford.

EXPLANATION OF PLATE XXVI.

PROTECTIVE RESEMBLANCE TO FLOWERS OF British East AFRICAN
FLATA NIGROCINCTA (WALK.).

About # of the natural size.

The sketch was made by Mrs. 8. L. Hinde at Kitui on Jan. 23,
1901, and represents an actual group painted in situ. A red and a
green form of imago are shown separately with their wings expanded.
The Plate is a three-colour reproduction of the original painting.

EXPLANATION OF PLATE XXVII.

Larv# oF British East AFRICAN FLATA NIGROCINCTA (WALEK.).

About 3 of the natural size.

The plate is a half-tone reproduction of Mrs, 8. L. Hinde’s original
sketch made from the larvee in the natural position, on an island in
the Athi River near Kitui, on January 20, 1901. The two larve
which are figured separately from the group were sketched in order
to show the curiously different curves of the waxy filaments in two
individuals.

its ASAOWI

Soc., Lond. 1902.

Ent.

Tvans.

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payay “ys1ag yp aapup

VIViY UPIMYY ISA YSU JO SIIMO]-y 0} IduL[quIVsay 3A19}99}0.1g

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Livans. Ent. \Soc., Lond. roo2. Pi. XXVill.

H. Hinde André & Sleigh, Limited.

About % of the natural size.

Larve of British East African Flata. Athi River,
Jan. 20, 190K.

Report of the Hope Professor of Zoology, 1900.

The additions in 1900 were numerous and valuable, but not
equal in number to those of 1899. The most important is
the extensive collection of N. Bornean butterflies made by the
late W. B. Pryer, Esq., presented by Mrs. Pryer. Important
gifts have also been made by Herbert Druce, Esq., F.L.S.
(many localities), R. Shelford, Esq., M.A. (N. Borneo), Guy
A. K. Marshall, Esq. (Mashonaland), S. L. Hinde, Esq., and
Mrs. Hinde (British East Africa), Professor Poulton (Majorca
and Minorca), Mrs. E. C. Bazett (Uganda), Dr. Henry
Strachan (Lagos), and R. C. L. Perkins, Esq. (Arizona).

The British Collections have also received numerous im-
portant accessions, among which I must specially mention
a splendid partial albino female of the High-brown Fritillary
(Argynnis adippe), captured at Monks Risborough in 1896,
and presented by Miss L. B. Evetts.

The final arrangement of the General Collection of butter-
flies was continued by Mr. Holland, the Morphinae (partially
finished in 1899) and Srassolinae being completed, and a con-
siderable part of the largest family, the WVymphalinae arranged
in about 150 drawers. Allowing a margin of cabinet room for
Dr. Dixey’s arrangement of the Pzerinae, the part of the
collection mentioned above occupied the whole of the con-
signment of 200 drawers delivered in March 1900. Another
consignment of equal size was presented to the Department
by Professor Poulton, and delivered in November. Such
consignments require six months or more for completion,
and therefore the Common University Fund was asked if it
could provide the £200 required for a further consignment of
200 drawers, to be delivered about the middle of the present
year. The sum was granted, and it is expected that the
cabinets will reach Oxford in about a month. A second-hand
cabinet was also purchased for the Department at Stevens’
sale in July, 1900.

B

2

The necessary arrival of fresh cabinets, and the inevitable
spreading out of our crowded collection, as the arrangement
proceeds, and as gaps are filled by the gifts which are con-
stantly reaching us from all parts of the world, makes it
absolutely necessary that more space should be provided.
Immediate needs can be met for some years by the use of
a small part of the south end of the space now occupied by
the Radcliffe Library, and when a new Laboratory is provided -
for the Wykeham Professor of Physics ample room will be

available.

In addition to the above-named work, Mr. Holland ar-
ranged the fine collection of Cicadidae and Fulgoridae, and
the consignments of Phytophaga (Coleoptera) which had been
submitted to Mr. Jacoby. He also worked through the large
collection of butterflies brought by Mr. Richard Evans from
Siam, and the duplicates from the British Museum of Natural
History.

Mr. Hamilton H. C. J. Druce having very kindly consented
to name the Hope Collection of Lycaenid butterflies (“Blues ”
and their allies), a group upon which he is so distinguished an
authority, Mr. Holland arranged examples of all the species
ready for conveyance to London. The working out of this
dificult group will be of inestimable value to the Hope
Museum.

A large amount of Mr. Holland’s time was also occupied in
an examination of the whole of the cabinet drawers and boxes,
and placing fresh naphthaline wherever necessary.

Mr. A. H. Hamm has re-set the butterflies of the General
Collection as far as the point reached at the close of the year
1900, as well as immense numbers of accessions of all kinds.
A large amount of printing, labelling, and cataloguing has
also been done, as the concluding pages of this Report will
indicate. A large number of specimens illustrating biological
problems, such as Mimicry, from Mr. Guy A. K. Marshall
(Mashonaland), and Mr. R.Shelford (Sarawak, N. Borneo), have
been specially set and labelled by Mr. Hamm for the Bionomic
Series. In this condition they have been most successfully
photographed as lantern illustrations by Mr. A. Robinson.

3

The visit of the Council of the Entomological Society took
place on June 30 to July 2. Mr. C. O. Waterhouse, Vice-
President, Mr. Edward Saunders, F.L.S., Mr. A. Hugh Jones,
Mr. H. Donisthorpe, and Mr. C. G. Barrett were present,
together with other students of the Insecta not on the Council,
viz. Mr. Roland Trimen, F.R.S., Hon. M.A., Mr. M. Jacoby,
Mr. Herbert Druce, F.L.S., and many Oxford naturalists.
The visitors were entertained by the Hope Curators and their
friends. The work of the Hope Museum was much assisted
by many kind suggestions. :

Mr. Sidgwick and Mr. Pogson Smith have continued the
arrangement of the British 7zzezva. Mr. C. G. Barrett kindly
assisted in this work on the occasion of his visit. Mr. Gilbert J.
Arrow and Mr. C. J. Gahan have worked through groups
of our Coleoptera, which they have been studying in the
Natural History Museum, and Mr. W. L. Distant has similarly
identified the Westwood types in our General Collection of
certain sections of the Hemiptera. He has published the
results in two papers in the “ Proceedings of the Zoological
Society,’ copies of which will appear in the Hope Reports.

Mr. Jacoby has continued his kind assistance with the
Phytophaga, a labour which will now soon be completed.

Dr. Dixey has continued the arrangement of the Pzerznae
in the new cabinets, and has carried the preliminary work
a long way ahead of the space at his disposal. On Feb. 28,
1go0o, he was elected a Hope Curator, in place of Mr. Edward
Chapman who had left Oxford. It is very pleasant to welcome
Dr. Dixey upon the governing body of the Hope Collections,
which he has done so much to render efficient. He has,
indeed, made his own special group, the Pzerzzaec, a model to
all Museums.

During the year Mr. W. J. Lucas has visited the Depart-
ment, and has helped us greatly in his own group, the
British Odonata. Mr. Percy H. Grimshaw has come to
inspect the General Collection of Diptera. Dr. P. L. Sclater,
F.R.S., Professor A. S. Packard, Professor W. T. Sedgwick,
and Professor J. W. Gregory have also visited the Hope
Museum.

B 2

4

Before concluding I desire as in previous years to express
my indebtedness to the members of the staff of the Insect
Department, of the British Museum of Natural History, as well
as my Satisfaction that the Hope Collection should have
afforded them assistance in their work.

As regards research during the year 1900, I have already
referred to the work upon bionomic questions carried on upon
the collections of Mr. Guy A. K. Marshall, and Mr. R.
Shelford. A condensed account of the chief results was pre-
sented to section D of the British Association at Bradford by
Professor Poulton.

Mr. N. Annandale continued and completed his paper on
his observations in Siam.

Mr. Malcolm Burr published a paper on the British Ortho-
ptera in the Hope Museum in the “ Entomologists’ Record”
for April, 1goo.

The important paper on Mr. C. V. A. Peel’s Somaliland
captures appeared in the “Proc. Zool. Soc.” for the year
1g00 (p. 2).

Volume II of the Hope Reports was not issued until the
beginning of the present year (1901).

ADDITIONS TO THE COLLECTION IN 1896.

Five butterflies from Cannanore on the Malabar Coast
(1896), part of a large consignment presented by A. G.
Cardew, Esq., M.A., Queen’s College, which had been put
aside as duplicates, have now been catalogued for the
General Collection.

ADDITIONS TO THE COLLECTION IN 1897.

Since the last Report the valuable collections made in
Somaliland in 1895 and 1897, by C. V. A. Peel, Esq., and
presented in 1897, have been catalogued and incorporated.
The delay in publishing the paper describing the collection
was the cause of the work being postponed to so late a date,

5

inasmuch as it is desirable to refer to the publication on the
printed labels. This valuable and very varied collection of
523 Insecta, Myriapoda, and Arachnida contains the types
or co-types of several new species (Rhopalocera, Heterocera,
Coleoptera, Diptera, Orthoptera and Arachnida) described by
distinguished specialists in the various groups in the “ Proc.
Zool. Soc.” for 1goo. Mr. Peel is greatly to be congratulated
on the numerous additions to knowledge which have resulted
from his two journeys to Somaliland.

ADDITIONS TO THE COLLECTION IN 1898,

A useful set of insects of many Orders, captured in the
“winter” season at Sao Paulo, S. Brazil, presented in 1898,
by Malcolm Burr, Esq., New College, has been catalogued
and incorporated since the appearance of the last Report.

The gifts of Herbert Druce, Esq., F.L.S., and of F. A. Dixey,
Esq., D.M., Wadham College, still remain uncatalogued, the
former in order that the work may be done in association
with the generous later gifts, the later in the hope that further
information may be forthcoming.

ADDITIONS TO THE COLLECTION IN 1899.

In the Report of last year nearly 7000 specimens, received
in 1899, were recorded as catalogued and incorporated. Great
progress has been made since then, and the following acces-
sions are now gratefully acknowledged.

Eighty-four butterflies and one moth from various localities
in the Bernese Oberland (1898) were presented by Hugh
Richardson, Esq. The data accompanying the specimens
are excellent.

A set of 120 butterflies and one moth from Banff, Alberta,
Canada, and one butterfly from Rush Lake, Ass., Canada
(1898), were presented by H. J. Elwes, Esq., F.R.S. The
data accompanying the specimens are admirable.

A set of 33g insects of various Orders, chiefly butterflies,

6

from Ceylon (1899), was presented by E. S. Goodrich, Esq.,
M.A., Merton College. The data are excellent and render
the collection of great value to the Hope Museum. Many
specimens showing injuries probably caused by enemies will
be added to the Bionomic Series.

Eighty-two insects of various Orders, chiefly butterflies,
from Fiji, and 4 from Vancouver City, B.C., Canada, were
presented by Professor Gustave Gilson of Louvain University.
The specimens are of great value to the Hope Collection on
account of the locality. A pair of convergent Zwploeina will
be a welcome addition to the Bionomic Series.

Collections of 125 Lepidoptera, chiefly butterflies from
various localities in Norway (1898), 20 from Eastern Carinthia
(1897), and 56 from various localities in Switzerland (1899),
were presented by Dr. T. A. Chapman. Many of the specimens
are very welcome additions, especially a set of Ercbia
flavofasciata from Campolungo, Fusio; but all are useful on
account of the data.

The following numerous and valuable accessions are due to
the generosity of G. C. Griffiths, Esq. :

Fifty butterflies and one moth from Chandolin, Switzerland
(z899). These specimens, collected by Professor C. Blachier,
are in beautiful condition.

Eighty-six insects, chiefly butterflies, from Zomba; 15
Lepidoptera from Sumatra; g fine Papilios from the Mo-
luccas; 2 fine Lycaenidae, new to the Hope Collection,
from Kapaur, New Guinea; 13 Lepidoptera from various
localities in Queensland (1897-99); 18 butterflies from the
Bombay Presidency (1898); a moth from Perak; a Pierid
butterfly from Tanganyika; 2 Syntomid Moths from the
Tugela River; 29 Hymenoptera, Neuroptera and Orthoptera
from Japan.

Probably the most important addition to the Hope
Collection of moths, since Mrs. F. W. Hope’s gift of the W. W.
Saunders’ Heterocera, is due to the generosity of Roland
Trimen, Esq., Hon. M.A., F.R.S., who presented the collection
made by Cecil N. Barker, Esq., in Natal, chiefly in the
neighbourhood of Durban (1887-96). The specimens, of

7

which there are 884, belonging to between four and five
hundred species, are in beautiful condition and have stood
the ordeal of re-setting remarkably well.

A Neuropteran (Ascalaphus) from Natal, and 4 specimens
of Melanargia occitanica, from Hyéres (1899), in fine condition,
were also presented by Roland Trimen, Esq.

Colonel J. W. Yerbury presented a valuable collection of
Diptera captured by him in the neighbourhood of Aden
(1895), including 84 Syrphidae described by G. H. Verrall,
Esq., F.E.S., with the type of elophilus africanus, and 93
Asilidae described by F. M. van der Wulp, with the types of
I5 new species. Accompanying the collection are Colonel
Yerbury’s valuable MS. notes on the habits, modes of occur-
rence, &c., of the species. The data on the specimens are
remarkably precise. A few of the specimens are placed in
the Bionomic Series, viz., Hristalts crassipes and a bee which
it resembles found on the same flowers, and two asilid flies
captured in the act of devouring butterflies.

Colonel Yerbury also presented Oestrus ovis from Simla
(1898), and 3 imagos and 4 puparia of Cephalomyia maculata,
the gad-fly of the camel, an insect of great interest and rarity,
bred by him from larvae obtained in the neighbourhood of
Aden in 1895.

A specimen of Argynnis lathonia from the neighbourhood —
of Carnac, Brittany, was presented by Professor Poulton.

Ten specimens of a local Pierid butterfly (Veophasia menapia)
from Vancouver Island (1896), were presented by Mr. A. H.
Hamm. .

In addition to the invaluable material from Mashonaland
presented by Guy A. K. Marshall, Esq., and described in the
Report of last year, a later generous consignment, captured or
bred in 1899, contained the following specimens, many of
which are of the highest interest for the study of Mimicry,
seasonal changes in appearance, &c.

Eighty-two butterflies, captured by Mr. Marshall in the
neighbourhood of Salisbury (1899), containing the very rare
Lycaenid, MWimacraea marshalli, new to the Hope Collection,
and a very fine set of Zeracolz captured at various dates.

8

Two specimens of Mimacraea marshalli, and two specimens
of its model, Lzmnas chrysifpus, taken on the same day
(April 9); for the Mimicry Series.

Twenty-four butterflies, with wings notched, probably as
a result of the attacks of enemies.

A series of 14 bred specimens of Precis sesamus (one being
the ectavia form), subjected to various conditions of moisture
and heat during the pupal period. Precise and accurate data
accompany the specimens, which constitute a solid mass of
evidence on which to build lasting conclusions as to the
manner in which this, the most wonderful seasonal change
hitherto recorded, is brought about.

Five bred specimens of Precis archesia (3) and P. pelasgis
(2), two other deeply interesting seasonal forms allied to the
above. One of the specimens of P. archesia had been sub-
jected to moist heat in the pupal stage. The dates of all the
stages are recorded in the case of the two specimens of
P. pelasgts.

Three female specimens of Zeracolus achine, and the 7
offspring reared from their eggs, some under normal con-
ditions, others in moist heat; four females of TZeracolus
omphale and their 14 offspring similarly treated. Most accu-
rate data are given, and in this case also enable firm conclusions
to be reached upon the much discussed seasonal changes of the
Teracolt.

Professor R. Meldola, F.R.S., presented 225 insects. of
various Orders, chiefly Lepidoptera, and Mrs. Meldola 3
moths, from Switzerland (1899). Professor Meldola also
presented 19 Lepidoptera from Cape Gris Nez. The data
accompanying all these specimens are extremely full and
precise.

Twelve fine Coleoptera (Dynxastidae and Lucanidae) from
Darjiling, were presented by Henry Ward, Esq.

Fifty-one butterflies and one moth were presented by Mark
L. Sykes, Esq., including a very useful set of specimens from
the Gabun River, and valuable specimens from a variety of
localities, many of which are specially wanted for the Mimicry
Series, others for the General Collection.

9

An Argynnis from Tonset, Norway (1899), was presented
by E. N. Bennett, Esq., M.A., Hertford College.

A flower-like spider from Sylhet (1899) was presented by
W. H. Jackson, Esq., M.A., Keble College.

Acraea serena, female, from Zanzibar (1899), was presented
by Mis. E..C.. Bazett.

A large number of specimens are still uncatalogued,
including those to be selected from the Collection of the
East India Company, and the great majority of the duplicates
from the British Museum of Natural History ; the majority
of the generous gifts of Herbert Druce, Esq., F.L.S.; the
Siamese captures of Richard Evans, Esq., M.A., B.Sc., Jesus
College; the great majority of the consignments ens by
R. Shelford, Esq., M.A., from Sarawak ; the butterflies pre-
sented by Abbot H. Thayer, Esq.,and by D. Chaplin, Esq. ; the
Lepidoptera by Osbert H. Howarth, Esq., and by Ronald W.
Poulton.

BRITISH COLLECTIONS, 1899.

Fifty-six Coleoptera from various British localities were
presented by H. Donisthorpe, Esq. The data accompanying
the specimens are very full and precise, rendering them a most
valuable accession.

Fifty-three Hymenoptera Aculeata from various British
localities were presented by Edward Saunders, Esq., F.L.S.
The species were specially selected to fill gaps in our very
complete collection of British Aculeates, and are corre-
spondingly valuable.

A fine series of 12 Caradrina ambigua from Boscombe
(1899) was presented by Major R. B. Robertson.

Three cocoons of the rare “ Hook-tip” (Drepana Sicula),
spun by larvae found in the Leigh Woods, Clifton, Bristol
(1898), were presented by G. C. Griffiths, Esq.

Eleven Hymenoptera Aculeata and mimetic Diptera from
various localities in Scotland (1898) were presented by
Colonel J. W. Yerbury.

Twenty-three insects of various Orders, beset Hymeno-

B3

10

ptera, from Charney, near Wantage, and from Oxford (1899),
were presented by Mr. H. Trim.

Eighteen insects of various Orders from the neighbour-
hood of Oxford (1899) were presented by Mr. W. Holland.
Mr. Holland also presented a most interesting series of 12
specimens of the beetle Cleonus sulcirostris, 4 from the reddish
sand of Boar’s Hill (1899), 4 from sandy soil on Shotover
Hill (1894), 4 from the pale coast sand of Deal (1896). Each
group shows a marked resemblance to the soil on which it
was found, that from Boar’s Hill being especially distinct
from the other two. The observation, the first of its kind in
the Coleoptera, has been described in “ Transactions of the
Entomological Society,” London, 1899, p. 430. The specimens
are a welcome addition to the Bionomic Series.

A hornet (worker) from Summertown (1899) was presented
by the captor, Mr. G. Webb. Curiously enough, the Hope
Museum did not contain a single specimen labelled as captured
in the Oxford district.

A male hornet from Birdlip (1899) was presented by
Mrs. A. G. Butler.

Three Coleoptera were presented by Miss Ruth Butler, and
an imago and pupa-case of JZacroglossa stellatarum by Miss
C. V. Butler. All were obtained at Birdlip (1899).

A set of 101 insects of various Orders from the neighbour-
hood of Oxford; Lingfield, S.E. Surrey; and St. Helen’s, Isle
of Wight (1899), was presented by Professor Poulton. The
series includes a group of Hymenoptera and mimetic Diptera,
for the Bionomic Series.

A beetle was presented by Janet Poulton, and a moth by
Ronald W. Poulton, both from Oxford (1899).

A local moth (Pyralis costalis) from Caversham (1899) was
presented by Miss Cora B. Sanders.

A hundred and sixteen insects of various Orders from
Oxford ; Fleet, Hampshire; and S. Devon (1899), were pre-
sented by Mr. A.H. Hamm. Six of these British Lepidoptera
will be added to the General Collection. ~

The following insects from Oxford (1899) were presented
by the captors:

ll

A dragon-fly by Miss Wright ; a dragon-fly (Gomphus), and
a specimen of “yristalis floreus and its puparium, by Mr. A.
Drew; 2 moths by Mr. A. Robinson ; a pair of Sphinx ligustri,
found on the same rose-bush, by Mrs. E. H. Hayes ; a specimen
of Smerinthus populi by Mr. H. Higgs; 4 specimens (3 green
and 1 red) of Casstda muraea bred from larvae found on /rala
dysenterica near S. Hinksey, by J. J. Walker, Esq., R.N.;
Sphinx ligustri and its pupa-case, by Mr. J. T. Long; a pair
of Smerinthus ocellatus, and a specimen of Cerura vinula and
its.cocoon, by P. J. Bayzand, Esq.

A specimen of Sphinx ligustri from Wotton, Herts (1899),
was presented by Mr. A. Overington.

The only British specimens, received in 1899, still uncata-
logued are the moths from N. Cornwall, presented by A. G.
Cardew, Esq., M.A., Queen’s College.

ADDITIONS TO THE COLLECTION IN 1900,

Large additions were made during the last year, although
not equal in amount to those of 1899. Fair progress has
been made with the labelling, cataloguing, and incorporating,
but a large majority are still unfinished and can only be
provisionally acknowledged in the present Report.

A valuable series of 310 butterflies and 1 moth collected
in April, 1897, in Southern Arizona, and 5 butterflies from
the neighbourhood of New Orleans, were presented by
R. C. L. Perkins, Esq. All the species were greatly wanted
for the accurate data accompanying them, while many were
new to the Hope Collection.

Very large and valuable accessions are due to the kindness
of R. Shelford, Esq., M.A. (Cantab.), the Curator of the
Sarawak Museum (Brit. N. Borneo). Only a part have been
labelled and catalogued, including 16 butterflies from Sarawak,
2 from the N. Shan States, 4 from Malacca,and 1 from Flores
Straits, and 58 insects of various Orders from Mt. Matang
(Sarawak), including some interesting specimens for the
Bionomic Series. The insects were collected at various dates.

B 4

12

One butterfly and 7 moths from various localities were
presented by the Zoological Museum, Tring, Hertfordshire.

A beautiful group of 4 similar butterflies belonging to
three Sub-Families from New Britain (1899) was presented
by Col. C. Swinhoe, Hon. M.A., constituting a valuable
addition to that part of the Bionomic Series which illustrates
theories of Mimicry. Col. Swinhoe also presented 4 butter-
flies of the genus Acraea from S. and S.E. Africa (1899).

A very interesting pair of insects was presented to the
Bionomic Series by H. Donisthorpe, Esq., viz. the Hemipicrond (aug
Lomechusa strumosa and its host Formica «pe, both capturedS y
in the neighbourhood of Roermond, Holland, by Father
Wasmann in 1897.

A fine series of 93 butterflies from British East Africa
(Mombasa, Machakos, and Machakos Road), 1g00, was pre-
sented by the captors, S. L. Hinde, Esq., and Mrs. Hinde.
The series included several interesting examples of seasonal
forms, including a most beautiful variety intermediate between
Precis octavia (wet season) and P. sesamus (dry), of forms
showing the influence of local conditions, such as height above
the sea and dryness. There were also valuable groups of the
different varieties of the same species captured in less than an
hour on a patch of ground a few yards in extent. The
excellent condition of the specimens and the admirable data
accompanying them render the gift especially valuable to
the Hope Museum.

Later in the year another valuable series of 113 specimens
from Machakos (June 6, 1900) and Kitui (Nov. and Dec.) was
presented by the same generous donors. ‘The series included
a very interesting set of 68 Lepidoptera, almost exclusively
butterflies, caught in two hours at a single spot at Kitui, on
Dec. 11. The group represents in a most interesting manner
a characteristic assemblage of Rhopalocera in this locality.
It contains several seasonal forms (wet) of butterflies of the
senus Precis and some interesting local varieties. There are
also species which are new to our collection, and others which
are almost wanting. Among the former one beautiful
Acraea, also absent from the British Museum, is conspicuous.

18

A comparison between the Machakos and Kitui seasonal
forms of the same species is of great interest.

A fine series of two forms of a Pierine butterfly, Cazopsilia
crocale (8) and C. catilla (6), and a pair of another species
(C. pyranthe), all captured flying together on August 11, 1900,
mn the Kangra Valley, W. Himalayas, were presented by L. de
Nicéville, Esq. A pair of the former varieties, captured 27
copula, in the same locality (August 13), was also presented
by the same donor. The specimens are of the highest interest,
showing that, in this locality, two forms which have been
described as seasonal, and may be seasonal in other parts of
their range, do undoubtedly occur together.

Twenty-four butterflies (1g00) from near Eshowe, Zululand,
and one from Natal, were presented by C. B. Russell, Esq.,
M.A., Balliol College. They include a Lycaenid, showing
injuries to the wings, probably caused by the attacks of an
enemy (for the Bionomic Series).

A Blatta from Maitland Camp, near Cape Town (1900), was
presented by Trooper H. E. Nash.

Twenty-one insects of various Orders, from Topeka, Kansas,
U.S.A. (1900), were presented by C. L. Pribble, Esq. They
include specimens showing the effects of probable attacks of
enemies, and the Danaine butterfly, Avosia plexippus, with its
Nymphaline mimic, captured on the same day (for the Bionomic
Series).

A rare Hesperid butterfly (Baorts netopha) from Salisbury,
Mashonaland (1900), was presented by Guy A.K. Marshall, Esq.

Twenty-four butterflies from German New Guinea (about
1897), including a Lwploea and an Elymniine mimic, for the
Bionomic Series, were presented by Mr. A. H. Hamm.

Hight Cefoniidae (Coleoptera), from various localities, were
received in exchange from O. E. Janson, Esq.

Fifty insects of various Orders, and a spider’s egg-cocoon
from Epe, the Mainland, Lagos, West Africa (1900), were pre-
sented by Dr. A. J. Brodie. Although much injured in the
journey, many of the specimens are very welcome to the
Hope Department.

Five specimens of the wide-spread butterfly, Aypolimnas

14

misippus, collected in the Mid-Atlantic in 1893 by Captain
E. P. Ellis, were presented by J. W. Tutt, Esq.

Thirteen Arthropoda of many kinds, chiefly Arachnida,
found alive (December, 1900) in the Oxford Museum, with
specimens from Yezo, Japan, were presented by H. Balfour,
Esq., M.A., Trinity College.

Thirty-one Lepidoptera, captured in Alberta, Canada (1 see
were presented by H. J. Elwes, Esq., F.R.S.

A valuable set of 184 insects, almost exclusively Lepido-
ptera, was presented by Mrs. E. C. Bazett. They were cap-
tured by Mrs. Leakey in 1900, at or near Mengo (chiefly at
Ndeje), Uganda, and include many valuable accessions,
especially a number of moths in fine condition, and the
very rare Danaine butterfly, Welinda mercedonia, new to our
collection. The locality renders all the specimens most
welcome.

A collection of 94 butterflies from Socotra (1898-9) and
one from the neighbourhood of Aden (1898) were presented
by the captor, W. R. Ogilvie Grant, Esq. The data are very
precise, and the specimens form a valuable accession.

Thirty-one butterflies from the neighbourhood of Freetown,
Sierra Leone (1895-6), were presented by Major W. G.
Clements.

Eleven butterflies from Sierra Leone (1897-8) were pre-
sented by F. W. J. Jackson, Esq.

Twenty-nine butterflies, collected in various localities in
Nyassaland (1899) by E. M. de Jersey, Esq., were presented
by the British Museum of Natural History, together with 33
butterflies collected in various parts of the colony of Lagos
(1898) by Dr. Henry Strachan.

Dr. Henry Strachan presented a collection of 212 insects
of many Orders and 3 Arachnida from Lagos, chiefly the Ogun
River basin (1899-1900). Many specimens are very welcome
on account of their rarity, and all on account of the locality.

The following specimens were purchased of J. C. Stevens:
Sixty-eight butterflies from Tenerife (1890). Fifty-two butter-
flies and 2 moths from the Andaman Islands. Forty-eight
Lepidoptera from Queensland. Seventy Lepidoptera and

15

1 Homopteron from Jamaica (1898). . Seventeen Lepidoptera
and 1 Neuropteron from China (almost exclusively from the
West). Fifteen Lepidoptera from Japan, and 15 from
Columbia. Twelve Lepidoptera from Honduras, 8 from
Venezuela, 2 from Colorado, 1 from Paraguay, 7 from Perak,
Malacca, 3 from the Moluccas, 8 from Borneo, 1 from New
Britain, 4 from Kashmir, 3 without locality.

A set of Lepidoptera, almost exclusively picked specimens
for the Mimicry Series, purchased of Watkins and Doncaster,
are as yet uncatalogued.

A fine collection of Orthoptera, including many types, was
purchased from Signor M. A. de Bormans. Many specimens
were unfortunately much injured in the carriage.

The following gifts are as yet uncatalogued :

One of the most important accessions of recent years is the
fine collection of butterflies made in British North Borneo,
chiefly from the East Coast Residency from 1878 to 18y8, by
the late W. B. Pryer, Esq.; presented by Mrs. W. B. Pryer.
Many moths are also included in the collection. The insects
will require a considerable amount of attention inasmuch as
the tropical heat had warped many of the boxes and let in the
dust and mould. The whole collection also requires re-setting,
a labour which is now being undertaken. The labels are
already printed. When completed and incorporated the
Hope Collections will be enriched by many hundreds of
specimens by this generous gift.

Next in importance is the collection of insects made in
Majorca and Minorca (April, 1g00), and to a small extent in
Barcelona (March, 1900), by Professor Poulton, including
captures by Mrs. Poulton and by E. S. Goodrich, Esq., M.A.,
Merton College. The collection consists of hundreds of
specimens of many Orders. All have been set and labelled,
so that nearly all the necessary mechanical labour has been
expended on them. The Hymenoptera Aculeata have been
worked out by Edward Saunders, Esq., F.L.S., the Diptera
by Colonel Yerbury. The former include some interesting
varieties and one fine new species of Momada, of which 4
specimens were obtained.

16

A valuable series of Coleoptera and other insects from the
neighbourhood of Mahon, Minorca, was Peeeaie by Sei.
Mauricio Hernandez.

A very fine and valuable collection of Orthoptera, chiefly
Acridiidae, from Salisbury, Mashonaland (1900), has been pre-
sented by Guy A. K. Marshall, Esq. The insects are now in
the hands of Mr. Malcolm Burr, New College, who is working
them out. Mr. Marshall also presented some extremely fine
examples of mimetic insects of various Orders from the same
locality, together with large numbers of the Hymenoptera
which form the models most commonly resembled. A very
useful collection of Hymenoptera and their mimics from the
same locality was also presented by R. H. Thomas, Esq. The
postal service was probably much deranged by the War, and
these consignments were more badly treated than any I have
previously received from Salisbury.

Extremely fine consignments of insects of many Orders
from Sarawak, British North Borneo, have been presented
by R. Shelford, Esq., M.A., the Curator of the Sarawak
Museum.

The collection of moths has been greatly enriched, as in
previous years, by the generosity of Herbert Druce, Esq.,F.L.S.,
who has also presented a number of butterflies and insects of
other Orders from many localities.

Collections of Lepidoptera from Queensland, of butterflies
from Sarawak, from the Khasia Hills, and from Tobago, and
of moths from Mexico, were presented by G. C. Griffiths, Esq.,
who has given so many valuable donations to the Hope
Museum.

A valuable collection of Lepidoptera from Trinidad was
presented by Prof. R. Meldola, F.R.S.

_ A small collection of insects from Iceland and the Farée
Islands was presented by N. Annandale, Esq., B.A., Balliol
College.

Two Lycaenid butterflies from Egypt were presented by
Prof. Wyndham R. Dunstan, F.R.S., and a moth from Paris
by Prof. Poulton.

Lepidoptera and Coleoptera from Port Elizabeth, and many

17

fine species of Orthoptera from Montenegro and other loca-
lities, were presented by Malcolm Burr, Esq., New College.

A fine collection of dragon-flies from Java was presented by
R. McLachlan, Esq., F.R.S.

A few Oriental butterflies from Col. Swinhoe’s collection
were presented by Mr. W. Holland.

Moths from Sarawak were presented by Col. Swinhoe,
~ Hon. M.A., Wadham College.

Insects of many Orders from S. Africa were presented by
E. N. Bennett, Esq., M.A.,: Hertford College.

Lepidoptera from the Fionnay Valley, Switzerland, were
presented by H. M. Wallis, Esq., and A. Wallis, Esq.

Butterflies bred from European larvae were presented by
F. Merrifield, Esq.

Coleoptera from the United States were spverented by
Ralph d’A. Morrell, Esq., and from the Oriental Region by
BW: Andrewes,; Esq., D:M., Christ»-Church, and) H.. E.
Andrewes, Esq.

In addition to the above-mentioned gifts, a very large
number of duplicate butterflies were generously presented by
the British Museum of Natural History. These will fill many
gaps in our series and supply innumerable examples from
localities which are unrepresented in the Oxford collection.

The British Collections have also been much enriched
although a large proportion of the accessions are not yet
catalogued and incorporated.

A valuable series of 186 Coleoptera and 36 Rhynchota
(both Hemiptera and Homoptera) from various British
localities, all very carefully recorded and dated, were pre-
sented by H. Donisthorpe, Esq.

A pair of Prosopis palustris, a bee new to science, from
Wicken Fen, Cambridge (1899), was presented by the captor,
ee Cabo Peskings hsq-, BeAx

A co-type of Lriocrania fimbriata, a moth new to science,
from Wellington College (1894), was presented by the captor,
Mr. A. H. Hamm.

Six specimens of the rare moth Lupoecilia degreyana, from
Thetford (1g00), were presented by Lord Walsingham.

18

A magnificent variety of Argynnis adippe (the High-brown
Fritillary) was presented by the captor, Miss L. B. Evetts.
In this specimen, a female, the whole of the amber brown
ground colour is replaced by white, the black markings
remaining unchanged. Partial albinos of this kind are known
to occur, although very rarely, in other species of the genus,
but this is I believe the first recorded example in A. adipfe.
The specimen has the additional interest that it was captured
in the neighbourhood of Oxford, on the eastern slope of White
Leaf Hill, Monks Risborough, in July, 1896.

A pair of Colas hyale, captured in I1g00 at Tackley, near
Bletchingdon, was also presented by Miss Evetts.

The following insects, &c., from Oxford (1900), were pre-
sented by the captors :—Cerura vinula, by Mr. G. Long;
Smerinthus ocellatus, by Mr. C. Bolton; 2 Lepidoptera and
a Vespa, by Mr. H. Trim; an ichneumon, by Mr. A. Robinson ;
Zeuzcra aesculi, by Mr. F.C. Hall; a Coleopteron, by Mr.
T. H. Walker ; a spider, by Miss Acland.

Four dragon flies from Newton Abbot, South Devon (1900),
were presented by A. E. Holdaway, Esq.

A very useful set of 47 insects, of many Orders, from
Surrey (including the rare dragon-fly S. faveolum, the local
S. sanguineum, and A. imperator, which is only captured with
ereat difficulty), Hampshire (New Forest), and the Oxford
district (11 specimens), was presented by W. J. Lucas, Esq.

Four cast nymph skins of Aeschna cyanea from South
Leigh (1900) were presented by the Rev. Arthur East.

The following accessions to the British Collections have not
yet been catalogued :

Diptera from Scotch and English localities,and Hymenoptera
from the former, including many specimens for the Bionomic
Series, illustrating mimicry, presented by Colonel]. W. Yerbury.

The year 1900 was remarkable for the abundance of the
larvae and perfect moths of the “ Death’s Head” (Achcrontia
atropos), and the Hope Department received larvae from
Mr. Austin, Mr. W. H. Greenaway, Mr. F. Lewis, Mr. R.
Jones, and Miss Churchill, two pupae from Mr. N. Brett, and
a perfect insect from Mr. G. Hunt.

19

British insects were also presented by the following, and
will be described in detail in the next Report when they are
catalogued :—Rev. J. W. B. Bell, M.A., W. H. Jackson, Esq.,
D.Sc., Keble College, R. W. Poulton, E. P. Poulton, Eustace
Palmer, Esq., Miss C. B. Sanders, H. E. Butler, Esq., Miss
Brown, G. C. Griffiths, Esq , F. A. Dixey, Esq., D.M., Wadham
College, Giles Dixey, Mrs. Gotch, Edward Saunders, Esq.,
Pes: Eh Ponisthorpe, * Esq.) Professor }’Poulton, "Mra Wi.
Holland and Mr. A. H. Hamm, H.' St. G. Gray,’ Esq,
H. A. Ormerod, Esq., Dr. Stark, W. G. Pogson Smith,
Esq., M.A.

Some of the above-named donors presented large numbers
of insects. When all are catalogued it will be seen that the
additions to the .British Collections in Igcoo were very
extensive.

A large series of Hepialus humult from the Shetland
Islands was purchased at Stevens’s sale.

ADDITIONS tO THE HOPE LIBRARY IN 1900,

The Trustees of the British Museum presented the “ Cata-
logue of the Lepidoptera Phalaenae,” vol. ii, by Sir George F.
Hampson, and also papers sent out to collectors of insects.

The Smithsonian Institution (United States National
Museum, Washington) presented the publications which deal
with the subjects of the Department, including the Annual
Reports for 1898 and 1899, and valuable monographs by
A. Busck, Esq., Dr. John B. Smith, Sc.D., Dr. Harrison G.
Dyar, Ph.D., William H. Ashmead, Esq,., 2 W. Coquillett, Esq.,
and Miss Niaiy J. Rathbun.

The University of the State of New York presented a fine
set of Reports of the State Museum (49. 3, 50.2, 51.1, 51. 2)
and the College Department (2.1 and 2. 2).

The Radcliffe Librarian, Oxford, presented the Catalogue
of Books added during 1899.

The Delegates of the Clarendon Press presented Part II
of the “Catalogue of Eastern and Australian Lepidoptera
Heterocera in the Collection of the Oxford University

20

Museum,” by Col. Swinhoe (Pterophoridae and Tineina by
Lord Walsingham and J. Hartley Durrant), Oxford, 1900.

The Boston Society of Natural History and the Bombay
Natural History Society presented their publications for the
year 1900.

_ The Transactions of the Entomological Society, and the

Transactions and Journal of the Linnean Society for the year
1900, were presented by Professor Poulton; also a list of
Lepidoptera of Folkestone, 1870.

The Superintendent of the Museum of Zoology, Cambridge,
presented the Report of the Syndicate for 1899.

“Fauna Hawaiiensis,” vol. ii, Part II (Coleoptera), and
vol. ii, Part IV (Entozoa), were presented by the respective
authors, R. C. L. Perkins, Esq., B.A., Jesus College, and
A. E. Shipley, Esq., M.A., Christ’s College, Cambridge.

The Liverpool School of Tropical Medicine presented
Memoir I, “Instructions for the prevention of Malarial Fever” ;
and Major Ronald Ross, D.P.H., M.R.C.S., presented “Malaria
and Mosquitoes,” and Memoir II, “Report of the Malaria
Expedition” (1900).

Copies of original papers on the Arthropoda have been
presented by the following authors:—M. Henri de Saussure
(2 memoirs); Dr. A. G. Butler (3 papers); W. L. Distant, Esq.
-(9 papers); W. F. Kirby (5 papers) ; Horace St. J. Donisthorpe,
Esq, (F.E.S. (3 papers) ; G.’ W. Kirkaldy,,Esq, Psa uae
papers); O. A. Sayce, Esq. (2 papers); Samuel H. Scudder,
Esq. (3 papers, including an important ‘‘ Catalogue of the
Described Orthoptera of the United States and Canada,”
1900); Sir G. F.: Hampson; B.A} F.Z:S.,:&ce>(“ Phe Moths
of South Africa,” Part 1); Rev. T. RoR. Stebbing iiss:
(“Crustacea from the Falkland Islands”); F. A. Dixey,
Esq., D:M., Wadham College; G. C. Bignell,. Esq: shan Sa
F. V. Theobald, Esg.,.M-A:. ; Gilbert J; Arrow, Psqy tabise
Malcolm Burr, Esq., F.E.S., New College (2 papers, including
a monograph on the Lumastacides); George H. Carpenter,
Fsq-,. B.Se.; W. P. Hay, Esq. Mis. ; Martin Jacoby, gi2say
F.E.S. (Phytophagous Coleoptera from S. and Central Africa) ;
M. André Sémenow ; Professor Roland Thaxter, Harvard

al

University ; Ernest W. L. Holt, Esq., and W. J. Beaumont,
Esq., B.A., Cantab.; M. W. D. Lepeschkin (a Monograph on
the Copepoda).

‘* Tllustrations of the Zoology of the Royal Indian Marine
Survey Ship ‘Inyestigator’” (Fishes Part VII, Crustacea
Part VIII,and Index Part I, 1892—1g00) was presented by the
Superintendent of the Indian Museum, Calcutta.

A valuable Catalogue of the Coleoptera of Minorca, by
'Dr. D. Francisco Cardona y Orfila Pbro (Mahon, 1872), as
well as a supplementary list of Coleoptera by the same author
(Mahon, 1878), and a supplementary list of the Balearic Fauna
by Dr. Juan J. Rodriguez (1887), were presented by Sen.
Mauricio Hernandez of Mahon, Minorca.

Several books were purchased, including “The Butterfly
Book,’ by W. J. Holland (New York, 1899), and among
second-hand works a fine copy of W. Lewin’s “ Insects of
Great Britain” (London, 1795). A fine series of mono-
graphs on Orthoptera by Fischer, de Saussure, Briinner von
Wattenwyl, and Bolivar was purchased from Sign. M. A. de
Bormans.

The parts of Barrett’s “British Lepidoptera,” the Ray
Society volume, and the parts of Rippon’s “ Icones Ornitho-
pterorum ” for the year were also pyrchased.

““Die Physiologie der facettirten Augen,” by Professor
S. Exner of Vienna, purchased in 1898, should have been
included in the Report of that year.

In addition to the gifts acknowledged above, a few works
presented in 1899 were accidentally omitted from the Report
of last year. I have pleasure in thanking Dr. John B. Smith ;
Vernon LU. Kellogs, Esq., M.S.; W. P. Hay, Esq, M.S.; and
Miss Harriet Richardson for presenting copies of their papers,
published through the Smithsonian Institution, Washington ;
Col. J. W. Yerbury for presenting copies of the papers of
Dr. F. M. van der Wulp, on the Aszlidac, and G. H. Verrall,
Esq., F.E.S., on the Syrphidae, collected near Aden; Miss
Edith M. Pratt, B.Sc., and Professor Sydney J. Hickson,
F.R.S.; and Dr. G. S. Brady, F.R.S., for presenting copies of
their papers.

22

A valuable set of 15 papers, chiefly upon the RAyuchota,
was presented by the author, W. L. Distant, Esq.

The Third Supplementary List of the Natural History of
Hastings and St. Leonards, presented by the author, the
Rev. E. N. Bloomfield, was also unfortunately omitted from
the Report.

Two papers “On the method of Organic Evolution,”
presented by the author, Dr. Alfred R. Wallace, in 1895,
were unfortunately not acknowledged in the Report of that
year.

EDWARD B. POULTON.

Report of the Hope Professor of Zoology, 1901.

The additions to the Collection in 1Ig01 were not so
numerous as in 1900. Nearly 6,400 specimens received in the
latter year have now been catalogued and incorporated, and
large numbers still remain to be included. The most important
of these is the very fine consignment from Sarawak, Borneo,
presented by R. Shelford, Esq., M.A., Curator of the Sarawak
Museum.

About 2,000 specimens presented in 1901 have been in-
corporated, and are acknowledged in detail in the later part
of this Report. When the whole of the Igor accessions are
included, the numbers will probably not exceed 4,000
altogether. The most important and valuable of the gifts
already catalogued are the following.

The Rhodesian insects collected and forming the material
of important experiments by Guy A. K. Marshall, Esq., are
of inestimable value to the Department. The specimens have
in large part been specially collected and specially observed
to afford evidence bearing on various biological problems of
the highest interest and importance. Some of these experi-
ments have been described by Mr. Marshall (Ann. and Mag.
INaimeeEtista LOOL,. Vol. ii. p. 3909), Dr./Bs "A. Dixey thas
discussed nearly the whole of Mr. Marshall’s material bearing
on seasonal dimorphism in butterflies and its causes, and has
compared it with a large amount of other material from other
parts of the world, in the Hope Department (Trans. Ent. Soc.
Lond., 1902, p. 189); while the remainder of Mr, Marshall’s
specimens and an immense mass of observations recorded by
him have been communicated to the Entomological Society
of London, and will appear later in the year. This last paper
describes the whole of the South African material bearing on
mimicry, warning colours, the struggle for existence in
insects, &c., accumulated by a matchless observer during the
five years ending with the close of 1go1. Owing to the
generosity of Mr. Guy Marshall, the whole of this invaluable

B

2

evidence on these much-disputed questions can be studied in
the Hope Department.

Of special value to the University Collections are also the
donations by C. J. M. Gordon, Esq., M.A., Balliol College, from
Southern Nigeria; by Col. J. W. Yerbury, from South
Europe; by F. W. Mark, Esq., H.M. Consul, Santos, Brazil,
from Bogota, Columbia; by Miss Mary G. Holmes, from
Manitoba; by W. L. S. Loat, Esq., from the White Nile; by
S. L. Hinde, Esq., and Mrs. Hinde, from British East Africa ;
by the Trustees of the British Museum of Natural History,
from Burmah.

The British Collections have been specially enriched by
a perfect specimen of Vanessa antiopa (“ The Camberwell
Beauty’’), captured (Aug. 19, 1900) and presented by Rev.
J. W. B. Bell, M.A., at Pyrton, Oxon; by Hymenoptera and
mimetic Diptera from various British and Irish localities, by
Col. J. W. Yerbury; by Lepidoptera from many English
localities, by F. A. Dixey, Esq., D.M., Wadham College; by
Lepidoptera from Perthshire, by G. C. Griffiths, Esq.

The welcome gift of nearly 400 beautifully set British flies
from the neighbourhood of Lyndhurst in the New Forest, by
F. C. Adams, Esq., was brought about through the Reports
of University Institutions which are now sent to members of
Convocation. The Rev. Harold T. Adams, M.A., kindly
directed the attention of the donor to the section dealing with
this Department.

Among the uncatalogued accessions by far the most impor-
tant are the specimens from Majorca, the Eastern Pyrenees,
and Montserrat near Barcelona, collected by Professor
Poulton, Mr. W. Holland and Mr. A. H. Hamm, during four
weeks in June and July, 1901. An immense amount of labour
must be expended upon this collection before it is in a con-
dition to be described and incorporated ; but it is confidently
anticipated that there will be several new species among the
less known orders, and many new and interesting records of
the geographical distribution of European species.

Of very great value and importance is the large series of
named American moths presented by W. Schaus, Esq., F.Z.S.

3

Valuable donations have also been made by Herbert Druce,
Esq., F.L.S., from many localities; by W. M. Geldart, Esq.,
WEA., Trinity ‘College, from! Greece; by) R. Shelford, Esq;,
M.A., from Borneo; and by W. C. Boyd, Esq., from many
British localities.

A very important set of Lepidoptera from China and Thibet,
belonging to the collection made by the late J. H. Leech, Esgq.,
was purchased at Stevens’.

The arrangement of the General Collection of Vymphalinae
has been continued by Mr. Holland, and is now nearly
complete. Allowing cabinet space for the Pzerznae which
were being finally arranged by Dr. Dixey and one 20-drawer
cabinet for a portion of the material bearing on mimicry, &c.,
the whole of the consignment of cabinets purchased by the
Common University Fund has been occupied by the butter-
flies of this immense sub-family, and the later groups are
still arranged in temporary quarters. The re-setting of the
General Collection has made but little progress during 1901,
because of the amount of work which has been expended
upon recent accessions, especially the vast series of insects of
many orders from Borneo, presented by Mr. Shelford.

The Lycaenidae, with very few exceptions, have now been
named and arranged in their order by Hamilton H. C. J.
Druce, Esq. It is a very great comfort to know that the
species of this difficult and comparatively little studied family
of butterflies are now accurately determined, and I desire to
express my grateful thanks to Mr. Druce for his kind help.

The usual visit of the Council of the Entomological Society
did not take place, because the expedition to Spain and the
Balearic Islands could not be postponed until after the
time of meeting, in the beginning of July. The Department
has however been visited by many resident and non-resident
naturalists, the latter including Mr. W. J. Lucas, Mr. R.
Shelford, M.A., of the Sarawak Museum, Mr. C. W. Dale,
Mr. W. C. Boyd, Professor C. W. Woodworth, of Berkeley,
California, U.S.A., Mr. F. Muir, Mr. C. J. M. Gordon, M.A.,
Balliol College, of Old Calabar, Professor Meldola, and the
Rev. Canon Fowler, M.A., D.Sc., Jesus College, President of
the Entomological Society of London.

B 2

4

It is a great pleasure to observe that the Department tends
more and more to become a centre for all resident naturalists
who are specially interested in insects.

Mr. Pogson Smith, M.A., St. John’s College, has continued
the identification and arrangement of the British Z7vezna.
Dr. F. A. Dixey has done a great deal of valuable work upon
the General Collection of Pzerznae. The large accessions of
butterflies of this group, constantly received from many
parts of the world, especially the Ethiopian region, have
involved much extra labour because of the re-arrangement of
the older material which became necessary. As a result of
Dr. Dixey’s patient researches and remarkable power in
using the materials of his investigations as a means for
elucidating the problems of evolution and unravelling the
tangled threads of phylogeny, the Oxford Collection of
Pierinae has become one which no serious student of the
group can afford to neglect.

I again wish to express my thanks to the members of the
staff of the Insect Department of the British Museum of Natural
History, who have helped me in the study of many groups.

The chief researches conducted in the Department during
1g0ot have been concerned with the material presented by
Mr. Guy A. K. Marshall and Mr. R. Shelford, and the
Balearic insects, collected by Professor Poulton. References
to the publications will be found in the parts of the Report
dealing with these donations.

ADDITIONS TO THE COLLECTION IN 1898 AND 1899.

The gifts which were uncatalogued at the publication of
the last Report remain in the same condition. The delay
has been in part due to the hope that further data may be
forthcoming, and in part to the advantage of printing the
donations of two or three years together in those cases in
which the same data are likely to be repeated frequently.
The immense labour required by the generous consignments
of R. Shelford, Esq., M.A., from Sarawak, Borneo, and by
the Siamese butterflies collected by Richard Evans, Esq.,

5

M.A., D.Sc., Jesus College, also severely taxed the Depart-
ment. The pinning and setting of the former is now complete
and the latter far advanced.

Mr. Shelford’s valuable and extensive material bearing on
the study of mimicry in Bornean insects presented in 1899
and the two subsequent years illustrates a paper by him
which was read before the Zoological Society in January,
1902, and will shortly be published in the Proceedings.

ADDITIONS TO THE COLLECTION IN 1goo.

Since the last Report a large number of specimens have
been catalogued and incorporated, and are now gratefully
acknowledged. .

A set of 6 butterflies collected by Lieut.-Col. Manning at
Zomba (1899).

EeESeeT or. 02 butteriies collected by Col. J. Bi, Yulesin
Nyassaland (1896).

A set of 36 Lepidoptera collected (1894) in the Arusa
Galla Country, Somaliland, by F. Gillett, Esq.

A set of 30 butterflies collected (1898) by A. Ross, Esgq., at
Johannesburg. ~

A set of 25 butterflies collected or bred by Rev. W. D.
Cowan, in Betsileo, Madagascar.

A specimen of Limnas chrysippus captured (1891) at Fort
Dauphin, Madagascar, by the Hon. Reg. Walsh.

Five butterflies from the forests near Antananarivo, Mada-
gascar (date uncertain).

A specimen of Hyfolimnas misippus collected (1887) by
D. W. Barker, Esq., R.N.R., at Massowah.

A specimen of Limunas chrysippus collected by G. Gulliver,
Esgq., at Rodriguez (1874).

Seven butterflies collected by C. W. Bewsher, Esq., at
Johanna Island, Comoro Islands.

The whole of the above-named specimens were duplicates
from the collection of the British Museum, kindly presented
by the Trustees.

6

A set of 5 Locustidae from the Bay of Antivari, Monte-
negro (1g0o), were presented by the captor, Malcolm Burr,
Esq., New College.

A male specimen of the moth O. dispar from Paris (1900)
was presented by the captor, Professor Poulton.

Nine butterflies from the Sandwich Islands were presented
by the Trustees of the British Museum, including several speci-
mens of Vanessa tammeana captured by R. C. L. Perkins, Esq.,
in 1895-6. This magnificent ally of our own “ Red Admiral ”
(Vanessa atalanta) is new to the Hope Collection.

The splendid accession to the Oriental species in the Collec-
tion, which we owe to the generosity of Mrs. W. B. Pryer, has
now been almost entirely catalogued and incorporated. Nearly
the whole collection was made by the late Mr. W. B. Pryer
at the East Coast Residency of British North Borneo between
the years 1878 and 1898. 1,043 butterflies from this locality
have been incorporated, viz. the whole of the families with
the exception of the Lycaenidae, which have been very kindly
worked out for us by Mr. Hamilton H.C. J. Druce. 92 moths
and 7 insects of Orders other than the Lepidoptera from the
same locality have also been added to the Collections. There
still remain the numerous Lycaenidae from the East Coast
Residency, and a comparatively few specimens from other
places, or from exact localities within the Residency. This
important gift, together with the fine series of Bornean butter-
flies presented by Mr. Herbert Druce in 1899, renders the
University collection from this island remarkably fine and
complete.

One Asilid (Diptera), and 45 Hymenoptera, comprising
40 Aculeata, 4 Chrysididae, and 1 Ichneumonid, from the
neighbourhood of Salisbury, Mashonaland (1899-1900), were
captured and presented by R. H. Thomas, Esq.

These Hymenoptera, together with the large collection from
the same locality presented by G. A. K. Marshall, Esq., are
being kindly worked out by Col. C. T. Bingham.

The following splendid additions to the University Collec-
tions are due to the kindness and skill of Guy A. K. Marshall,
Esq. The locality, except when otherwise stated, is Salisbury,
Mashonaland, and the captor Mr. Marshall.

7

A fine general collection of Hymenoptera Aculeata, con-
taining 610 specimens. Some of these have been found by
Col. Bingham to be new species. Other Hymenoptera :
5, Chrysididae, 13 Chalcididae, and 3 other Terebrantia.

The following Orthoptera: 2 Forficulidae, 23 Blattidae.
198 Acridtidae, 23 Locustidae, 6 Gryllidae, 47 Mantidae, and
1 Phasmid. Also 1 Locustid and 1 Blatta from the Umfuli
district (1899).

The following Rhynchota: 10 Hemiptera and 3 Homoptera.

Thirteen Diptera: 1 M/antispa (Neuroptera), 10 Coleoptera.

The following fine additions to the bionomic series, espe-
cially the parts illustrating Batesian and Miillerian Mimicry,
Fifteen Hemiptera, forming components of various mimetic
groups: 2 Reduviids (Hemiptera), a Longicorn and a Can-
tharid beetle, all dark iridescent blue-black, and resembling
an important group of Aculeate Hymenoptera. The wonderful
group of insects mimetic of the well-known Ethiopian Lycid
beetles, with a tawny colouring anteriorly and black poste-
riorly, formerly presented by Mr.- Marshall, has now been
greatly strengthened by 8 beetles, 4 moths, 2 Hemiptera,
3 Ichneumonids, and 2 Aculeate Hymenoptera. Thirty-four
specimens are divided between the following groups, the
models of which appear always to belong to the stinging:
Hymenoptera: (1) black insects with yellow or red head and
sometimes thorax, and in some species black and yellow legs—
Aculeates, Ichneumonids, Hemiptera,beetles, flies ; (2) greyish
black anteriorly, red-brown posteriorly—Aculeates and an
Asilid fly; (3) black with red or yellow apex to abdomen
—Aculeates, Bombylid fly; (4) black ant-like group—ants,
Coreid and Reduviid Hemiptera.

A small proportion of Mr. Marshall’s generous gift, which
is here gratefully acknowledged, has been accidentally
transferred to the catalogue for Ig00 instead of IgoI to
which it properly belonged.. Hence his donation for the
latter year appears smaller than it actually was, and that for
the former year larger in the same degree. Inasmuch as the
printed locality labels on the specimens remain strictly
accurate and contain a statement of the correct date of pre-

8

sentation, it was not thought necessary to undertake the con-
siderable labour of altering the catalogue and the small label on
the specimens which record the catalogue number and year.

A fine collection of 419 insects of all Orders from British
Central Africa was presented by C. V. A. Peel, Esq. The
insects were collected in 1899 by C. H. Pemberton, Esq., over
a very extensive tract of interesting country to the W. of
Lake Nyassa, from Kota-Kota on the shore of the Lake to the
Loangwa River Valley, the Mushinga Mountains and Chitala.
The numerous localities and dates are accurately recorded.
The collection is especially valuable, inasmuch as the Hope
Museum possesses very little material from this part of Africa,

A small collection of 36 Lepidoptera made by the same
naturalist at Delagoa Bay on Aug. 10, 1899, and a Mantid
captured by him on the Lower Zambesi in July of the same
year, were also presented by C. V. A. Peel, Esq.

A set of 55 insects of various Orders, and 10 other Arthro-
poda from different localities in Cape Colony, were captured
(1899-1900) and presented by E. N. Bennett, Esq., M.A.,
Hertford College. Although the species are apparently not
rare the localities render them of much interest and value to
* the Hope Collection.

The collection of insects of many Orders from Majorca
and Menorca captured in March and April, 1900, has now
been catalogued, and some of the groups worked out and lists
published. This collection, together with the much larger
one made in Igo0l, constitutes by far the most important
addition to our European collection during recent years.
The following is an account of the 1900 captures.

Five hundred and ninety-eight specimens were captured
and presented by Professor Poulton. The great majority of
these are from Majorca (many localities), a relatively few
from Menorca (Mahon). A list of the Diptera has been
published by Col. J. W. Yerbury (Ent. M. Mag., 1890,
p. 272), the species of principal interest being Phorantha sub-
coleoptrata and Bombylius pictus. The Hymenoptera Aculeata
have been described by Edward Saunders, Esq., F.R.S. (l.c.,
p. 208). . Forty-eight species were obtained, two of which are

9

new to science, viz. a fine bee of the parasitic genus Vomada |
described as NM. poultont (Saunders), and a new F/lalictus,
fT. soror (Saunders). Of the latter 2 females were obtained
(Castle Bellver, Palma, March 26 and 30); of the former
1 male and 3 females (Castle Bellver, 400 ft., on flowers,
1 female March 25, the others March 30). The common
British Humble-bee, Bomdbus terrestris, was very abundant
on flowers; but all the specimens were of the variety ferru-
gineus, from SW. France, Spain, and Portugal, which differs
from the type in having the hairs of the tibiae fulvous instead
of black. Of the Hemiptera 14 species were obtained, and
a list has been published by Mr. Saunders (I. c., pp. 239, 240),
who points out that all the specimens of Lygacus pandurus,
very abundant in Majorca, are of the variety with milky-white
unspotted membrane, like those from Algeria.

Twenty-eight insects and arachnids from the same localities
were captured and presented by E. S. Goodrich, Esq., M.A.,
Merton College, and 17 Coleoptera by Mr. Goodrich and
Professor Poulton. Of the latter, 15 were captured in
Menorca, where beetles were much more abundant than in
the larger island, at the time of our visit.

A valuable set of Coleoptera and a few other insects from
the neighbourhood of Mahon were captured and presented
by Sen. Mauricio Hernandez. They include 42 Coleoptera
captured about 1885, together with 27 insects of various
Orders captured in 1900. The few Hymenoptera Aculeata are
included in the Mr. Saunders’s published list alluded to above.

Forty-one Coleoptera and one Hemipteron were captured
(April 6, 1900) near Mahon, and presented by Mr. Goodrich,
Sen. Hernandez, and Professor Poulton.

Forty-three insects of various Orders from the neighbour-
hood of Barcelona and Montserrat (about 3,000 ft.) were
captured in March, 1900, and presented by Professor Poulton,
and 6 insects from the same localities by E. S. Goodrich, Esq.,
while 6 from Barcelona were captured and presented by these
two naturalists jointly. Specimens from this part of Spain
have hitherto been wanting in the Hope Collection, and are
rare in European collections generally.

By

10

Two specimens of Deudorix antalus from Manashi, near
Cairo, were presented by E. A. Floyer, Esq. The larvae had
been injurious to “ /zga dulcis,’ feeding upon the seed-pods.
The specimens were kindly transmitted by Professor Wyndham
R. Dunstan, F.R.S.

Seven specimens of Aporia cratacgé were bred from pupae
(Germany, 1900) presented by F. Merrifield, Esq.

A pair of the Aculeate, Prosopis variegatus, and of the fly
(Paragus bicolor) which mimic it, were presented by Edward
Saunders, Esq., F.R.S. The specimens were from Algeria
(1896), Italy (1895), and Brittany (1900). The interesting
resemblance of the facial markings of the male fly to those of
the male bee, and the facial markings of the female fly to
those of the female bee has been described by Mr. Saunders
(Ent. Monthly Mag. 1900, p. 83).

The following kind donations, mentioned in the Report of
last year, are still uncatalogued. The Bornean insects of
R. Shelford, Esq., M.A.; the moths from many localities, and
other insects of Herbert Druce, Esq., F.L.S.; the Lepidoptera
from the Khasias, Borneo and Mexico, of G. C. Griffiths,
Esq.; the insects from Iceland and the Farodes of N. Annan-
dale, Esq., B.A., Balliol College; the South African and
Eastern European insects (in addition to those from Monte-
negro, catalogued above) of Malcolm Burr, Esq., New College ;
the Javan dragon-flies of R. McLachlan, Esq., F.R.S.; the
Oriental butterflies of Mr. W. Holland; the Bornean moths of
Col. Swinhoe, Hon. M.A., Wadham College; the Swiss butter-
flies of H. M. Wallis, Esq.and A. Wallis, Esq.; the American
Coleoptera of R. dA. Morrell, Esq.; the Oriental Coleoptera
of F.7W. Andrewes, Esq., D.M.,Christ Churtch,-and) Hig
Andrewes, Esq.; and a large proportion of the duplicate
butterflies presented by the Trustees of the British Museum
of Natural History.

The specimens purchased from Sign. M. A. de Bormans
and Messrs. Watkins and Doncaster are also uncatalogued.

The catalogue of British insects presented in 1g0c has now
been completed, and the specimens incorporated in the
collection. The gifts which were not catalogued at the
appearance of the last annual Report are acknowledged below.

iL!

A specimen of Plusia moneta, captured on honeysuckle at
Pyrton (1900), was presented by Rev. J. W. B. Bell. This
interesting moth, which was unknown in this country not
many years ago, has been hitherto unrepresented in our
British Collection.

A beetle, Balaninus glandium, from near Bladon (1g00) was
presented by Miss C. A. Brown, and Hedobia imperialis from
Oxford (1900) by J. E. Pogson Smith.

A bred specimen of the moth Axarta myrtilli, together
with its cocoon and pupa-case, from Dartmoor (1899), was
presented by H. E. Butler, Esq.

A Dipteron from Oxford (1900) was presented by Dr. W. H.
Jackson, of Keble College.

A specimen of Polyommatus corydon from Boar’s Hill
(Sept. 12, 1900) was presented by H. St. G. Gray, Esq.
together with C. edusa and JN. lucina from Dorset and
Wiltshire. The former capture is of great interest, for the
“ Chalk-hill Blue” has never before, so far as I am aware,
been observed so near to Oxford.

An interesting series of 8 Sierinthus populi were bred in
the Hope Department (June-July, 1900). In the excessive
heat of the latter month in Ig00, the moths appeared in a
week or two after pupation, instead of emerging in June,
1901, after a pupal period of normallength. All the specimens
are of an unusual shade of colour, a probable result of the
high temperature.

Four moths and a dipterous insect were captured in York-
shire (1g00), and presented by Professor Poulton and E. P.
Poulton, Esq.; a remarkably small worker wasp (Vespa
vulgaris), from St. Helens, Isle of Wight (1900), was pre-
sented by Ronald W. Poulton.

Sixty-nine insects of several Orders from the neighbour-
hood of St. Helens, Isle of Wight, and 18 from various
localities in and near Oxford, were captured (1900) and
presented by Professor Poulton.

A pair of Vanessa urticae (“The Small Tortoise-shell
Butterfly”), captured in the meadows by the Cherwell at
Oxford (May, 1goo), was presented by Professor Poulton.

12

The later stages of courtship were observed in these specimens,
for the first time in the case of this common species. The
notes will be recorded.

Twenty-two insects of different Orders from the neighbour-
hood of Oxford (1900) and 11 moths from near Reading
(1891) and Basingstoke (1892) were captured and presented
by Mr. W. Holland.

A very useful set of 115 insects of different Orders from
Mundesley, Norfolk, was captured (1900) and presented by
Mr. W. Holland. The series includes a fine series of the
varieties of the moth Zygaena trifolii Mr. Holland also
captured in the same locality and presented 62 insects belong-
ing to the Hymenoptera, Diptera and Coleoptera. These
insects consist of three groups, each captured in a single day,
and in one place. The species in each group thus found
together exhibit more or less marked similarity in colour and
pattern, and will be kept together in the bionomic series.

A specimen of the Pentatomid bug, Aethus flavicornis (Fab.),
was presented by Mr. W. Holland, who captured it at Fresh-
water, Isle of Wight, in July, 1895. The specimen is of great
interest, having been identified and recorded by Mr. Edward
Saunders as “a genus and species new to the list of British
Hemiptera” (Entomologists’ Monthly Magazine, 1899, p. 155).

One hundred and thirty insects of different Orders from
various localities in and near Oxford were captured (1900)
and presented by Mr. A. H. Hamm. They include Aculeate
Hymenoptera with their dipterous foes, and sets of insects
of very different kinds, with a similar appearance, taken
together at the same place and time. Four specimens of
a butterfly new to the Oxford district, Zhecla W. album
(“The White Letter Hairstreak”), from Tubney, are also in-
cluded, together with 3 Colias hyale (“ Pale Clouded Yellow”)
from Cowley Marsh.

Another set of 54 insects from the neighbourhood of
Oxford and S. Devon was presented by Mr. A. H. Hamm,
having been captured or bred by him in 19co. They in-
clude a fine series of bred Callimorpha hera (“The Jersey
Tiger”) from Dawlish, a moth probably introduced into this

13

country and now steadily extending its range; and 7 speci-
mens of Pyrrhosoma tenellum, from Newton Abbot, a dragon-
fly new to our British Collection. Many of the specimens
illustrate the biological relations between insects and their
enemies, and will be added to the Bionomic Series.

Mr. Hamm also presented an interesting set of 6 cocoons of
the moth C. zeustria (Oxford, 1900), 5 of which had been
opened by birds and the pupae abstracted, while one had
produced a parasitic insect : for the Bionomic Series.

Colonel Yerbury’s donation of Scotch and English Diptera
and Hymenoptera were erroneously catalogued as presented
in 1901 (see p. 20).

The 5 specimens of Acherontia atropos, bred from larvae
from the Oxford district, presented by Mr. Austin, Mr. W. H.
Greenaway, Mr. F. Lewis, Mr. R. Jones, and Miss Churchill,
together with the specimen of the perfect insect presented by
Mr. G. Hunt, are now catalogued and incorporated. The two
pupae presented by Mr. N. Brett failed to produce moths.

Two specimens of Papilio machaon were bred from pupae
from Wicken Fen (1899), presented by F. Merrifield, Esq.

Four Vanessa polychloros were bred from larvae-from West
Sussex (1900), presented by Miss Cora B. Sanders.

A specimen of the rare moth Bombyx trefolii, from
St. Helens, Isle of Wight (1900), was presented by
Mrs. Gotch.

A specimen of the very rare butterfly Pzeris daplidice (the
“Bath White”), with the locality Ascot (1897), was presented
by H. A. Ormerod, Esq.

A specimen of Agrotis praccox, captured at “sugar” at
St. Helens, Isle of Wight (1900), was presented by E. P.
Poulton, Esq.

Interesting examples of a British beetle (Polydesmus un-
datus) protectively resembling parts of the birch (Tilgate
Forest, 1891), and of Creptodera transversa resembling the
seeds swept with them (Oulton Broad, 1900), were presented
by H. Donisthorpe, Esq., together with a specimen of Cassida
equestris (Wicken Fen, 1900) with injuries probably caused

14

by the attacks of enemies. This material is a welcome
addition to the rapidly growing Bionomic Series.

A set of ants, and the immature Hemiptera which mimic
them, were presented by Mr. A. H. Hamm. The insects were
captured together on the bark of an apple-tree at Oxford
(1900).

Six specimens of the beetle Wzptus hololeucus, from Oxford
(1900), were presented by Dr. Stark; and 7 of the same
species, from Shropshire (1900), by W. P. D. Stebbing, Esq.
The latter were found in decayed fot pourri fifty years old.

Twenty-nine insects of various Orders from Oxford and
various other English localities (1900) were captured and
presented by Dr. F. A. Dixey. Twenty insects of various
Orders from the neighbourhood of Totland Bay, Isle of Wight
(1900), were captured and presented by Giles Dixey. Ex-
cellent data accompany both these sets of insects.

Seventy-five specimens of the interesting northern variety,
hethlandica, of the “ Ghost Swift,” Hepialus humuli, from Unst,
Shetland, were purchased (1g00) at Stevens’s sale. Eight
specimens are added to the General Collection.

ADDITIONS TO THE COLLECTION IN 1g0I.

A valuable set of 113 Lepidoptera from Trinidad (about
1874) were presented by Professor R. Meldola, F.R.S.

Three specimens of a rare Satyrine butterfly (Leptoneura
bowkert) from Natal (1901) were presented by Colonel Swinhoe,
Hon. M.A., Wadham College.

A small but valuable set of 75 Lepidoptera, 1 Cicada, and
1 Longicorn beetle, from Southern Nigeria, were captured
(1900-1901) and presented by C. J. M. Gordon, Esq., M.A.,
of Balliol College. The exact locality of capture was the
stretch of thirty miles intervening between the Forcados
mouth of the Niger and Warri, a most interesting district, and
one hitherto unrepresented in the Hope Collection. Among
the butterflies an Erycinid and all the Lycaenzdae were espe-
cially wanted, together with some very interesting mimetic
specimens.

15

A valuable set of 77 butterflies from Southern Spain (Alge-
ciras, Cordova, Granada, Malaga, Gibraltar) were captured
(1901) and presented by Col. J. W. Yerbury. The excellent
condition of the specimens and the full data accompanying
them render the gift most acceptable. The specimens of the
Pierine, Zegris eupheme, with full data, were specially wanted.

Col. Yerbury also presented a pair of the Bombus-like fly,
Mallota fuciformis, captured by him at Hyéres (1898), and
a pair of the wasp-like fly, Cerza eumenoides, captured at
Matheran, Bombay (1879), by Major C. G. Nurse: for the
Mimicry Series.

Material of inestimable value from Rhodesia, illustrating
seasonable changes and their causes, mimicry, &c., was pre-
sented by Guy A. K. Marshall, Esq., together with a fine
series of specimens for the General Collection. Unless other-
wise mentioned, the specimens were captured or bred at
Salisbury, Mashonaland, in 1901.

Twenty-five specimens of the Pierine genus Zerzas include
the results of many experiments in which Mr. Marshall at-
tempted to reproduce the wet season form by placing the
larva and pupa or the pupa alone in a damp atmosphere.
The species experimented on were 7. senegalensis, T. regularts,
and 7. brigitta. The results have been recorded and discussed
by Mr. Marshall (Ann. and Mag. Nat. Hist., 1901, vol. ii,
p- 398), and by Dr. Dixey (Trans. Ent. Soc. Lond., 1902, p. 189).

Fifteen specimens of the two species of the Nymphaline
genus Byblia, B. ilithyia and B. acheloia, included 4 captured
females and offspring reared from eggs laid by each of them ;
thus demonstrating the parallel seasonal alternation in pattern
and colouring of the under side of the wings in both species.
The results of these investigations, which in certain cases
included the use of a damp atmosphere, have been recorded
and discussed in the papers already referred to by Mr. G. A. K.
Marxshall and Dr. F. A. Dixey.

Three specimens of the Nymphaline butterfly Preczs
archesia had been subjected in the pupal state to a moist
atmosphere. The forms were nevertheless those which are
characteristic of the dry season, apparently indicating that

16

the change is not in this species due to the direct influence
of environmental conditions. This result has also been re-
corded by Mr. Marshall (1. c.).

Nineteen butterflies showing injuries to the wings, probably
caused by the attacks of enemies.

Five butterflies captured on the same day, Sept. 28, 1900,
including 3 similar species of Acraea and 1 Lycaenid with
an Acraea-like under side.

The Lycaenid Alacna nyassa and the Hesperid Cyclopides
qwillemt with a somewhat similar under side, captured on the
same day, Feb. 23, 1901; and a similar pair captured
March 3, 1901.

A dark-winged fly and Zygaenid moth (Dec., 1900),
mimetic of the large group of black dark-winged Aculeate
Hymenoptera.

Three Ichneumonids and 2 Longicorns (1900, 1901) form
part of an interesting group of black insects with the anterior
parts of an ochreous or reddish colour. An Aculeate and
a sesiid moth (1900), with the same general type of colouring,
afford beautiful examples of model and mimic. Four Ichneu-
monids (1900) constitute a very characteristically coloured
group of black-barred ochreous insects mimicked in the most
perfect manner by a Reduviid bug of a new species. Two
other Reduviids entered respectively the large group of which
the models are many species of black Aculeates, and the
characteristic group formed round the Lycid beetles.

A beautiful group of 4 black Aculeates with yellow- or
orange-tipped abdomen from Umtali, Mashonaland (1900),
are beautifully mimicked by a sesiid moth from the same
locality: 2 specimens of the latter were presented. Three
specimens of Acraea encedon, 2 of a Lycaenid (Catochrysops
peculiaris), and 2 of a rare Hesperid (Adantis tettenszs), all
from Umtali (1g00). The two latter species show probable
incipient mimicry of the Acraeine type.

A most interesting group of insects was captured together
on a plant at Salisbury on Feb. 17, 1901. Four specimens of
black ants were the models mimicked by a Locustid, A7yrme-

17

cophana sp., and 2 specimens of a Coreid bug of a new genus
and species to be described by Mr. W. L. Distant in Mr.
Marshall’s forthcoming paper describing the whole of his
material which bears on mimicry and allied questions.

Two specimens of a dark Ichneumonid with a transparent
area in the opaque fore-wings. This species may be the
model of a Mashonaland Bombylid fly with a somewhat
similar character.

A specimen of an Asilid fly, with its prey a Lycaenid
butterfly (1900). Another Asilid, caught (1901) in the act
of devouring a dragon-fly far larger than itself.

The General Collection has also been further enriched by
Mr. Marshall’s great generosity.

Fifty-nine butterflies (Salisbury, 1g01) include several
interesting and valuable additions to our series of the genus
Precis, with the exact data which is so especially necessary in
studying their seasonal forms. The rare Hesperid Baoris
netopha is represented by 3 examples. Eleven other insects
include a male specimen of the rare mosquito discovered by
Mr. Marshall, Wegarrhina marshalti.

Thirty-five insects from Umtali, Mashonaland, 3,700 ft.
(1900), include further valuable additions to the genus Precis,
and several rare Hesperidae, &c.

Mr. Marshall also presented the following insects not of his
own capture :—Three moths from Beira (1900), 2 insects from
Umtali (1900), and a pair of the Danaine butterfly Lzmnas
chrysip~pus, var. klugit, from Dar-es-Salaam, German East
Africa (1899).

A fine series of 7 specimens of the beautiful Morphine
butterfly Zeuridia horsfieldit, and 10 specimens of Z. ame-
thistus from Sandakan, Borneo (1895-6), were presented by
Herbert Druce, Esq., F.L.S., together with a set of 7 Limuas
chrysippus and 1 Argynnis lathonia from Tenerife, and a large
number of insects as yet uncatalogued.

Fifty-two insects of many Orders, and 3 spiders from
Madeira, were captured (Dec. 1900—Jan. 1901) and presented
by E. S. Goodrich, Esq., M.A., Merton College.

Two hundred and forty-five Lepidoptera, collected (1848-57)
almost, if not entirely, in the neighbourhood of Bogota,

18

Columbia, by the late Edward W. Mark, Esq., H. M. Vice-
Consul, were presented by F. W. Mark, Esq. Eight of the
specimens are moths, and the remainder butterflies. All are
much wanted in the University Collection, several species
being entirely unrepresented. Considering the date of capture
and the fact that no special technical skill has been employed
in preventing deterioration for so long a period, the condition
of the majority of the specimens is remarkably good. I desire
to: thanle E. J. Trevelyan, Esq., M.A. B.C: All tSenis
College, and Mrs. Trevelyan, for rendering generous assistance
to the University Collections by bringing the matter to the
attention of the kind donor.

A large number of insects other than Lepidoptera, with the
same history as the above, were also presented by F. W.
Mark, Esq. These are as yet uncatalogued.

A valuable set of 80 insects of various Orders, and 1 Arachnid
from near Laurier, Manitoba, Canada, were presented by Miss
Mary G. Holmes. The data are most precise and detailed, and
the specimens a most useful addition to our Canadian series.

Two specimens of the extremely local Erebta arete were
captured (July, 1877) at Stelzing, Carinthia, and presented by
RoW Lloyds Esq) and another specimen from ine same
locality by W. E. Nicholson, Esq.

A series of 88 butterflies, collected (1885-1891) at various
localities in Burmah by the late Captain E. Y. Watson, was pre-
sented by the Trustees of the British Museum, together with
a specimen of Limenitis calidava captured (1892) in Ceylon
by Col. J. W. Yerbury. The data accompanying all these
specimens are precise and detailed. The additions to our collec-
tion of Oriental Danaina and Euploeina are especially useful.

Twenty-seven butterflies and 1 moth from Tobago (date
unknown), and 36 Lepidoptera from the neighbourhood of
Brisbane (1897-99), were presented by G. C. Griffiths, Esq.

Eleven insects from the Klondyke area (1901) were presented
by Professor H. A. Miers, M.A., D.Sc., F.R.S., Magdalen
College. Although the specimens are very interesting from the
point of view of locality, they appear to belong chiefly, if not
entirely, to well-known North American species.

19

Eighty-three butterflies from near Kaka and Gharb-cl-Aish
on the White Nile (1901) were presented by the captor,
W. L. S. Loat, Esq. The locality (between 10 and 11 N. Lat.)
renders the specimens of great interest. An account of this
consignment, together with another from the more southern
reaches of the White Nile, kindly sent by Mr. Loat during
the present year, will be published by Dr. F. A. Dixey.
I desire to thank A. E. Tutton, Esq., B.Sc., F.R.S., for kindly .
calling the attention of the donor of these specimens to the
needs of the Hope Department. The data accompanying
the specimens are admirably complete, and greatly enhance
their yalue.

A valuable set of 262 insects of various Orders from Kitui,
British East Africa (Ig01), was presented by the captors,
S. L. Hinde, Esq., and Mrs. Hinde. The butterflies of the
Nymphaline genera, Precis, Neptidopsis, and Charaxes, to-
gether with the Acraeinae, are specially valuable to the
Collection, but the whole of the species are most acceptable
on account of the excellent data which accompany them.

Nineteen Lepidoptera from Machakos (1901) were also
presented by the same kind donors, and these also are a very
useful addition to our tropical East African Collection.

Two hundred and eight Lepidoptera from various interest-
ing British East African localities, especially Lomogo,
N’Gongo Bagas River (1898-1900), together with 17 Lepido-
ptera from Mombasa (1900), were also presented by S.L. Hinde,
Esq., and Mrs. Hinde. All the specimens were much wanted
on account of locality and the accurate data, and many of
them are unrepresented or barely represented in the Collection.
I may specially mention a fine series of Syuchloe johnstoniz,
three specimens of J7/ylothris rubricosta, and the very
interesting series of the different forms of Lzmunas chrysippus,
and of its chief mimic the female of Hypolimnas misippus.

A specimen of a Mylabrid beetle from Sierra Leone (1858),
with the head of a Termite fixed to one of its legs, was
received in exchange from the British Museum.

The purchased Chinese and Thibetan Lepidoptera are not
catalogued.

20

The following donations are as yet uncatalogued :—the
numerous specimens from Majorca and Spain, by Professor
Poulton, Mr. Holland, and Mr. Hamm ; many specimens, espe-
cially Lepidoptera, from various localities, by Herbert Druce,
Esq., F.L.S. ; insects of many Orders from Borneo, by R. Shel-
ford, Esq., M.A.; Grecian butterflies, by W. M. Geldart, Esq.,
M.A.; a valuable and very numerous series of named American
moths, presented by W. Schaus, Esq., F.Z.S.; a set of butter-
flies from the Italian Riviera, by Hugh Richardson, Esq., and
from Germany by E. L. Meyer, Esq.; an immature Mantid
from British Bechuanaland, and a spider introduced in bananas,
by H. Ward, Esq.; a set of insects of various Orders. from
Germany and Denmark, by Professor Poulton; orficulidae
from Japan, by Harold Hornsey; a A4latta from Jamaica, by
Mr. E. Wheal; a specimen of JZelitaeca artemis from Italy, by
Miss Butler; specimens from Norway, by Colonel Swinhoe
and by E. N. Bennett, Esq., M.A.; from Europe, by H. M.
Wallis, Esq.; from Topeka, Kansas, by C. L. Pribble, Esq.

The following additions to our British Collections in 1g01
have been catalogued and incorporated :

A magnificent specimen of Vanessa antiopa (‘‘ The Camber-
well Beauty ”) was presented by the Rev. J. W. B. Bell, M.A.
The butterfly was captured, August 19, 1g00, by Mr. Bell, at
rest on a post which had been “sugared”’ for moths, in the
Rectory Gardens, Pyrton, Oxon. The capture is recorded in
‘The Entomologist” for Igco, p. 250. The specimen is of
the highest interest and value to the Department, having
been taken nearer to Oxford than any other specimen in the
University Collection, and also because of its perfectly fresh
condition. It is most improbable that such an insect could have
emerged from the pupa at any great distance from Pyrton.

Six specimens of the Noctuid moth Vonagria geminipuncta,
bred (1901) from pupae in reeds found (August, 1901) at
Bournemouth, were presented by Major R. B. Robertson,

Colonel J. W. Yerbury presented 9 specimens of Hymeno-
ptera Aculeata and mimetic Diptera, captured (1901) in
various localities in Co. Kerry. These are the first examples
of mimicry from Ireland in the Hope Collection. Colonel

21

Yerbury also presented 17 examples of models and’ mimics
from the same Orders captured (1900) in Inverness, Cromarty,
and Sutherland ; also a pair of the mimetic fly Gasterophilus
equi from Aldeburgh (1900), and 2 worker bees captured on
May 24, 1901, at Belvedere, near Woolwich, together with
a pair of the fly Stratiomys longicornis which superficially
resembles them. All the specimens were captured by Colonel
Yerbury, and bear precise statements of locality and date.

This gift constitutes a most valuable addition to our Special
Collection of the British illustrations of mimicry.

A fine collection of 378 Diptera from the Lyndhurst district
of the New Forest was presented by F. C. Adams, Esq.
All the specimens were captured (1900, 1901) by Mr. Adams,
and bear exact dates. The condition of the specimens and
their beautiful setting are such as to render the gift a most
acceptable addition to our poor collection of British Diptera,
_in which the old specimens are mostly in a bad state, and
almost invariably without data.

One hundred and fourteen British butterflies from exact
localities in Hampshire, North Devon, Herts, Berks., Oxon.,
Bucks, Suffolk, Kent, and Sussex, and from Finchley, N., and
St. Helens, Isle of Wight, were presented by Dr. F. A.
Dixey, Wadham College. The data accompanying the speci-
mens are most precise and accurate; the dates of capture
range from 1873 to1go1. The great majority of the specimens
were captured by Dr. Dixey, but some were taken by members
of his family and friends including Dr. G. B. Longstaff,
New College. The gift will be of much value to the University
Collection of British insects, a most important section of the
Department in great need of assistance.

Twelve equally valuable specimens of British Satyrine
butterflies from Berks. and North Devon were captured (1877-
1901) and presented to the General Collection by Dr. Dixey.

A set of 8 males and 53 females of the Chalcid parasite
Pteromalus puparum were presented by Professor Poulton
together with the pupa of Vanessa atalanta, in which the
parent had been seen laying its eggs (Sept. 7-8, 1900). The
parasitic Hymenoptera emerged on May 22, 1901.

22

A valuable set of 43 Lepidoptera captured (1899) by Mr.
William Reid, in Perthshire, was presented by G. C. Griffiths,
Esq., together with a series of 6 Limenitis sybilla and 4
Tephrosia crepuscularia from the New Forest (1900). Two
specimens of the former are added to the General Collection.

British insects as yet uncatalogued were also presented by
Rev. G. B. Simeon, Miss Cora B. Sanders, Major Regi:
Robertson, B. Tomlin, Esq., Mr. Mullis, H. Donisthorpe, Esq.,
W..E. Sharpe, Esq., W. -G.. Pegson -Smithy, Esq i@ueye
Bayzand, Esq., W. J. Lucas, Esq., H. Thompson, Esq., Mr.
W. Alder, Mr. T. Baines, Mr. W.'D. Rowles, Mr. AH. imu
Mr. A. Robinson, Mr. W. Holland, Mr. A. H. Hamm, as well
as by Professor Poulton and the members of his family.
A very important and valuable set of Lepidoptera from many
British localities was presented by W. C. Boyd, Esq.

ADDITIONS TO THE. HOPE LIBRARY IN Giger

The Trustees of the British Museum presented the “ Cata-
logue of the Lepidoptera Phalaenae,” vol. iii (Arctiadae and
Agaristidae), by Sir George F. Hampson.

The Smithsonian Institution (United States National
Museum, Washington) presented the publications which deal
with the subjects of the Department, including valuable
memoirs by Nathan Banks, Esq. (2 papers), Miss Harriet
Richardson, D. W. Coquillett, Esq. (3 papers), James .E.
Benedict, Esq., Jerome M°Neill, Esq., and Ralph V. Cham-
berlin, Esq.

The Boston Society of Natural History and the Bombay
Natural History Society presented their publications for the
year 1901.

The United States Department of Agriculture presented
a memoir by F. E. L. Beal, Esq., B.S., on the food of certain
N. American birds.

Reports of the following Museums, Libraries, and Natural
History Societies have been presented to the Hope Depart-
ment during the year 1901 :—

R. Accademia delle Scienze dell’ Instituto di Bologna
(Transactions, vols. vii and viii; Proceedings, vols. ii (1897-8),
iii (1898-9), and iv (1899-1g00)).

23

The Sarawak Museum.

The Bristol Museum and Reference Library.

The Kansas Academy of Science.

The Radcliffe Library, Oxford University Museum.

The Public Museum of the City of Milwaukee.

The Ashmolean Natural History Society of Oxfordshire.

The University Museum of Zoology, Cambridge (“A
Revision of the Coleopterous Family Erotylidae’’).

The Yorkshire College, Leeds.

The Annual “Report of Injurious Insects” for 1900 was
presented by the late Miss Eleanor A. Ormerod, LL.D.,
Tek Wet: Soc.

Parts I and II of the “Monograph of the Membracidae”
were presented by B. Buckton, Esq., F.R.S., F.L.S.

Dr. Samuel H. Scudder presented his “Index to N. American
Orthoptera” together with 4 papers dealing with the same
Order of insects.

« Zoological gleanings from the Royal Indian Marine Survey
Ship ‘Investigator’” (1901) was presented by the Super-
intendent of the Indian Museum, Calcutta.

Baron C. R. Osten Sacken presented 5 memoirs upon the
Diptera, including “ An Introduction to the Record of my
Life Work in Entomology.”

Seven papers upon American Lepidoptera were presented
by W. Schaus, Esq., F.Z.S.

Four papers on Lepidoptera were presented by Herbert
race lsq., F.L.S.

Four papers on Australian Crustacea were presented by
Ora sayce, Esq,

Five papers on Lepidoptera were presented by Dr. Arthur
See bubler, eh). FLAS. .F.Z.5:, together with a setiof
corrected proof-sheets of his “Illustrations of typical speci-
mens of Lepidoptera Heterocera in the collection of the
British Museum ” (1886) and many plates and coloured draw-
ings of Lepidoptera.

A valuable series of 15 papers on the economic entomology
and vegetable pathology of Australia, from the “ Queensland

Agricultural Journal,’ were presented by the author, Henry
Tryon, Esq.

ee the for Walter Ropechiid: Ph. ND. a and 2
Jordan, Ph.D.

year I90I, were presented by Professor Poulton. .

Copies of original papers on the Arthropoda h
presented by the following authors:—George H. F.
Esq., M.A., D.M., Ph.D., and Arthur E. Shipley, Es
(1 paper); W. L. Distant, Esq. (2 papers) ; Gilbert J
_ Esq., F.E.S. (3 papers) ; Miss oe Evans ee Pape
Bignell, Bets F; E. Str ee

staff, Esq., D. M., Bak G: P).
The parts of Barrett’s “ British Lepidopties
Society volume and the volume of the Zoe

49

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