THE HOPE REPORTS

VOL, Ut

1897—1900

» EDITED BY

EDWARD B° POULTON, M.A.. D.Sc., F.RS.
Hon. LL.D. PrincEron

FELLOW OF JESUS COLLEGE, OXFORD
HOPE PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF OXFORD

Orford
PRINTED FOR PRIVATE CIRCULATION
RY HORACE HART, PRINTER TO THE UNIVERSITY

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THE HOPE REPORTS

VOT

1897—1900

EDITED BY

EDWARD B. POULTON, MA. DSc, F.RS.
; Hon. LL.D. Princeton

FELLOW OF JESUS COLLEGE, OXFORD
HOPE PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF OXFORD

Orford
PRINTED FOR PRIVATE CIRCULATION

BY HORACE HART, PRINTER TO THE UNIVERSITY

IQOI

PRE A Cly

TureE and a half years have elapsed since the first
volume of the Hope Reports was issued, in June 1897.
During this period the materials for the second volume
have been gradually accumulating, representing certain
important sides of the work done in the Department, or
upon portions of its collections which have been removed
for study and comparison with those of other museums.
I had hoped that this second volume would have been
ready in a somewhat shorter time, but other sides of our
work have prevented an earlier appearance. The Reports
of the Department for the years 1897, 1898, and 1899
(Nos. 21, 22, and 23) sufficiently indicate the nature of
this work, and prove that much labour has been expended
upon the manipulation, naming, and systematic arrangement
of this vast collection, together with the numerous accessions
due to the kindness of naturalists in many parts of the
world, A large amount of cataloguing and the printing of
many thousands of labels have also been rendered necessary
by the new material. Furthermore, a considerable pro-
portion of the papers in this volume are abstracts of
much longer and more detailed memoirs, many of which are
far advanced towards completion. This is the case with
the papers of which Nos. 4, 7, 8, 9 and 10 are abstracts:
these, when published, will contribute an important share to
the third volume of the Hope Reports.

4 ~ PREFACE

The contents of this volume have been arranged accord-
ing to their subjects, the first ten papers (omitting 3c)
being almost exclusively concerned with Mimicry, Warning
Colours, and the Struggle for Existence in Insects, Nos. 10 to
13 with seasonal dimorphism, and the colour of pupae, &c.,
as influenced by the surroundings of the larvae, No. 14
with Hybridisation, No. 14 4 with the ‘Organic Selection’
of Baldwin, Lloyd-Morgan and Osborn, Nos. 15 to 19 with
systematic work upon parts of the existing Hope Collection
or new material which has recently been added. It should
be noted that Mr. Champion’s paper on the Zzxgztidae,
No. 15, is founded on the study of material from many
collections: it is included here because a large proportion
of the types of the new species described are in the Hope
Museum. Nos. 3 c and 20 deal with the manipulation and
labelling of insects for collections, while Nos. 21 to 23 are
the Reports of the Hope Professor for the years during
which the materials for this volume have been accumulating.
I venture to hope that the contents of the volume indicate
some activity and energy in the Department. Great
encouragement has been received from the manner in which
naturalists in many countries have looked to the Hope
Museum as a centre for the study of insect bionomics, and
have sent material bearing on these difficult and deeply
interesting problems to Oxford. When the specimens now
rapidly accumulating have been arranged, the student will
have unique opportunities for the study of Mimicry, Warning
Colours, Protective Resemblance, and allied questions. Nor
will the arrangement be long delayed, for the chief part of
the printing, labelling, cataloguing, and preliminary group-
ing has already been done.

One chief duty of our ancient University is to contribute
its share, and may it be a very large one, towards winning

PREFACE 5

fresh territories from the unknown. This Department
would not be doing its duty to the University, to the
memory of its Founder, and to that of the first Hope
Professor, if some advance were not made in that part of
the boundaries of knowledge which stands opposite to it.
I would that every College, and every University depart-
ment, felt this responsibility as keenly as it is felt in
a happily increasing number of Oxford laboratories and
scientific departments. I do not wish to be misunderstood
in making these high claims for the exponents of the
natural and physical sciences. Our scientific men are
fighting side by side with the Oxford students of other
branches of learning. The interests of the ancient studies
in Oxford are sometimes supposed to conflict with the more
modern. But the real contest is between those who would
and those who would not devote their energies to widening
the boundaries of knowledge, and there can be nothing but
comradeship and the warmest sympathy among the men
whose keenest efforts are put forth in order that Oxford
may be known far and wide as a centre of research and
learning.

EDWIN eB: POULTON:

Hore DEPARTMENT OF ZOOLOGY,
University Museum, Oxrorp,

January 4, got.

CONTENTS OB) Orn

t. Preface.

2. Mimetic Attraction, by Frederick A. Dixey, M.A., M.D., F-E.S., Fellow
of Wadham College, Oxford. (From ‘ Transactions of the Entomo-
logical Society of London,’ 1897, p. 317.)

3. Abstract of the above Paper, and discussion which followed it. (From
‘Proceedings of the Entomological Society of London,’ 1897,
pp. XX-Xxxii, xxxiv—xlvii.)

3 4. Mimicry in Butterflies of the Genus Hypolimnas, and its bearing on
older and more recent Theories of Mimicry, Abstract, by Edward B.
Poulton, M.A., F.R.S., Fellow of Jesus College, Oxford, and Hope’
Professor of Zoology in the University. (From ‘ Proceedings of the
American Association for the Advancement of Science, 1897,
vol, xlvi, p. 242.)

3.8. Mimicry as evidence for the ancestral home of a wide-ranging species,
Abstract, by Edward B. Poulton. (From ‘Proceedings of the
American Association for the Advancement of Science,’ 1897,
vol. xlvi, p. 244.)

3c. A method of labelling type specimens in Collections of Insects,
Abstract, by Edward B. Poulton. (From ‘Proceedings of the
American Association for the Advancement of Science,’ 1897,
vol. xlvi, p. 244.)

4. Theories of Mimicry, as illustrated by African Butterflies, Abstract, by
Edward B. Poulton. (From Report of the Meeting of the British
Association at Toronto, Section D, 1897, p. 688.)

5. Protective Mimicry as evidence for the validity of the Theory of
Natural Selection, Abstract, by Edward B. Poulton. (From Report
of the Meeting of the British Association at Toronto, Section D,
1897, p- 692.)

CONTENTS: OF VOL IL 7

10,

II.

FS:

14.

_ Natural Selection the Cause of Mimetic Resemblance and Common

Warning Colours, by Edward B. Poulton. (From the ‘ Linnean
Society’s Journal—Zoology,’ vol. xxvi, 1898, p. 558.)

. An Experimental Enquiry into the Struggle for Existence in certain

Common Insects, Abstract, by Edward B, Poulton and Cora B.
Sanders. (From Report of the Meeting of the British Association
at Bristol, Section D, 1898, p. 906.)

. Observations on Mimicry in South African Insects, by Guy A. K.

Marshall, Abstract, arranged by Edward B. Poulton. (From Report
of the British Association at Bradford, 1900, Section D.)

. Observations on Mimicry in Bornean Insects, by R. Shelford, B.A.,

Curator of the Sarawak Museum, Abstract, arranged by Edward B.
Poulton. (From Report of the British Association at Bradford, 1900,
Section D.)

Guy A. K. Marshall’s proof that Prec?’s ocfavia is the summer form of
P. sesamus, Abstract, by Edward B. Poulton. (From ‘ Proceedings of
the Entomological Society of London,’ 1898, p. xxiv.)

Abstract of F. Merrifield’s paper on ‘The Colouring of Pupae of
P. machaon,’ &c. (see below, No. 12, for the complete memoir), and
discussion thereon, (From ‘Proceedings of the Entomological Society
of London, 1898, p. xxx.)

. The Colour-relation between the pupae of Papzlio machaon, Pieris napt

and many other species, and the surroundings of the larvae ‘preparing
to pupate, &c., by F. Merrifield, F.E.S., and Edward B. Poulton.
(From ‘ Transactions of the Entomological Society of London,’ 1899,
p- 369.)

Seasonal dimorphism in tropical American Pierine butterflies, Abstract,
by Frederick A. Dixey. (From ‘Proceedings of the Entomological
Society of London,’ 1898, p. xxxix.)

Recent Experiments in Hybridisation, by Frederick A. Dixey. (From
‘Science Progress,’ vol. vii: vol. ii of New Series: No. 7, April, 1898.)

144. Modification and variation and the limits of Organic Selection, Abstract

ree

of paper by Professor Henry F. Osborn, of Columbia University,
New York, and of a criticism thereon, by Edward B. Poulton. (From
‘Proceedings of the American Association for the Advancement of
Science,’ 1897, vol. xlvi, p. 239.)

Notes on American and other Tingitidae, with descriptions of two new
genera and four species, by George Charles Champion, F.Z.S.
(From ‘Transactions of the Entomological Society of London,’
1898, p. 55:)

16.

17.

18.

IQ.

20.

21.

22.

23.

CONTENTS “OF: VOL. Ti

Descriptions of some new species of Syntomidae, chiefly in the Oxford
Museum, by Herbert Druce, F.L.S., &c. (From the ‘ Annals and
Magazine of Natural History,’ Ser. 7, vol. i, May 1898, p. 401.)

On a Collection of Insects and Arachnids, made by Mr. E. N. Bennett
in Socotra, with descriptions of new species, by F. A. Dixey, M.A.,
M.D., Malcolm Burr, F.Z.S., and the Rev. O. Pickard-Cambridge,
M.A., F.R.S., C.M.Z.S. (From the ‘ Proceedings of the Zoological
Society of London,’ 1898, p. 372.)

On a Collection of Insects and Arachnids made in 1895 and 1897 by
Mr. C. V. A. Peel, F.Z.S., in Somaliland, with descriptions of new
species by C. V. A. Peel, F.Z.S., E.E. Austen, P. Aj Dixeys Was
M.D., Herbert Druce, -F.L.S., F-Z.S., C. J. Gahan, M-A., Gilbert J.
Arrow, R. McLachlan, F.R.S., Malcolm Burr, F.Z.S., and R. I.
Pocock. (From ‘ Proceedings of the Zoological Society of London,’
1900, p. 2.)

On the British Orthoptera in the Hope Museum, Oxford, by Malcolm
Burr. (From the ‘Entomologist’s Record,’ &c., vol. xii, April 1900,
P. 97.)

The methods of setting and labelling Lepidoptera for Museums, by
Edward B. Poulton. (From the Report of the Proceedings of the
Museums Association, Oxford, 1897, p. 30.)

Report of the Hope Professor of Zoology for 1897. (From the
‘Oxford University Gazette.’)

Report of the Hope Professor of Zoology for 1898. (From the
‘Oxford University Gazette.’)

Report of the Hope Professor of Zoology for 1899. (From the
‘Oxford University Gazette.’)

@ size)

XIII. Mimetic Attraction. By Frepericx A. Dixey,
M.A., M.D., F.E.S., Fellow of Wadham College,
Oxford.

[Read May 5th, 1897.]

Pirate VII.

In a former contribution* to the Transactions of this
Society, I endeavoured to trace, by means of actual
examples, the successive steps through which a complicated
and practically perfect mimetic pattern could be evolved
in simple and easy stages from a form presenting merely
the ordinary aspect of its own genus. In the present
paper I propose to enter somewhat further into the
subject of mimetic change, and in the first place to show
how the process of gradual assimilation, starting from
one given point, may take not one direction only, but
several divergent paths at the same time; in other
words, how the members of a single group may assume
several different mimetic developments, each one corres-
ponding to a distinct model, but all derived by easy
stages from the same original form.

In the paper just referred to it was shown that a very
complete transition could be demonstrated, by means of
closely-allied and still existing species, between an
ordinary Pieris such as P. phaloe, presenting only the
usual features of its genus, and a form of such widely
different aspect as Mylothris pyrrha @ ; the latter being
a nearly exact copy of Heliconius nuwmata. But although
these facts are sufliciently striking, it is perhaps stall
more remarkable that from the same or closely allied and
very similar forms of typical Pierine aspect, at least four
other lines can be traced, each showing almost as perfect
a transition as that from P. phaloe to M. pyrrha, and
each leading up to a presumably distasteful model ; these
models being in appearance entirely different from H.
numata and from each other.

* “On the Relation of Mimetic Patterns to the Original Form,”
Trans. Ent. Soc. Lond. 1896, pp. 65-79, pls. ITI.—V.

TRANS, ENT. SOC. LOND. 1897.—PparT I. (SEP'.)

318 Dr. Frederick A. Dixey on

A full illustration of this position would require a
larger series of diagrams than could well be given in this
place; it is hoped, however, that the series of figures on
Plate VII. may suffice to show the more important
links in each chain, or at any rate to make the drift of
my statements intelligible. It should be borne in mind
that all the forms here spoken of are neotropical.

1. The “agna” line. The first of these lines of
development may be called the agna line, inasmuch as it
leads towards a well-marked group of distasteful forms
of which Aeria agna, Godm. and Salv., (Pl. VIL., fig. 6) is
a good example. Starting from a white Pieris of ordinary
aspect like P. phaloe, we find that the first step in the
growth of this mimetic pattern is furnished by the pro-
longation of the diagonal dark bar, which in P. phaloe ? *
extends from the costa to the distal end of the cell in
the forewing, to meet the dark hind margin as in P. caly-
donia § (fig. 1, kh). All stages in the development: of
this first feature can indeed be traced by comparing
specimens of both sexes of P. calydonia itself. The next
step is the extension of tne dark hind border of the
forewing, already more pronounced in P. calydonia 2
than in P. phaloe 2, along the inner margin, as in P.
demophile 2 (fig. 3, l). Simultaneously with this change
the dark border of the hindwing is much broadened
(ib., 2), and in some specimens of P. demophile 2, as in
the one figured, the white ground colour is replaced by
yellow. ‘These changes are sufficient strongly to suggest
the general aspect of the protected group referred to,
and it seems difficult to believe that the appearance of
the yellow female of P. demophile has not a significance
derived from this fact.

2. The “ atthis”’ line. But the last-named form, viz.,
the yellow P. demophile 2 , though the final Pierine term
in one transitional series, is but an intermediate term in
another. This second line of development, starting
afresh from P. demophile ° , passes into an unmistakable
mimetic relation with the protected group that centres
round such forms as Heliconius atthis (fig. 8) and
Tithorea pavonit.

Comparing P. viardi 2 with P. demophile &, we find
that in the former insect a further stage of divergence

* See Trans. Ent. Soc. Lond. 1896, pl. TIT.; fig. 2, z.

Mimetic Attraction. 319

from the original Pierine form has been reached (1) by
a slight modification in shape and proportions of that
area of the pale ground colour included between the
diagonal bar and the dark inner margin of the forewing,
and (2) by the appearance of a submarginal series of
pale spots on both fore and hindwing (fig. 5,7). Both
of these changes mark an approximation towards the
characteristic aspect of the protected Heliconius atthis
group ; but the final link in the Pierine chain, viz., the
form represented in fig. 7, which appears to be the female
of P. locusta or of a closely allied species, brings us
nearer still. Here the pale area on the apical side of
the diagonal bar (nv) shows a tendency to be broken up
into separate spots ; and the pale ground colour, which
in P, viardi 2 is uniformly yellow, in P. locusta 2 only
retains the yellow tint proximally, the subapical area (7)
and the submarginal spots (7) having resumed their
original white. These submarginal spots have also
diminished in size and become more compact in outline.
In all these respects a further approach is made towards
H. atthis, and the whole series from P. phaloe or P. caly-
donia to P. locusta 2 and H. atthis affords as striking a
succession of transitional forms as that before traced from
the same ordinary Pierine types up to Mylothris pyrrha ?
and Heliconius nwmata.*

3. The “inachia” line. In this line an early step in
advance of the usual Pierine pattern is taken by Pieris
pandosia.t On the hindwing underside of this species
appears a narrow chestnut-coloured streak in the midst
of the dark hind-marginal band, running paraliel with
the hind border of the wing. In P. leptalina, Bates (=
P. pisonis, Hew.), both surfaces of the forewing show a
diagonal dark bar as in P. calydonia ¢ and other species ;
there is also a transverse dark bar crossing the under-
surface of the hindwing nearly parallel with the costa
(fig. 2). In P. pandosia, where the distal end only of this

* Trans. Ent. Soc. Lond. 1896, pp. 65-70. It should be added
to the foregoing that P. viardi 9 itself is probably attracted by
H., charitonia, which may perhaps be considered an outlier of the
H. atthis combination. This is analogous to the attraction of
another intermediate term in the atthds line, viz., P. demophile 9,
by the agna group.

+ Figured by Hewitson, Exot. Butt., “ Pieris,” pl. IL., fig. 14,
and ibid., pl. V1., fig. 39.

320 Dr. Frederick A. Dixey on

latter bar is indicated, the chestnut-coloured streak takes
no part in its formation; in P. leptalina, however, the
same chestnut-coloured streak is better pronounced along
the hind margin than in P. pandosia, and is prolonged
for a short distance along the transverse bar (fig. 2, 0).
In this way a considerable resemblance is brought about
between P. leptalina and some of the well-known trans-
parent neotropical Danainz, which are presumably
protected, such as Napeogenes inachia (fig. 4). It is
worth noting that the bar o, which belongs to the hind-
wing in P. pandosia and P. leptalina, appears to
represent the dark inner margin of the forewing in the
model; and the presence upon its distal portion in P.
leptalina of a shght prolongation of the chestnut marginal
streak may perhaps stand for the turning inwards of the
corresponding chestnut marginal band in N. inachia
along the costal border (figs. 2 and 4).

4. The “numata” line. This series needs here no
more than a mention, as it has been already discussed
and illustrated in the paper above referred to.*

5. The “tarracina”’ line. This is an offshoot of the
last, or nuwmata line, diverging in the neighbourhood of
Mylothris malenka § , or perhaps somewhat further back
towards the original Pieris. ‘The remarkable form M.
alethina 2 shows a near approach to the pattern of
Tithorea tarracina and other associated species; its own
aspect being probably derived from a type like M. lovena ?
or M.malenka 2, im which the yellow of P. demophile &
has persisted, and the base of the forewings has become
overspread with black.t

General Considerations. There is, therefore, in each one
of these cases, a continuous line to be traced; starting
in every instance from the same ordinary Pierine form,
and passing through a graduated series of closely-allied

“ Vide supra, p. 319, note.

+ Three at least of the above five lines of development, viz.,
the agna, the numata, and the inachia lines, can be paralleled from
the genus Dismorphia and its allies, and it is interesting to see in
the latter case how the same ultimate result is brought about by
somewhat different means. But I have not attempted to include
species of Dismorphia within the above series, because the affinity
of this genns with Piers and Mylothris is not close, and its own
course of mimetic development, so far as phylogeny is concerned,
must be regarded as completely independent.

Mimetic Attraction. 5 7Al.

species untilit terminates in a Pieris or Mylothris bearing
an intimate mimetic relation with some insect of entirely
different affinities. It is further to be observed that, in
every instance, the species here considered as the model
towards which these diverging series tend, does not
present an isolated and independent scheme of coloration,
but is itself a member of a larger or smaller group of
forms, in addition to the Pierine mimic, all of which are
endowed with an aspect similar to itself—in other words,
that the mimetic associations do not run simply in pairs,
but in groups. This latter fact has long been recognised ;
and the existence of such mimetic groups has been
shown by F. Miiller, Meldola, and Poulton to possess
a further significance than that originally detected by
Bates. The elaborate work of Haase* contains an
attempt to give a systematic account of the chief cases
of mimetic grouping. But in spite of what has already
been written by these and other authors, it may be
doubted whether the importance of the principle of
mimicry among the factors that have determined the
facies of the insect fauna in such a region as the neo-

* “Untersuchungen tiber die Mimicry,” Stuttgart, 1893. It may
here be mentioned that several of the above-named insects have
been noticed by Haase ; who, however, has not attempted to trace
in any detail the lines of mimetic assimilation that diverge from the
common Pierine stock. He speaks, for instance, of MWylothris lorena
Qand M. malenka 9 , which undoubtedly belong to the nwmata group,
as having arisen from such forms as P. demophile 2 , and considers
that the transition took place through forms resembling P. viardi
Q. Buta careful examination will I believe show that, as stated
above, neither P, demophile 9 nor P. viardi ? is in the direct
line passing from the unaltered Pieris towards M. pyrrha and
Heliconius numata. Both are, in fact, intermediate terms in the
series leading up to an entirely distinct assemblage, that typified
by Tithorea pavonii and Heliconius atthis (Haase’s “ Bonplandi
Tracht”); while P. demophile shows evidence of attraction by the
protected agna group, and P. viardi by the dominant form Heli-
conius charitonia. Again, the red streak on the underside of the
hindwing in the males of M. lorena, etc., is attributed by Haase to
“inheritance from the female,” but its origin is not traced by him
to the primitive basal red common to many Pierine genera. (See
Trans. Ent. Soc. Lond. 1894, pp.283-289 ; zbid., 1896, pp. 72, 73.)
Pieris leptalina, Bates (P. pisonis, Hew.) is spoken of by Haase as
representing the first partial assimilation to certain Ithomias ;
rightly, so far as the main fact is concerned ; but it may also be
noticed that earlicr stages of the same assimilation exist in P.
kiguha, and, as shown above, in P. pandosia.

d22 Dr. Frederick A. Dixey on

tropical has ever been fully recognised. The mimetic
groups here referred to are no doubt mainly of the
“ Miillerian ” kind; that is to say, they are associations
between inedible species of various affinities ; each asso-
ciation possessing a conspicuous and distinctive pattern
of its own, more or less perfectly reproduced by all its
members. In the paper already cited and elsewhere* I
have given reasons for considering the principle of Miil-
lerian mimicry to be far more widely operative than has
generally been supposed ; and I have also endeavoured
to supply a test by which, even in the absence of
information as to the edibility and relative abundance of
the members of a mimetic group, a conclusion may some-
times be arrived at as to whether the assemblage is
Batesian or Miillerian. As there are grounds for suppos-
ing that the arguments just referred to have sometimes
been misunderstood, it will be attempted in the following
section to re-state, in as simple language as possible, what
appears to be the best interpretation of the facts at present
known.

Mimetic Attraction. When a species of butterfly has
become established in such a region as the neotropical,
where life of all kinds is very abundant and competition
extremely keen, it may be taken for granted that the
species possesses some efficient means of defence, failing
which it would be unable to maintain its position. In
very many instances, as is well known, the required
protection is essentially afforded by the possession of a
nauseous flavour, which causes the butterfly in question
to be avoided, when recognised, by some at least of its
insect-eating enemies. ‘The possibility of easy recogni-
tion in such a case constitutes, of course, an important
factor in the safety of the species ; since there would be
no advantage in being inedible, if the fact only became
known in each individual case as the result of an
experiment fatal to its subject. It is in consequence of this
necessity for ‘advertisement ” that, as is also well known,
inedible species tend to assume gaudy and conspicuous
colours, and to adopt habits calculated to display their
warning signals with the utmost publicity. In this
manner the members of a distasteful and conspicuous
species are enabled to profit by the experience gained at

* Trans, Ent. Soc. Lond. 1894, pp. 297, 298 ; ibid., 1896, p. 75.

Mimetic Attraction ‘ 323

the expense of other individuals of the same species, some
of which must necessarily fall victims to their insectivorous
enemies during the “ education ” of the latter as to the
kinds of prey to be sought or avoided. Hach form that
thus succeeds in establishing itself becomes, in propor-
tion to its nauseous character and the ease with which it
can be recognised, a centre of attraction for other species,
which, by assimilating their own aspect to that of their
model, are enabled to share in the immunity from attack
enjoyed by the latter. It is now considered probable by
many that this power of attraction is exercised by
dominant inedible species over edible and inedible forms
alike; there is, however, one important difference be-
tween the two cases, which I have before endeavoured to
point out, but which seems to need a more explicit state-
ment than has yet been given to it.

Let us first take the case of a species which is edible,
and therefore liable to extermination by insectivorous
animals. The chances of the survival of such a species
depend on the excellence of its means of defence ; such as
superior swiftness, or the power of concealment, whether
by resemblance to inanimate objects, or to some other
species protected by a disagreeable flavour. The force
which impels an edible species to seek protection by the
last-named method, viz., by sheltering itself under the
reputation of a conspicuous anedible form, is the well-
known ‘“ Batesian mimicry.” With reference to this,
which is the most complete kind of mimicry, it is to be
observed that the advantage of association is all on the
side of the mimic, and is not shared in the least degree by
the model. Indeed, the existence of the edible mimicking
species is a source of danger to the form mimicked,
inasmuch as any experience gained by tasting the former
would be used to the detriment of the latter. From these
considerations two consequences follow; the first being
that such an association can subsist only when the
numbers of the mimic are insignificant compared with
those of the model, for otherwise the latter’s reputation
for inedibility would be interfered with, and eventually
destroyed. The second consequence is that the attrac-
tive force leading to assimilation between the two forms
can act only in one direction ; 7.¢., the model attracts the
mimic, but the mimic can exert no reciprocal influence
upon the model. The latter stands secure upon its own

324 Dr. Frederick A. Dixey on

footing, any departure from which might be attended with
danger to itself; while the only part played by the former
is to shelter itself under as close an approximation to the
aspect of the model as circumstances permit.

We may now take the case of a species possessing
a nauseous flavour, and requiring some means of adver-
tisement in order to make its inedibility available for
purposes of protection. ‘'T'wo courses may be said to lie
open for such a species. First, it may seek advertise-
ment by acquiring a conspicuous and easily recognised
aspect of its own, distinct from that of any others; or,
secondly, it may obtain a share in the notoriety already
attaching to some dominant inedible form by assimilating
its aspect to that of the latter, instead of striking out a
new line for itself. An examination of the lepidopterous
fauna of such a region as the neotropical makes it certain
that the latter of these methods, viz., the method of
‘ Miillerian mimicry,” has been very extensively fol-
lowed. Its advantages, as compared with the former
method, are obvious. In the first place it assists the
memory of predaceous foes by keeping at a low figure the
number of distinct inedible types to be learned and so
avoided, and in the second place it benefits at least two
species at the same time instead of one, and both have
therefore an interest in keeping it up; for inasmuch as in
this case, as distinct from that of Batesian mimicry, the
mimic is inedible as well as the model, the results of
experimental tasting will be uniformly the same, and will
be favourable to the immunity of both species. It
follows that (1) there is no such limit as exists in
Batesian mimicry to the number either of individuals or
species forming a Miillerian group. An assemblage of
this latter kind is only strengthened, not weakened, by
fresh accessions; all being alike inedible, and so all
contributing to the common safety. (2) The benefit
of Miillerian association being mutual, there is a distinct
reason, which we saw does not exist in the case of
Batesian mimicry, for the model to help on the process of
assimilation by itself advancing to meet the mimic.

To summarise the foregoing. 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

Mimetic Attraction. 325

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.

This doctrine of the mutual attraction between inedible
forms, leading not merely to the copying of one by
another, but to the departure of each from its original
aspect by the adoption of features belonging to the other,
is not simply a speculation, nor does it rest only on
a@ priort reasoning. There is much evidence that
it represents a fact which does actually take place in
nature ; and in the two papers above referred to* I have
brought forward cases which seem imexplicable by any
other principle. I may be allowed to add in this place a
further instance, which appears to me for several reasons
remarkable.

The instance of P. locusta §. In Trans. Ent. Soc.
Lond. 1896, p. 72, note, I spoke, though somewhat
doubtfully, of P. locusta § as a mimic. My suspicion
that this was the case at that time fell short of actual
conviction. Now, however, after a further careful
examination of P. locusta $ from this particular point
of view, I have little or no hesitation in pronouncing it to
be a member of a mimetic association of an exceptionally
interesting kind.

It will be remembered that the males of Mylothris
lorena, M. pyrrha, etc., form a good illustration of the
accurate manner in which the appropriate habits are
correlated with adaptive colouring—the mimetic pattern
in these instances being confined to the underside, and
being in all probability useless as a protection except
during the resting position; while the habits of these males,
as testified to by Wallace,+ are such as would probably
render a Heliconiine resemblance during flight a source of
danger to its possessors rather than of safety. Similarly
in P. locusta ~, it is only on the underside that the
mimetic pattern appears, and here again there can be

* See also an abstract in “ British Association Reports,” 1894,
p. 692.

t+ “ Tropical Nature,” 1878, p. 205. See also Haase, op. cit.,.
p. 68. It should, however, be stated that neither of these authors
assigns any protective value to the underside of the male forms in
question.

326 Dr. Frederick A. Dixey on

little doubt that its use has reference only to the resting
position.

The aspect suggested is rather that of several forms of
Heliconius in general than that of any one in particular ;
the most definite relation, however, is with the group of
which Heliconius cydno and H. galanthus are examples, a
group characterised by the presence on both surfaces, but
especially beneath, of a good deal of light yellowoften paling
further to white. When the undersides of Preris locusta
and one of these Heliconw, say H. cydno (which inhabits
the same part of the neotropical region), are compared
together, both insects being in the resting position, we
find not indeed an exact resemblance, but a general
similarity which, judging from other instances, we may
consider as probably sufficient to suggest the possession of
like qualities. In both there occurs on the forewing a
certain amount of white ground colour; on one side
bounded by a dark tip, and on the other more or less
limited by the dark area of the hindwing ; which latter
is traversed by a bright yellow streak, and beset about
the base with red or chestnut patches. The elements of
the pattern in both insects are the same, and their
general relation to one another much alike, though the
pale marginal band on the hindwing that occurs in
this species of Heliconius is not found in the Pierine.
A further point of interest is the manner in which
the partial breaking up of the white colour is effected
in the Heliconius and the Pieris respectively. There
is, I think, little doubt that the dark marks on
the costa of the hindwing in P. locusta represent
m a general way the dark discoidal spot and a
portion of the inner dark area of the forewing in the
Heliconius. The bright yellow line of the hindwing will
be seen in each case to terminate in relation with a dark
patch; but in the first instance (that of the Pieris) this
belongs to the same wing; and in the second instance
(that of the Heliconius) it is contributed to by both
wings. This seems to exemplify a principle repeatedly
met with in mimicry ; viz., that exact homology is dis-
regarded, and the whole exposed surface of the insect is
taken as it were as a canvas on which the mimetic
picture is painted with a free hand. The relation of this
particular group of Heliconius with the Pierines is
remarkable, and deserves more detailed treatment; here

Mimetic Attraction. oe

it will be sufficient to point out that there is more reason
to suppose that the Heliconius has adopted certain
features from the Pieris (for example, the whiteness of
the ground colour, and the disposition, if not the exist-
ence, of the basal red marks) than that the converse
alone has taken place.

Reciprocal Mimicry and Convergence. ‘This fact of the
reciprocal copying of two or more species by each
other is perhaps implied, though not distinctly so,
in the term ‘ convergence,” which has been used
by many authors to express the phenomenon of Miil-
lerian mimicry; but I am not aware that any writer
who so employs the term has laid stress on the mutual
character of the changes involved, or has traced in
any instance the actual modifications undergone by
both species of a Mullerian couple under the influence
of the attractive force existing between them. It seems
hitherto to have been taken for granted that a dominant
form will attract or retain other species within its own
sphere of influence, without being itself attracted in
return ; whereas the fact is, as we have seen, that each
member of an inedible association has more or less
influence upon all the rest. The respective value of the
attraction exercised or suffered by any member of a
Miillerian group will depend on its numbers, its nauseous
qualities, and its notoriety. The stronger any species is
in these respects, the stronger will be its power of
attraction, and the weaker in comparison will be any
force tending to draw it in the direction of other mem-
bers of the group. The actual mimetic path taken by
any species will be the resultant of the various forces
acting uponit. If the form happens to be a dominant one,
these external forces will be insignificant in comparison
with its own stability ; and it will therefore resist change
to a large extent, or perhaps altogether. The most com-
plete intermingling of characters given and taken on
both sides may be expected when two species meet on
equal terms, neither being strong enough to predominate
over the other.

While there can be no doubt of the convenience of the
term “convergence,” and its suitability to express
relations of the kind just discussed, there would seem to
be no sufficient reason for disallowing in their case the
earlier term “mimicry.” ‘This latter word may be quite

328 Dr. Frederick A. Dixey on

legitimately employed to designate the adoption of any
new features borrowed from another species. In so far as
A copies the appearance of B, it may properly he said to
“mimic” the latter, whether the object be to suggest
the presence of a disagreeable flavour which it does not
really possess, or merely to convey the impression
that it and its model are alike in all respects. Muil-
lerian assimilation may be quite as deceptive as Batesian,
in the sense of leading to confusion between species
essentially distinct ; and in the case of a “ weak’’ species
being associated with a “strong” one, the departure of
a form from the typical aspect of its congeners by the
development of strictly imitative features may be as well
marked in the one kind of mimicry as in the other. But
although either of the terms ‘ Miillerian mimicry ” or
‘convergence ” would appear to express quite adequately
the general idea of the mimetic relation between inedible
species, a separate term is wanted to designate the
peculiar give-and-take changes which we have seen are
theoretically possible to a greater or less extent in every
case of Miullerian association, and which in fact do actually
occur in several. It is to supply this want of a term that
I have proposed the expression “ reciprocal mimicry,’”*
which is meant to convey, besides the general idea of
convergence, the special information that in the cases to
which the term is applied, the convergence is brought
about not by the simple imitation of one form by another,
but by the interchange of features between forms, and
their consequent simultaneous approach to an intermediate
position.

The foregoing remarks will, I think, have made it
sufficiently clear, (1) that reciprocal mimicry can only take
place in Miillerian associations, not in Batesian; and that
it 1s therefore, as I have elsewhere said “ good evidence
of the distastefulness of all the forms between which it
can be shown to occur;” (2) that although a mimic
which is of relatively plentiful occurrence must be Miil-
lerian, it does not follow that a mimic which is scarce
must necessarily be Batesian. An inedible mimic may
be either rare or common ; an edible mimic must be rare.
Judging by these principles, we must conclude that the
association of Pieris locusta f$ with Heliconius cydno is

“ Trans. Ent. Soc. Lond. 1894, p. 298.

Mimetic Attraction. 329

Miillerian, and an addition is thus supplied to the evidence
already existing in favour of the distasteful qualities of
this and other Pierine genera.*

Conclusion. If we take a comprehensive survey of the
whole butterfly facies of the neotropical region, we cannot
fail to be struck with the numerous cases of mimetic assimi-
lation which it presents. It would in fact almost seem
that scarcely any conspicuous form is completely isolated.
In a region where enemies to insect life are so numerous,
and competition for existence is so keen, a butterfly can
hardly afford to be conspicuous unless it is also distaste-
ful; nor is the mere individual possession of these
qualities in the majority of instances sufficient for safety.
Any form that requires to establish a reputation for
inedibility, must as it were seek allies; and no sooner does
it make a bid for survival as a nauseous species, than it
becomes subject to the influence of mimetic attraction,
and probably finds itself drawn into the vortex of one of
the great Miillerian associations.

It is no doubt true that the process of mimetic assim1-
lation is subject to limitation by the operation of other
forces. Thus it may be needful that the resemblance to
a model, though close enough to deceive enemies, should
not be so close as to interfere with due recognitzon
between the sexes. We know little of the means by
which insects recognise each other, but there are at least
some grounds for thinking that they are assisted at times
by external marks. ‘his may atford one reason for the dif-
ferent ways in which the sexes of the same species occasion-
ally react to mimetic influences ; and it may possibly be
the meaning of the retention of a portion of the original
white ground-colour in the males of certain species of
Dismorphia. It is also quite conceivable that the aspect
of every species is to some extent controlled by its
physiological constitution ;+ and this fact may tend to
disturb the perfect operation of the process of mimetic
change.

But notwithstanding all limitations, it remains the
fact that any inedible form, in the midst of competing

* Vide Trans, Ent. Soc. Lond. 1894, pp. 297, 298; ibid., 1896,
pp. 74, 76.

+ Vide Presidential Address by Prof. Meldola, Proc. Ent. Soc
Lond, 1896, pp. Ixxx. et seqq.

330 Dr. Frederick A. Dixey on

mimetic systems, is liable to be carried off in this or that
direction as it comes under their influence. Sometimes,
as in the forms that were discussed at the beginning of
this paper, we find several species, closely related to one
another by affinity, being drawn away in different direc-
tions by the attractive power of the Miillerian groups
that surround them; and the same assemblage of cases
illustrates the fact that a mimetic change from the original
form in the direction of one protected group may serve
as a stepping-stone for a further de ane towards
another.* Sometimes again, as in Mylothris lypera, M.
lorena, M. malenka, and M. pyrrha, we see that in the
males a compromise is struck between the ordinary
Pierine aspect (used for flicht), and a mimetic dress like
that of the female (used for repose); while in Pieris
lucusta we find the same compromise in the male, with
the curious difference that here even the sexes of the same
species have been wrested apart into separate mimetic
relations.

Finally, the comparison will perhaps not seem too far-
fetched, if the several mimetic groups, each with its own
type of coloration, are likened to the solar and stellar
systems of astronomers. Sometimes, as in the solar
system, there is one central body (7.e., species) dominat-
ing the whole, and influencing its attendant planets (7.e.,
mimics) to an extent im comparison with which the force
they themselves can exercise is insignificant. At other
times, as in the systems of double and multiple stars,
there are bodies (7.e., species) more nearly equal in mass
and importance, bound together by mutual attraction
into a single combination, where each one effectively
controls and is controlled by the rest. We may even
push the comparison so far as to find an analogy between
those irregular wanderers through cosmic space which
from time to time get drawn within the limits of some

* E.g., a8 was pointed out above, forms hke Pieris pandosia,
P, leptatina, etc,, show the result of attraction by the énachia group
upon Pieris of the ordinary kind. Similarly the yellow female
of P. demophile exemplifies the ordinary form attracted in another
direction, that of the agna combination, Again, the last-named
development of Pieris has served as a basis for a further attraction,
that by Heliconius charitonia, as seen in P. viardi 9 ; and this latter
form has given scope for the influence of the atthis group as shown
by P. locusta 9.

Mimetic Attraction. 331

established system, and certain species which seem to
hover on the outskirts of mimetic groups, undecided as it
were whether to throw in their lot with one association
or another.

I am indebted to Professor Poulton, F.R.S., for free
access to the Hope Department, and for permission to
figure insects from the collection under his charge.

Nore.

In the. course of the discussion which followed the reading of
this paper, my friend Professor Poulton expressed the opinion that
the term “mimicry” should be restricted entirely to cases of
Batesian association, and should not be applied to resemblances
between distasteful forms. I am, of course, entirely af one with
Professor Poulton as to the essential difference between the false
warning which is the leading feature in the one case, and the true
warning which characterises the other ; and I agree that it would
be most desirable to mark the distinction by the use of separate
terms. Though the present paper must stand as it was read, IT am
willing in future to attempt the restriction which he recommends,
Perhaps ‘“ Miillerian assimilation” and “ reciprocal assimilation,”
though a little cumbrous, may serve instead of the terms used in
the text ; and it may be hoped that the advantage of greater pre-
cision thus gained will outweigh the disadvantage of having to
drop such convenient words as ‘ mimic,” “mimetic” and ‘ model ”
when speaking of a Miillerian group.—June 2, 1897,

332 Dr. Frederick A. Dixey on Mimetic Attraction.

List oF Species MENTIONED.

PIERINZE. HELICONIINA.
Pieris phaloe, Godt. Heliconius numata, Cram.
P. calydonia, Boisd. H. atthis, Doubl.

P. demophile, Linn. H. charitonia, Godt.
P. viardi, Boisd. HI, cydno, Doubl. & Hew.
P. locusta, Feld. Hi. galanthus, Bates.

P. pandosia, Hew.
P. kigaha, Reak,
§ P. leptalina, Bates.

( IE pisonis, Hew. DANAINA.
Mylothris lypera, Koll.
M. lorena, Hew. Aeria agna, Godm. & Saly.
M. malenka, Hew. Tithorea pavonii, Butl.
M, pyrrha, Fabr. T. tarracina, Hew.
M. alethina, Butl. Napeogenes inachia, Hew.

EXPLANATION oF Prats VII.

. Pieris calydonia 9.

. P. leptalina, underside.
. P. demophile 2.

. Napeogenes inachia.

5 ER Goin

. Aeria ugna.

» Plocusiar®y.

. Heliconius atthis.

ADANanFrwwnwre

IN ALL THE FIGURES

k, diagonal dark bar of forewing.

1, dark bar of forewing parallel to inner margin.
m, pale area on apical side of diagonal bar.

n, submarginal pale spots.

o, transverse bar of hindwing.

N.B., k and / asin Trans. Ent. Soc. Lond, 1896,
Plates III.-Y.

Trans. Ent. Soc. Lond. 1897. PL. Vil

L
7 ig
is VETLS VLO ai )
ba
ane
17.7 )
D 7 + Heltsconwis atthars
CLEP US LOCALS TOL O -
Y g
8
FAD mey pinx' West,Newmsan chrorno

Mametic Attraction

[Reprinted from the Proceedings of the Entomological Society of
London, 1897, pp. xx.-xxxii., xxxiv.-xlvii.]

PAPER, EXHIBITION, AND DISCUSSION ON
MIMICRY. (May 5th, 1897.)

Dr. Dixy read a paper on ‘‘Mimetic Attraction.” He
began by pointing out with the aid of diagrams that the
process of mimetic assimilation might start from a given
point and proceed along several divergent paths. For
example, from an ordinary non-mimetic form of neotropical
Pieris, such as Pieris phaloe, not one only but several diver-
gent series of mimetic modifications could be traced; each
passing through a graduated series of closely-allied forms
until it terminated in a Pieris or Mylothris bearing a more or
less intimate relation with some protected form or forms of
entirely different affinities. Such were the series leading
through P. calydonia to P. demophile 2, a mimic of Aeria
agna ; the series starting from P. demophile and passing into
forms such as P. locusta 2, which was in mimetic relation-
ship with Heliconius atthis; the series represented by P. pan-
dosia and P, leptalina, which approached Napeogenes inachia ;
the series dealt with in a previous communication (Trans,
Ent. Soc. Lond., 1896, pp. 65-79), which led up to forms
associated with Heliconius numata; and lastly, a series
derived from the one last-mentioned, which ended in
Mylothris alethina 2, a mimic of Tithorea tarracina,

In all these instances, the model towards which the series
tended did not present an isolated scheme of colour, but was
a member of a larger or smaller group of forms, associated in
external features; such as were shown in the series to be
exhibited by Mr. Blandford. These mimetic groups were no
doubt mainly of the ‘‘ Millerian” kind, that is, they were
associations between inedible species of various affinities.
Reasons were given for attributing great importance to the
operation of the Miillerian principle in producing such a
facies as that of the neotropical fauna, and for assigning to
Batesian mimicry a comparatively subordinate position. Stress
was laid on the facts (1) that Batesian mimicry could exist
only when the numbers of the mimic were insignificant
compared with those of the model, whilst a Miillerian group
was strengthened by every fresh accession; and (2) that the

a

ao

attractive power in Batesian mimicry acted only from the
model towards the mimic, whereas in Miillerian association it
was mutual, and tended to produce, reciprocal changes. For
this mutual modification of characters the term ‘‘reciprocal
mimicry’ was suggested. A relation of the latter kind
appeared to exist, e.g., between Pieris locusta ¢ and Heliconius
cydno, and it must therefore be concluded that the Pierid was
inedible, a conclusion previously reached in the case of other
Pieridze on more than one kind of evidence. A consequence of
the keen competition for life in such a region as the neotropical
was that scarcely any conspicuous form was completely isolated.
If edible, it would generally be a Batesian mimic; if nauseous,
it would be drawn into the vortex of one of the great Miil-
lerian groups. The force exerted by these latter was well
exemplified by the facts dwelt upon at the beginning of the
paper, which showed the steps by which the members of a
single genus such as Pieris, and sometimes even the sexes of a
single species such as P. locusta were drawn apart into more
or less intimate relationships with separate Miillerian groups.
Mr. Buanprorp then exhibited and described the series of
Neotropical butterflies from the Godman-Salvin collection,
which were shown by him on a previous occasion (Proc. Ent.
Soc. Lond., 1896, p. xxxviii.); he stated that he used the
term ‘‘homceochromatism,” originally employed in the
‘* Biologia Centrali-Americana,”’ in preference to Dr. Dixey’s
term ‘‘reciprocal mimicry,” to denote the phenomenon
exhibited by ‘‘ Miillerian”’ groups, because it gave a sufficient
indication of its nature without assuming any theoretical con-
sideration as to its origin. The series shown were :—

J. Homa@ocHRoMATISM BETWEEN Parrep SPECIES OF THE SAME

GENUS.
Heliconius galanthus and H. luce. Guatemala.
H. chioneus and H. sappho. Panama.
H, cydno and H. eleuchia. Colombia.
H. alithea and H. primularis. EKeuador.

Each pair was closely alike on the upper-side; the under-
sides were different, there being common types for the right-
hand and left-hand members of the pairs respectively.
There was also a progressive modification in the patterns of
the upper-side in proceeding from North to South.

( iw)

Il. HomaocurRoMATISM BETWEEN ParrepD SPECIES OF DIFFERENT
GENERA.

Tithorea humboldti and Heliconius cassandra. Colombia.

Tithorea candollei and Heliconius sp, Antioquia.
(with these associated E’picalia chromis, @ ).
Tithorea bonplandi and Heliconius hecuba. | Colombia.
Tithorea pavonit and Heliconius atthis Ecuador.

(with these associated Pieris locusta, 2).

Il]. Groerapuicat Mopirications or HomamocHromatic SPECIES
OF THE SAME GENUS.
Heliconius thelviope and H, vesta,

a. Parallel varieties of both species in Cayenne, inoscu-
lating with H. melpomene at one end of the series,
and extending to typical H. thelwxiope and H. vesta at

- the other end; the varieties occurring without refe-
rence to geographical distribution.

b, Fixed geographical paired races of both species from
British Guiana, Lower Amazons (Para), Upper
Amazons (Pebas), Ecuador and Bolivia, and accom-
pained in the first three localities by homeochro-
matic races of H. ade,

c. Intermediate forms connecting H, vesta with H. phyllis
in Bolivia.

TV. Exrenstve Homa@ocnromatic (Minuertan) Groups.*

1. North Central-American Type.—Guatemala to Nicaragua.

Lycorea atergatis. Heliconius telchinia,
Melinea imitata. Eiueides zorcaon,
Tithorea sp. Eresia philyra.
Mechanitis doryssus. Protogonius cecrops.
Ceratinia dionea. Dismorphia praxinoe,
C. fenestella, Mylothris malenka.

* The genera in the left-hand columns of the list, except Lycorea,
a true Danaid, belong entirely to the Heliconioid Danainz (Neotro-
pine). Of those in the right-hand columns, Heliconius and Eueides
belong to the Heliconiine ; Hresia, Anwa and Protogonius to the
Nymphaline ; Archonias, Mylothrisand Dismorphia to the Pierine ;
and /thomeis to the Erycinide.

( iv.)

Intermediate between Groups 1 and 2.
Melinea scylax. Heliconius clarescens.
Mechanitis lycidice.

Napeogenes tolosa.
Ithomia heraldica.

2. South Central-American Type.—Costa Rica to Panama.

Thyridia melantho, Heliconius zuleika.
Tithorea helicaon, HZ, formosus.

T. duenna. Hueides vulgiformis.
T. pinthias. Eresia nigripennis.
Mechanitis isthmia. E. pecillina,
Ceratinia decumana. Anaa jansoni, &.

C. callispila, Ithomeis imitatria.
Napeogenes tolosa. Archonias lyceas, ¢ .
Dircenna relata. Dismorphia deione, ?.
Callithomia hezia. D. sororna, ?.
Hyposcada adelphina. Mylothris malenka, ?.
Pteronymia notilla. Papilio zalates.

Hypoleria libera.

Colombian modification of 2.

Tithorea hecalesina, Heliconius hecalesia.
Napeogenes peridia. Eresia ithomioides.
Pteronymia picta. Anea panariste, 2.

3. East Brazilian Type.

(a.) The apical spots on the forewing yellow.

Lycorea halia. Heliconius dryalus.
Melinea ethra. Hueides dianasa.
Tithorea sp. Hresia esora.
Mechanitis nesaa, Eresia sp.

Napeogenes xanthone. Dismorphia astyonome.

Ceratinia laphria.

(6.) The apical spots on the forewing white.
Mechanitis lysimnia. Heliconius narcea.
Napeogenes euryanassa.

Ceratinia dacta.

4. Guiana Type.

Lycorea ceres,
Melinea mneme.
Tithorea harmonia.
Mechanitis polymnia.
Ceratinia philidas.

5. Upper Amazons.
a. Kga Type.

Lycorea cinnamomeana,

Melinea pardalis.
Tithorea harmonia.
Mechanitis egaensis.
Ceratinia fluonia.

b. Sao Paulo Type.

Lycorea cinnamomeana.

Melinea cydon.
Mechanitis olivencius.
Ceratinia tigrina.

Heliconius numata.
Eresia sp.
Dismorphia amphione,

Heliconius pardalinus.
Protogonius castaneus.
Dismorphia egeana,

Heliconius isabellinus.

c. Lower Rio Napo Type.

Melinea sp.
Mechanitis sp.
Callithomia sp.
Ceratinia sp.

6. Ecuador Type.

Melinaa cydippe.
Mechanitis mothone.
Ceratinia semifulva.
Napeogenes sp.

FTeliconius sp.
Eueides sp.
Eresia sp.

Heliconius aristiona.
Eueides acacetes.
Acrea acipha.

Evesia tthomiola.
Protogonius semifulvus.
Papilio bacchus.

( wy
7. Central Colombian modification of 6.

Melinea messenina. Heliconius messene.
Mechanitis menophilus.
Napeogenes sp.

The greater part of these groups, which also contained a
few moths belonging chiefly to the genus Pericopis, consisted
of species of the Heliconioid Danaine or, as they had been
recently called, Neotropine ; one or more species of Heli-
coniide were constantly present, and the associated forms of
Heliconius might conveniently be regarded as representatives
of the respective colour-types. A few Nymphaline, Pierine,
Papilionins, and in one case an Erycinid, had been drawn
into the vortex. The colour-types which they represented
were so far diagnostic of the geographical distribution that
a strange specimen belonging to one or other of these groups,
which were a few only out of many such that could be put
together, could be unhesitatingly assigned by its facies to its
original habitat in Tropical America.* But whereas certain
groups, such as 1, 2, 8, and 4, were of extensive geographical
range, others, such as the Ega group, 5a, with its peculiar
suffused brown coloration, or the Rio Napo group, 5c, were
restricted to a comparatively small area. This was intelligible
in the Andean region, where every valley had its own special set
of forms and where isolation played the same part as in the
evolution of insular faunas, but, as Bates had pointed out,
it was a very remarkable phenomenon in the Amazon valley,
throughout which the physical conditions were very uniform.
He did not propose to dispute either of the theories of mimi-
cry which were associated with the names of Bates and
Miller, but they rested very largely on hypothesis and were

* As an example, group 4 is distinguished mainly by having
the hindwings almost or quite black behind the transverse black
bar usually present. The same character exists in British Guiana
examples of H. thelxtope and H. vesta, not in other respects true
components of the group; it is absent in their Amazon repre-
sentatives, at least as far as Bolivia, where it turns up again in
association with very different groups.

(. vii.)

in want of further support from observation and experi-
ment, which would afford a large if arduous field of
work to the enterprising naturalist; the difficulties of the
subject did not appear to him to be fully overcome by these
theories, which should not be pushed so far as to lead to the
disregard of other factors which might have influenced the
genesis of these groups. One difficulty was that of distribu-
tion. As before mentioned, the groups of the Upper Amazon
valley were often of limited range; but if they were geneti-
cally connected and the conditions of their environment were
constant, the causes which brought about association under
a common type, if prohibiting deviation therefrom under
penalty of destruction, should have operated to extend the
limits of a group as widely as possible by acting as a check
to variation on its outskirts. If it were assumed that one
form were so far dominant as to drag its associates with it in
any given direction, it must be also recollected that the prin-
ciple on which these large ‘‘ Millerian’’ associations were
supposed to be based was prohibitive to variation of any
component species, either as a whole, or in any part of the
range of the system. ‘The logical tendency of such a group
would be to extend its limits indefinitely and not to give rise
to repeated changes of the colour-type.

Another difficulty was presented by the very close resem-
blance, at times amounting to identity in external characters,
between certain pairs in these groups, a resemblance to
which Brunner’s epithet, ‘‘ hypertelic’? might be applied.
Existing theories postulated a selective elimination by insecti-
vorous birds, ete. ; but the birds’ discrimination of members of
a ‘* Millerian,” z.c., protected group must be in relation to
its distinctness from the other insects co-existing in the same
region. If, as was frequently the case, such a group was
immediately recognisable by its broad features as protected
and inedible, such further discrimination between its members
as would be necessary to bring about the intimate likeness
found, e.g., between many species of Melinea and Heliconius
was not adequately accounted for by Miuller’s hypothesis.

With regard to ‘‘ convergence,” which had been put forward
as a necessary phenomenon in Millerian mimicry, the

CY yi)

possibility of parallel variation ought not to be excluded. To
take the case of H. thelxiope and H. vesta, the ‘ typical”
forms might be assumed, ex hypothesi, to be the most ancestral,
and to have acquired their common resemblance by conver-
gence; but from these forms were derived, as the series
exhibited showed, a number of paired varieties which were
progressively modified in relation to their distribution. There
was nothing to show that either species had been influenced
in its variation by the other, and that it would not have
followed the same course if it had been isolated. And it was
conceivable that the causes, in most cases unknown, which
brought about modifications in the colour and markings of a
species in association with its geographical range, might have
produced identical results in two species of the same genus,
with a common facies, under common conditions.

Prof. Pouttron : He congratulated Dr. Dixey on his care-
ful work, and on the deep interest of the results he had
obtained. It was a great source of satisfaction to him that
this research had been conducted in the Hope Department of
the Oxford University Museum.

He then exhibited a further series of neotropical butterflies
from the Godman-Salvin collection, illustrating the various
members of the group which had been formed round Methona
confusa, Butl., and Thyridia psidii, L.; the best known moths
(of the genera Castnia and Hyelosia) which fell into the group,
were supplied by specimens from the Hope collection. This
group which, as regards most of its members, had been
originally described by H. W. Bates, occupied a very wide
geographical area, and was of special interest, not only on
account of the number of forms which entered into it, but
also because of the perfection of the resemblance.

Further, he exhibited the smaller group which converges
around [tuna lamira, Latr.,and species of Olyras, Thyridia, ete.,
the specimens being selected from the Godman-Salvin collec-
tion; he also showed several groups characteristic of Honduras,
Surinam, Eastern Brazil, etc., the specimens having been
recently acquired by the Hope collection for the purpose of
illustrating the principles of Warning Colours and Mimicry.
Many of these specimens possessed the special interest that

Ci)

they were captured by one collector nearly at the same time,
and in one locality. In fact, in the case of the Honduras
insects (presented by Col. Swinhoe), examples of several
different species and genera had been sent in one set of papers
as a single species. Thus in the case of these groups evi-
dence was actually forthcoming that the separate species do
live together intermingled, and are liable to be confused, at
any rate by a human collector.

The term “ homcochromatism”’ was criticized on the
ground that it was a mistake in science rigidly to exclude
theory and interpretation. A theory might be a good guide
to discovery, even if it turned out in the end to be imperfect
or wrong. And in this case it was contended that the theories
of Mimicry and of Warning Colours had by no means been
proved to be wrong, but remained as the only hopeful inter-
pretation of the facts.

He also objected to Dr. Dixey’s phrase, ‘reciprocal
mimicry,’ inasmuch as the resemblances alluded to were
those of specially defended insects, and not of forms which,
being themselves harmless, lived on the reputation of their
better defended neighbours, as in mimicry proper. Protective
Mimicry had been defined by the speaker as ‘‘ False Warning
or Deceptive Warning Colours” (pseudaposematic), while,
according to Dr. Dixey’s contention, there was nothing false
about the warning colours of the insects described.*

It was contended that the peculiar local groups, such as
those of Ega and the Rio Napo do not offer any difficulty
to the theory. The ordinary laws of variation continued to
operate after the formation of a group, only in this case the

* Tt has since occurred to me that terms accurately descriptive
for those who accept the theories of Mimicry and Warning Colours
may be obtained by an extension of the terminology proposed by
me in 1890 (“The Colours of Animals,” pp. 336 e¢ seqq.). In the
majority of cases there is reason for the belief that Miillerian
groups have been formed by a gradual approximation towards the
appearance of some aggressive and abundant species, or towards
that of the most prominent general characteristics of several such
species belonging to a specially defended section of Lepidoptera,
such as the Euploeine or Acrzinz. Miilleriau groups of this kind—
probably by far the commonest—may besaid to possess Synaposematic

es)

change which so often occurs in a species with a wide geogra-
phical range, ultimately splittmg it up into subspecies and
true species, would draw the associated forms after it by the
operation of the very principles by which the group was
originally formed. If this process were sufficiently gradual
no principle of Miullerian association need be violated. Or
the facts might be reasonably explained in another way: the
dominant (a word which here implies only the commonest
and best known, in fact, the most widely advertised
of the specially defended Lepidoptera of the district)
form might have inhabited the region in question, and
assumed its peculiar aspect before the formation of the
group, and may have then separately ‘‘ converted ’’ each new-
comer as it arrived in the district. The tendency of a group
was certainly, as Mr. Blandford maintained, to extend its
limits indefinitely—a tendency which had operated with great
success in certain cases. But the spread of species always
encountered opposing forces which in many cases acted as
effective barriers.

With regard to Brunner’s ‘ Hypertely,’’ he maintained
that one knew far too litle of the details of the struggle for
existence to justify the conclusion that it was incompetent to
produce such effects. What little was known confirmed the
belief that very minute differences might serve to turn the
scale. The differences between extremely perfect resem-
blances and those which were less perfect or only very rough,
were probably to be explained by the relative age of the
association in the former, or the more complete and rapid
operation of natural selection on account of a special reliance
on this among other modes of defence possessed by the

(abv, together ; ard, away : ofa, sign) colours, pattern, or appear-
ance, the noun being Synaposeme. They may also be said to possess
Millerian Warning Colours or Common Warning Colours. For
those extremely interesting but, as I believe, relatively uncommon
cases, in which the approach is mutual—a process of ‘give and
take,” so well described by Dr. Dixey—the term Diaposematic
(Sd, used to express mutual relation, as in “ dialogue,” ard, and ojua)
may be employed, the noun being Diaposeme. These cases may
also be spoken of as Reciprocal Warning Colours. Mr. Arthur
Sidgwick has kindly helped in the formation of these new terms.
—E. B. Poulton, June 14th, 1897.

( xi)

species. One was compelled to believe that every perfect
resemblance began as an imperfect resemblance, and then
passed through stages in which the likeness was gradually
increased ; and it was only to be expected that examples of
all such stages should exist at the present day among the
numberless forms which exhibited mimicry and common
warning colours.

Gradual changes in the geographical distribution of the
constituent species along the borders of groups would tend
from time to time to bring certain of them within the
influence of other groups, and so begin a change in another
direction. Furthermore, there was no reason for concluding
that the detached members of Miillerian groups must become
extinct, as in the case of the Batesian or true mimic. In the
presence of other dominant members of a group, any ten-
dency towards resemblance might well be of selection value :
in their absence it was by no means necessary to assume that
a species, which, ex hypothesi, was specially protected in some
way, must become extinct, although any further advance
towards the likeness would be checked, and the ground gained
in the past lost after a longer or shorter interval. To enter
the area of another group would tend towards a rapid modifi-
cation of the old appearance.

The suggestion that parallel variation assisted in the for-
mation of these resemblances was strongly opposed by the
fact that the superficial characters were alone affected, and
that the closeness of resemblance bore no relation to degree
of affinity. For. instance, the resemblance between a
Melinea and a Heliconius was frequently much closer than
that between the former and a Mechanitis, or the latter and
an Eueides, in the same groups. Although the interesting
facts brought forward by Mr. Blandford with regard to Heli-
conius thelxiope and H. vesta, etc., would, taken alone, seem
to support his suggestion of parallel variation, yet when they
were considered as part of the whole phenomena of Millerian
warning colours, as exhibited and as at present known and
understood, in the Neotropical Rhopalocera, one was led to
believe that one set of principles had been at work, and that
natural selection, which, he contended, offered the only hopeful

(xr)
solution in the vast majority of cases was the true explanation
of the others also.

Further strong support for this conclusion and further
difficulty in the way of any other interpretation as yet offered
was to be found in the similar behaviour of the groups which
in other tropical countries represented the Danaine, Nectro-
pine, and Heliconiine of 8. America. Thus unmistakable
indications of Miillerian association were to be found among
the Acreine of Africa and among the Danaine and Kupleine
of the Oriental Region. It was impossible to contend that
these representative groups possessed the monopoly of parallel
variation, or of change under direct influence of the environ-
ment.

The Presipent: In treating of the Miillerian associations of
species closely resembling each other—many of which were so
well illustrated by the admirably arranged series of Tropical-
American Lepidoptera exhibited by Mr. Blandford—there
was always, in his opinion, great risk, in the case of species
of the same genus or even of nearly allied genera, of mis-
taking for mimicry the similarity really due to close affinity in
blood. It also occurred to him that in these Miillerian com-
panies, where each component species is admittedly protected,
some certainly and the rest presumably, by offensive and dis-
tasteful secretions, there did not apparently exist the same
necessity for exact imitation as was demanded in the case of
the Batesian mimicries, when the very existence of the unpro-
tected edible individual and species depended on the closest
simulation of the protected inedible form. The opinion had
been expressed that in the latter cases mimicry had some-
times been carried to an exactness in minutie quite unneces-
sary; but he thought that no one, who considered the life
conditions under which these mimicries had been brought
about—the intensity of competition, the overwhelming fer-
tility, the complex inter-relations of organisms, so charac-
teristic of tropical regions, could seriously conclude that these
special modes of protection could by any possibility be too
perfect. We might rest assured that imitations so complete
as many of them are would certainly not exist if they had not
become necessary.

(7 xin)

In the discussion of these extremely interesting phenomena
it was only too evident how great was the need for prolonged
and continuous systematic study of the living forms in their
natural environment. The collector in tropical regions,
however able and enlightened, and however desirous of con-
tributing to the elucidation of these and kindred problems,
was too much hampered by the hindrances and interruptions
incident to his main occupation to admit of his undertaking
such a series of observations as was essential. What was
wanted were Biological Stations dealing with the terrestrial
fauna on the lines of late so successfully followed in marine
biological research.

The discussion was then adjourned till June 2.

Discussion on Mimicry, ete.

The discussion on Mimicry and Homeochromatism in
Butterflies, adjourned from May 5, was then resumed.

Dr. Dixry: He wished to add his voice to the chorus of
approval and thanks which had greeted the very fine exhibit
arranged by Mr. Blandford from the collection of Messrs.
Godman and Salvin. He thought that his own views were
supported by the contents of the drawers shown, and that the
series of examples contained therein were, in most instances,
Miillerian associations of an extensive character.

He agreed that the term ‘‘ homceochromatism”’ had an
advantage in merely denoting the facts, without reference to
any theory ; but he thought that Mr. Blandford ought not to
restrict it, if such were his intention, to Millerian mimicry.
It was equally true that Batesian mimics were homcochro-
matic with the models they represented.

He was not sure whether Mr. Blandford intended his
remarks about the intimate resemblance existing between
certain species included in these groups to be considered as a
criticism of the Millerian theory. The course taken by any
species was a resultant of the various forces acting upon it,
and the perfection of a mimetic resemblance would therefore
stand in relation to the facility for its acquisition which was
allowed by the other forces tending to modify the external
appearance.

( xive 1}

Professor Poulton had criticised the use of the term
“mimicry” in the case of Miillerian resemblances. The
speaker agreed that any ambiguity between these phenomena
and Batesian mimicry was to be avoided, but it had not
occurred to him that his use of the term ‘reciprocal
mimicry’’ was open to this objection. He had also been
charged with saying that Miillerian mimicry was necessarily
reciprocal. Possibly he had conveyed this impression ; he did
not, however, intend to assert that this form of mimicry must
always so demonstrably act as to produce reciprocal alterations
among species associated under its influence; but merely that
it exercised a constant potential force, in some cases becoming
actual and capable of demonstration, towards mutual converg-
ence, whereas in Batesian mimicry the mimic was neces-
sarily without any such influence on the model.

He was glad to hear Professor Poulton say that he was
becoming convinced by the speaker’s arguments that the
Pierine were a protected group. He should not himself,
however, care to assert more than that much evidence point-
ing to this conclusion existed in respect to many members
of the subfamily—for instance, the genera Delias and Mylo-
thris; while in some cases the evidence for inedibility was
considerably strengthened by the presence of reciprocal
change.

Canon Fowrrer: Could anyone explain the existence of the
‘*predominant partner’ which was assumed in the explana-
tion given of these groups? He could not imagine why one
species should be stable and the others unstable and depen-
dent upon the former for their characters.

Mr. Exwes: The society was much indebted to Messrs.
Godman and Salvin for the loan and to Mr. Blandford for
the arrangement and exhibition of these valuable specimens.
He doubted if there were any other collection in the world
from which such an exhibit as that shown could be got together.
To his mind the specimens were of infinitely greater scientific
value in their present arrangement than they would be if
dispersed throughout a collection in their proper systematic
positions.

He therefore pleaded that when the question of their being

( xv )

sent to their ultimate destination, the British Museum,
arose, the possibility of their being kept permanently to-
gether might be considered, and their retention in the present
arrangement stipulated for, The absence of any rare or
unique examples from their proper places in the collection
would be more than compensated for by the increased interest
which would thus be secured, and the series would serve as
a model to curators of other museums in the art of making
butterfly collections interesting and instructive.

He had to speak that evening without the advantage of
having heard the first part of the discussion, and what he
had to say was therefore based mainly on the abstract which
had been furnished of it. He thought, and his opinion was
formed on personal experience gained in collecting in many
countries, tropical and otherwise, that there was too much
assumption about either the Batesian or Miillerian theories
of mimicry. In many supposed cases he doubted whether
the so-called models were protected either by taste or smell,
and he thought that the importance of birds as enemies of
butterflies had been overrated. It seemed to him that the
protection supposed to be given to the imago was of little
account in the perpetuation of the species when compared
with the destruction which took place in the larval or pupal
stage by climatic influence, as well as by insect and other
enemies.

However beautiful these explanations of the phenomena of
mimicry were in theory they ought not in most cases to be
treated as proved scientific axioms. He did not deny that
there might be some truth in each of the theories which had
been put forward, but it appeared to him that altogether too
much stress was laid on them. Had any observations been
made which would justify the statement that the members of
a Miillerian group were inedible, as was asserted by Dr. Dixey
in his quoted remarks ?

He desired to call attention again to a passage in his
Presidential Address on Geographical Distribution (Trans.
Ent. Soc. Lond., 1894, pp. lxv., Ixvi.). He had said:
‘* What is most remarkable is the existence, at high eleva-
tions in various parts of the Andes, and at sea level in South

(sya)

Chili and Patagonia, of several genera and species elsewhere
unknown in the Neotropical region, and which are isolated
from their congeners in North America by an enormous area
of country.

‘‘ Among these Trifurcula huanaco is a remarkable species
which occurs in the Andes of Bolivia, at 16,000 to 17,000
feet, and has a marvellous likeness to Baltia shawi, found at
a similar elevation in Ladak.

“‘ Phulia, a genus of three or four nearly allied species
also occurring at great elevations in the Andes and Chili, has
a striking resemblance to Synchloe butleri, a species which
accompanies Baltia in Ladak. If similar conditions of en-
vironment do not produce similar effects, how can these
extraordinary cases of resemblance in remote and discon-
nected areas be accounted for ?”’

He hoped that entomologists who resided in places where
any of the supposed instances of protective mimicry occurred
would pay special attention to the life-history of the species
affected by it, as such observation alone could prove or dis-
prove the question.

Mr. Verratt : Homcochromatism was not a phenomenon
confined to the tropics. Homcochromatic resemblances
existed even in Europe between Diptera and Hymenoptera,
and it must be assumed that they stood in relation to the
protective armature of the Hymenoptera.

Col. Swrnnor: A challenge had been held out as to whether
the distasteful qualities of protected Lepidoptera had been
experimented on. As an example of a species, of which such
qualities were placed beyond doubt, he instanced Danais
chrysippus, one of the most widely-spread protected models.
This insect was so free from attack, owing to its nauseous
character, that the protection extended even to dried
examples. These would be found untouched in a box of
insects, although the remainder of its contents had been
destroyed by mites or Anthrent. He did not doubt that
there were many other protected Lepidoptera, including many
Pierine genera—for example, all the species of Teracolus
appeared to be inedible.

Mr. Jacosy: If protected and inedible species were so

(y xvile *)

widely distributed, why were they not universally imitated
by unprotected insects? He did not think that sufficient
proof had been given of the existence of protection.

Sir Grorce Hampson, Bart.: In his personal experience
in 8. India he had found that it was quite an exceptional
thing to see birds catch, or even attempt to catch, butterflies,
whilst it was a matter of daily occurrence to see them taking
other insects, especially moths, ete., started from a position
of rest. In the cases he had witnessed the Huplee and
Danaidae were caught as often as any others, but usually
escaped eventually from the beak of the bird and flew away
none the worse owing to the toughness of the integuments.

The only bird he had observed frequently to pursue butter-
flles was the common Indian Bee-eater, which he had seen
hawking Pieride, and among them Teracolus, which Col.
Swinhoe had expressly referred to as a protected genus.

He thought the cause demanded by these theories was
inadequate to produce the results assigned to it.

The Hon. Watrer Roruscuitp: It had struck him that it
was much more conceivable that certain climatic influences,
etc., had played a part in bringing about these resemblances,
and he thought that these groups assumed the same appear-
ance because one given influence was at work on them.
Such a case as that of Papilio merope, and its various
representative forms inhabiting Africa (except the Palearctic
North part) and Madagascar were inexplicable by mimicry
alone. This Papilia was monomorphic in Madagascar, the
female being similar to the male both in colour and form of
wings, while the representative found in Abyssinia had the
hindwing tailed like the male, but possessed a decidedly
‘‘mimetic’’ pattern. In this respect it resembled the female
forms of S. and W. Africa, which were, however, tailless.
He contended that these geographical forms of Papilio merope
had probably sprung up under the direct influence of the
extarnal conditions of the respective areas rather than in
consequence of the direct selection of specimens with a more
‘‘mimetic” pattern.

Canon Fowirer: There was too much assumption about
the current theories.

( xviii)

Mr. McLacutan: A point that appeared to him to be a
good deal overlooked in this matter was the possibility that
two species might go on independently and yet apparently
mimic each other by arriving at the same results in their
modifications.

Prof. Weupon, F.R.S.: He did not intend to enter upon
the details of mimetic resemblances or upon the theories
which had been put forward, but, speaking as a visitor, he
would like to take the opportunity, as he did whenever an
occasion presented itself, of pointing out to entomologists
that the truth or falsity of these theories was capable of being
tested to a large extent experimentally, and especially by
experiments which would give a basis for statistical deter-
minations.

Thus, if one took a mimetic species distributed over a
large area, homceochromatie with one species in one district,
and with another in another district, such a species on
theoretical grounds must be variable and susceptible to a
moderately rapid selective action.

In any given area, although the examples caught might
conform fairly closely with a common type, the species must
still be variable and capable of selection ; if the theories were
true there should be evidence of continued selective destruc-
tion, and this could be ascertained by experimental breeding
on a large scale from eggs of wild individuals. A comparison
of the examples thus reared with an equal number of wild
specimens from the same locality would show whether there
was any greater variability among the bred forms. Assuming
that sources of error had been excluded, such a result would
be good evidence of the existence of continued selection
tending to ensure conformity with the model.

Mr. J. J. Watxer: He was unable to believe that birds
were effective agents in causing mimetic resemblances.
During all his experience as a collector in different parts of
the world he had never seen a bird pursue and catch a
butterfly but once. On the other hand, butterflies were often
eaten when at rest by lizards, small mammals and monkeys.

Col. Yersury: During his personal experience of many
years in India and Ceylon he had hardly ever seen a bird

GC sis })

touch a butterfly. Some years ago the question was raised
by the Bombay Natural History Society, and he, with others,
took notes on the subject. He recorded two cases only
during three years’ observing. It was significant that while
the flocks of locusts and white ants were attended by
vertebrates of all orders, the tlocks of butterflies in Ceylon
(locally known as ‘‘snowstorms’’) were attended by one
species only of bird, and that but seldom.

In his opinion the enemies of butterflies were chiefly, if not
entirely invertebrate. In Ceylon two protected species,
Euplea core and Delias eucharis, were largely taken by a
mantis, Gongylus gongyloides, while two of the large Asilide,
Promachus maculatus and Scleropogon ambryon preyed largely
on Danais limnace.

Mr. Buanprorp: In criticizing the term ‘‘ homceochroma-
tism.’’ Prof. Poulton had, he thought, somewhat mistaken
the speaker's attitude. 2 had no intention whatever of
excluding theoretical considerations, even if he could not
accept them at their full value. But it was obviously unjust
that a class of facts, about which there could be no dispute,
should be labelled with a collective name implying the
acceptance of a theory which, however well it might stand
criticism, had certainly not yet been established. He
preferred to keep one terminology for the facts and another
for the explanatory theory.

The wideness of meaning which he proposed to attach to
the term ‘‘homeochromatism’’ required some explanation.
Certainly he conceded that it covered cases of Batesian
mimicry; but if generally adopted, it would probably prove
convenient to give it a more restricted and conventional
meaning by their exclusion: such a conventional limit had
constantly to be applied in terminology. In order to keep
the nomenclature of these facts independent of speculation it
seemed desirable to employ the words ‘‘ mimic” and ‘“ model”
without reference to the questions of Batesian or Miillerian
mimicry, the essential character ot a ‘“‘ mimic ’”’ being that of
a wide departure from the general type of its genus or
subfamily with a resulting likeness to a model which was not,
or scarcely, modified thereby; in homcochromatism, as he

( xx )

desired to limit the term, and as he had illustrated it, no
such one-sided departure was manifest.*

The President had called attention to the risk, in the case
of species of the same genus, or of nearly allied genera, of
mistaking for homeochromatism a similarity due to blood-
relationship. Of course, the value of resemblances had to be
estimated in relation both to the range of form existing in
the genera or sections of a genus involved, and to distribution ;
and the speaker did not admit that such a mistake had been
made in respect to any single species of his exhibit. As far
as his knowledge went, instances of non-mimetic hom«o-
chromatism among Neotropical butterflies were entirely
confined to the Neotropine, Heliconiine, and Hesperiide,
although other subfamilies afforded mimetic forms.

He had brought forward certain difficulties attending the
current theories; Prof. Poulton’s epicriticism thereon was
based largely on the assumption that each group possessed
some dominant form. The speaker had already pointed out
that the Millerian theory was opposed to the existence of
dominant forms, and he knew of no direct evidence that such
actually were present—that was, species which could so far
influence a group as to compel its components to change
when they changed. His objections as to geographical
distribution had been met by two counter-suggestions, one of
which presupposed that a series of dominant forms had
preoccupied the country and had influenced the appearance
of the protected species which subsequently invaded it; but
one could not bring oneself to believe without strong evidence
that the groups of insects concerned, the Heliconiine
and Neotropme, were not coeval in distribution. Prof.
Poulton’s comments on ‘“ hypertely ’’ again presupposed the

* Homeeochromatic pairs, such as those of Series I. in my
exhibit, are not demonstrably in the relation of ‘ model” and
“mimic,” but in one for which some other word must be found.
I would suggest that each member of such a pair, or group, which
does not show the departure indicative of a mimetic form, be
called the “ homotype” of its associates. Thus Heliconius galanthus
would be the homotype of, or homotypic with, A. luce—W. F.
H. Blandford, July, 1897.

Csr J

existence of a dominant form in each group, the other com-
ponents of which were capable of being arranged in a
descending order of resemblance to it. That was scarcely
borne out by the specimens. A group such as that shown
from Panama presented several hypertelic pairs and did not
support the idea that the species could be arranged in degrees
of adaptation towards one particular model. Whilst the
facts of geographical distribution afforded, in his view,
areal objection to the Millerian theory, he did not look to
distribution alone as likely to have hada considerable share in
the production of these groups.* It might have had some ;
but their production might conceivably be due to a variety of
causes, and not one alone.

He had certainly intended his remarks on hypertely to be
taken as a criticism of the Miullerian hypothesis.

However effective a destructive agency might be in pro-
ducing change, directly it became non-selective, the resulting
change must stop short at the point reached; and _ his
argument was that the process of discrimination by birds,
the only available agents, would be limited to the recognition
of a group of associated forms as an inedible whole, and being
superfluous if carried farther would not be exercised so as to
bring about such minute resemblances as were often met
with. As an instance, he might mention the two Brazilian
groups, differing in the white or yellow colour of the apical
spots of the forewing. ‘To use a rough illustration, a person
whose object was to avoid the society of a policeman would
betake himself off at the first sight of the familiar uniform
and would not stop to decipher such minutiz as the distin-
guishing number thereon. His view was strengthened by
the President’s admission that in Miillerian groups there
apparently did not exist the same necessity for exact
imitation as was demanded in the case of Batesian mimics ;
it practically conceded his point.

Every one who had listened to the discussion must, he
thought, be struck with the amount of doubt thrown on the

* The objections to any explanation based on distribution alone
have been forcibly stated by Fritz Miiller himself. “ Kosmos,”
1882, p. 262.

@ xa}

bird-theory by speakers whose competence and opportunities
for observation were quite beyond dispute. Further impor-
tant evidence on this point had been lately given by M. Piepers
(Congr. Internat. Zool., III., p. 460) who had studied the
question in Sumatra and Java for twenty-eight years. In
that time M. Piepers had seen four cases only of butterflies,
two belonging to the “ protected’? genus Huplwa, being
attacked by birds; and his paper referred to the fact that
neither Pryer, after twenty years’ observation in Borneo,
Skertchley nor Scudder had geen or accepted such a
phenomenon. M. Piepers had arrived at the sufficiently
striking result that mimicry had nothing to do with Natural
selection.

The premisses necessary to support either theory of
Mimicry had been unduly neglected. For example, though
evidence existed to show that the models were protected and
inedible, the proof of the edibility of mimetic butterflies had
not received enough attention; it was necessary to Bates’s
theory, and all the more so since Dr. Dixey’s work on the
“‘ Millerian ” character of certain Pierine. These theories,
indeed, were really working hypotheses, the object of which
was to suggest experimental work tending to prove or
disprove them ; they were not yet to be put forward dogmati-
cally as true and as convincing proofs of Natural selection.
To insist that these homceochromatic groups owed their
origin merely to the educational requirements of birds had
led, in his view, not so much to a development as to a
stifling of broad speculation on and inquiry into the problem.

The facts presented by the Ithomie (s. lat.) had scarcely
been touched upon in the discussion. Almost every colour-
type among these insects, however insignificant in appearance,
was represented by species of two or more of the genera into
which the old genus /thomia had been divided; so that the
Ithomie might be said habitually to exist as homceochromatic
pairs. The coloration of many of these pairs, consisting of
nothing but afew black patches and a white or yellow patch on
a transparent ground was far from exhibiting the striking
features which one was led to believe were characteristic of
warning colours.

(- Sxxi >)

Personally, he was not disposed to reject Bates’s theory
which, even if of less general application than was commonly
supposed, was strongly supported by evidence from other
orders of insects, but he was unable to accept that which had
been called by the name of Fritz Miller. What was to take
the place of the latter? At present, nothing. Until more
information had been collected on the habits and variation of
homeeochromatic species, and on selective agencies, there
were no data on which a new theory could be legitimately
founded. But he desired to call attention to a significant
passage in Bates’s original paper on Mimicry (Trans. Linn.
Soc., 1861, p. 501). Bates said therein: ‘“‘ The process of
the creation of a new species I believe to be accelerated in
the Ithomie and allied genera by the strong tendency of these
insects, when pairing, to select none but their exact counter-
parts ; this also enables a number of very closely allied ones
to exist together, or the representative forms to live side by
side on the confines of their areas, without amalgamating.”
Such a statement indicated the possibility that sexual
selection, or the segregation of forms might take place as a
direct act of p2rception on the part of the insects themselves.
If such a phenomenon were shown really to exist, it would
remove many of the difficulties which the present theories
entailed, and in view of Bates’s definite and repeated state-
ments, some proof or disproof of them should be attempted
before the Miillerian theory came to be regarded as more than
a merely provisional suggestion.

The Prestipent: The point had been raised by Mr. Elwes
that the destruction of species in the imago stage was of little
importance when compared with the much greater destruction
of larve and pupx which took place. As a matter of fact,
he could bear witness that certain species at any rate of
- Danais and Acrea were distasteful and protected in all stages.
The larve and pupe of Danais chrysippus and of five or six
species of South-African Acrea were rejected by cage-birds
and common fowls; they were highly conspicuous (especially
the pup of Acrea, which are white or yellowish-white with
orange and black bars and spots, and suspended indiscrimin-
ately on green leaves, dark-brown bark of trees, tarred

(" Sai 9)

palings, etc.) and all gave out an odour weaker than, but of
the same character as, that emitted by the perfect insects.

With respect to the term ‘‘ homceochromatism,” it had this
disadvantage, that it was at once too wide and too restricted —
too wide, because it did not exclude cases of resemblance due
to mere relationship, and too restricted, because it left out of
sight the similarity in shape of wings or body, or of move-
ment and habit which often made up part of a mimetic
likeness.

He did not agree with Mr. Rothschild as to the case of
Papilio merope, considering that cases in which the 9 alone
was exactly modified in imitation of a protected form (or, as
in the instance under notice, of three or four differing
protected forms) were, by reason of the extraordinary contrast
with the unmodified ¢, more striking and unmistakable
instances of obvious and indisputable mimicry than even
those in which both sexes were similarly modified. As
regards the Abyssinian representative of P. merope named
P. antinorii, which until recently was thought like the
Madagascar and Comoro representatives (P. meriones and P.
humbloti) to have the sexes alike, without any mimetic
modification of the 9, Prof. Kheil had described and figured
in 1890 two forms of the 9? (collected, with seven ¢ ¢ and
two ? ? coloured like the ¢, at Lake Tana by thelate Dr. A.
Stecker) respectively closely imitative of D. chrysippus and
Amauris niavius, but still retaining the conspicuous tails
on the hindwings which all the other known forms of 9?
of the allied species on the African continent have lost.
There, it seemed to him, was a most interesting and con-
clusive case of mimetic modification still actually im progress,
the @ usually resembling the ¢ in both colouring and
pattern as well as in outline of wings, but also presenting
two other forms, each profoundly modified in simulation of:
a protected Danaine butterfly, yet an imcomplete mimicker
in so far that the tailed outline of the hindwings remained
unaltered.

He must admit that the capture by birds of butterflies was
rare, but he had himself seen birds, especially the Drongo
shrike, chasing butterflies.

Caan)

Mr. Mansel Weale had recorded his having witnessed the
capture of the ¢ Papilio cenea (the S. African representative
of P. merope) by the Fly-catcher, Tehitrea cristata; and Mr.
T. Ayres has observed that the small King-hunter, Zspidina
natalensis, fed almost entirely on butterflies.

The larger Madagascar Chameleons also ate butterflies, but
appeared to show no discrimination. Lizards attacked them,
but must necessarily see the undersides only, which were
protectively coloured in many cases.

As regarded the known protected butterflies, it should be
borne in mind that there was very little difference in colour-
ing and pattern between the two surfaces of the wings in the
Danaine and Acreine, so that warning indications of dis-
tastefulness were shown almost as conspicuously when at rest
as When in flight. The importance of this as a means of
protection was manifest, and was further evidenced by the fact
that it was exhibited very markedly by the mimicking forms
as well.

The chief invertebrate enemies he had noticed were
Asilide and dragon-flies.

It seemed to him impossible to explain except by the theory
of mimicry such cases as that of Danais chrysippus, a widely
distributed and very common insect which was attended by a
troop of mimetic species wherever it went.

Dr. Drxry: It was too late to deal fully with all the points
which had been raised, and he must necessarily leave some
unanswered for that reason, and not because he undervalued
their importance.

He agreed with Canon Fowler as to the danger of making
too much assumption upon a matter such as mimicry—it
was a subject upon which he had no desire to dogmatise.
His position was rather this:—Supposing the theory that
such and such forms were inedible seemed to supply a
provisional explanation of observed facts, it was desirable
to work out fully the logical consequences of such a
theory, and then to make a fresh appeal to observers for
verification.

Mr. Elwes had laid stress on the importance of getting
more observations from naturalists resident in the country

c

( -xkmi- ))

where these phenomena existed; with that the speaker fully
agreed.

With respect to Mr. Elwes’s question as to the reason for
concluding that the various members of a ‘ Millerian ”’
group were inedible; that was an inference which was
drawn from several data, in some instances resting on
direct observation, in others depending mainly on the
accordance of the characters exhibited with the logical
requirements of the theory. Cases of the latter .kind
awaited verification. |

The suggestion made by Mr. Rothschild with regard to the
similar results produced by a similar environment did not
remove the difficulty ; for these effects were not uniform, and
even closely allied species inhabiting the same region might
differ widely in aspect.

[From the PROCEEDINGS OF THE AMERICAN ASSOCIATION FOR THE
ADVANCEMENT OF SCIENCE, VOL, XLVI, 1897. ]

MIMICRY IN BUTTERFLIES OF THE GENUS HYPOLIMNAS AND ITS BEARING ON
OLDER AND MORE RECENT THEORIES OF MIMICRY. By Prof. EDWARD
B. Poutton, M.A., F.R.S., University of Oxford, England.

Tue theory of mimicry suggested by H. W. Bates in 1862 explained
the superficial resemblance of a rare to a common species in the same
locality by supposing that the latter possessed some special means of de-
fence (such as unpleasant taste, smell, etc.), and that the former, without
the special defence, was mistaken by enemies for the latter, and thus es-
caped a considerable amount of persecution. The relation may be com-
pared to that existing between a successful well-known firm and another
small unscrupulous one which lives upon its reputation. On the other
hand, Bates thoroughly recognized the existence of resemblance between
the specially defended forms themselves. These he could not explain by
his theory of mimicry, and suggested that they were a result of the in-
fluence of locality. Many years later Fritz Miiller satisfactorily explained
this difficulty by suggesting that a common type of appearance simplified
the education of enemies and thus was the means of saving life. The
lives of many individuals must be sacrificed before enemies have learned
to recognize and to avoid the colors and patterns which indicate some
special means of defence, and the fewer such patterns in any locality the
smaller the sacrifice. The relation may be compared to that between two
successful firms which combine to use a common advertisement.

This latter theory, although received rather coldly at first, las gradu-
ally made way, and seems now likely to occupy a good dealof the ground

(242)

943 ZOOLOGY.

formerly believed to be covered by the former theory. Thus, Dr. F. A.
Dixey, of Oxford, has recently shown that South American Heliconine are
affected by the color of certain Pier‘ne which have hitherto been looked
upon as true Batesian mimics of the former.

The old-world nymphaline genus Hypolimnas has been regarded as one
of the best examples of mimicry, but an unbiassed examination leads to
the opinion that it affords a case of Millerian rather than Batesian resem-
blance.

In India the female of the common species H. bolina resembles Huplea
core while the male is a dark butterfly with a large white spot, shot
with blue, on each of the four wings. Throughout the Malay Archipel-
ago representative species occur with males like that of H. bolina and
females reseinbling the local Euploeas. Occasionally, as in Ke Island and
the Solomons, species of the genus occur in which the male as well as the
female resembles a Huplea. In Fiji the male is as in the Indian species
while the female is extremely variable, ranging from forms like the male
through intermediate varieties, to be brown and straw-colored individ-
uals. The Euploeas of Fiji are not sufficiently known, but it is very
improbable that all the forms of the female Hypolimnas are mimetic. A
still more instructive case is that of the nerina form of female found,
with a male like that of H. dolina, in Australia, Celebes, New Guinea, and
other E. Indian Islands and in many of the Polynesian groups.

This conspicuous and abundant butterfly has, in addition to the four
white-and-blue spots of the male, a large reddish brown patch upon each
fore-wing. This well-marked form resembles no other butterfly except
the Danais chianippe of Celebes, and, as this latter appears to be very
rare, it is far more probable that the resemblance has come from the other
side, and that the Danais has approached the Hypolimnas.

In Africa the sub-genus Huralia is represented by several species which
resemble in both sexes species of the Ethiopian Danaine genus Amauris.

Finally there is the well-known and widespread Hypolimnas missippus
which accompanies Limnas chrysippus throughout its range; while the
female of the former resembles the latter very closely. In this case it is
certain that we have to do with no struggling hard-pressed form, for the
Hypolimnas has recently established itself in some of the West Indian Isl-
ands and in Demerara — localities in which its model, L. chrysippus, is as
yet unknown.

Tosum up —the genus Hypolimnas is distinguished among nymphaline
genera for the extent to which its numerous and widespread species re-
semble the local distasteful forms of Euploeins or Danaine.

Upon the older theory of Bates this would be explained by supposing
that the genus is very hard-pressed in the struggle and has thus been
driven to mimicry almost everywhere. Upon the newer Miillerian theory
it is supposed that the genus is distinguished among nymphaline genera
by some special defence, probably in the way of taste or smell or indi-
gestibility, and that it has been to its advantage to adopt the advertise-

SECTION F. 244

ment of still better-known and probably still more distasteful forms in
its locality.

The abundance of the various species, the conspicuous nerina form of
female, and the resemblance of a rare Danaid to it, the recent spread of
H. misippus beyond the limits of its model all support this latter inter-
pretation.

MIMICRY AS EVIDENCE FOR THE ANCESTRAL HOME OF A WIDE-RANGING
specius. By Prof. Epwarp B. PouLton, M.A., F.R.S., University
of Oxford, England.

We know on historic evidence that Anosia plexippus has spread and is
spreading through the warmer parts of the world. If we had not this
evidence, it might still be inferred with safety that North America is-
the ancestral home as compared with the other countries now inhabited
by this species. In North America we find a very perfect mimic in
Limenitis misippus, while in no other country has the occupation been
long enough to permit of such a resemblance growing up.

Similar evidence indicates that Africa is the ancestral home of Limnas
chrysippus which now ranges through almost all the warmer parts of the
Old World. The far greater effect which has been produced by this
species upon the Lepidopterous fauna of Africa as compared with that of
other lands, proves a far longer sojourn in the former.

A METHOD OF LABELLING TYPE SPECIMENS IN COLLECTIONS OF INSECTS.
By Prof. Epwarp B. PouLTon, M.A., F.R.S.,University of Oxford,
England.

THE method adopted in the Hope Department of the University Museum,
Oxford, consists in printing a form of type label to which the reference
to the author’s description of the species can be added. Such a label is
placed on the pin, below the specimen, while a duplicate label is pinned
beside it, in the most convenient position for the student. All essential
facts are recorded on this label and a second one stating the locality, date
of capture, name of captor, and date of presentation. This latter (which
is prepared for all specimens recently added to the collections) is simi-
larly placed beside as well as on the insect. Ordinary printers’ ink is used
for both labels because of its permanence; but the type labels are distin-
guished by a redline just within the margin.

Theories of Mimicry, as vlustrated by African Butterflies.
By Epwarp B. Poutron, WA., 7. R.S., Hope Professor of Zoology, Oxford.

(An abstract of the paper read before Section D of the BRITISH ASSOCIATION at
Toronto on Friday, August 20, 1897. Reprinted from the ‘Report’ of the
Meeting, pages 688-91.)

H. W. Bates, in his epoch-making paper (‘ Trans. Linn. Soc.,’ vol. xxiii.
1862), first gave an intelligible theory of mimicry, and accounted for the superficial
resemblances which had been known for so long by supposing that the most
dominant, well-defended, and conspicuous forms in a country become the models
towards which natural selection leads many of the weaker hard-pressed species
in the same locality. The material on which Bates’ theory was formed was con-
fined to tropical America, and his generalisation remained incomplete until it
could be applied to the other great tropical regions. This want, however, was
soon supplied by A. R. Wallace for the East (‘ Trans. Linn, Soc.,’ vol. xxv. 1866),
and by Roland Trimen for Africa ( ‘Trans, Linn. Soc.,’ vol. xxvi. 1870).

In Bates’ original paper a certain class of facts—frequently mentioned and
abundantly illustrated—cannot be explained under his theory of mimicry. This is
the strong resemblance which is apt to exist between the dominant forms them-
selves, and which is as minute and as remarkable as the resemblance of the weaker
for the stronger species. Bates pointed out that this was unsolved by his theory,
and both he and Wallace were compelled to suggest the direct action of some
unknown local influence as the possible cause. There the matter rested until
Fritz Miiller, in a paper published in Kosmos for May 1879, suggested an explana-
tion, viz., that the dominant forms gain an advantage by this resemblance, inasmuch
as it facilitates the education of their enemies by giving them fewer patterns to
learn. The necessary waste of life by which the education of young birds, &c., is
brought about is here divided between the various species of a closely convergent
group, instead of being contributed by each member independently. The chief
sub-families of butterflies which in tropical America appear to be specially dis-
tasteful to insect-eating animals, and which are specially mimicked by others, are
the Danaine, Ithomiine, Heliconine, and Acreine. Of these the second and
third are confined to this part of the world. The resemblances which Fritz Muller
explained are those which occur very commonly between the Danaine, Ithomine,
Heliconing, and less commonly the Acreine of any locality. In order to complete
this theory it was necessary to test its application in other parts of the world.

In the East the butterflies which take the place of the four above-named sub-
families belong almost exclusively to the Danaine, the Acreine being represented
by very few species. The Danaine are, however, extremely rich in species, and
F. Moore first pointed out in ‘ Proc. Zool. Soc.,’ 1883, p. 201, that there is the same
relationship between the species of this dominant group that obtains between those
of tropical America. Not only do Danaine of very different genera closely re-
semble each other, but there is often a strong likeness between the species belonging
to the two chief divisions of the sub-family—the Danaina and Eupleima, As in
America, these resemblances are always between the species of the same locality.

While, however, Miiller’s theory received full confirmation from the facts observed
in India and the tropical East generally, no attempt has been made until now to
apply it to the African lepidopterous fauna. I have therefore examined this
fauna from the Miillerian standpoint, and find that in it too the same relationship:
can be traced.

The dominant distasteful groups of Africa are the Acreine, which have their
metropolis here, and the Danaine. The latter are chiefly represented by the
species of the peculiar African genus Amauris, and by the abundant and wide-
spread Danais (Limnas) chrysippus. I first looked for evidence of convergence

between the dereine and Limnas chrysippus, and soon found what appeared to
d 3

“3

be evident traces of it. Such species as Planema esebria (certain forms of), Acrea
petrea (female), A. oppidia, and, above all, A. encedon (lycia) bear a considerable
resemblance to L. chrysippus, inasmuch as all of them possess a dark tip to the
fore wing crossed by a white bar, as in the Danaine butterfly. Looking at the
near allies of these species and at the Acreine@ as a whole, we may feel confident
that this black-and-white tip is not an ancestral character of the group, but a
comparatively recent modification. Again, the fact that this character is some-
times more strongly developed in, and sometimes confined to, the female sex agrees
with the corresponding relationships in other parts of the world, and furthermore
supports the conclusion as to the recent acquisition of the markings.

Convergence between the Acreine and Danaine of the genus Amauris was
next looked for and many examples found. Thus Acrea johnstoni of Kast Central
Africa certainly suggests the appearance of one of the echeria group, such as
A, hanningtonz, found in the same locality; while in West Africa Acrea lycoa
resembles the black-and-white Amawris damocles and A. egialea. Similar resem-
blances in the West are to be seen between the large black-and-white females of
the numerous species of the Acreeine genus Planema and other Acrzeas in the
same locality, such as A. carmentis (female) and A. jodutta (female), while the
species referred to, of both Acreeine genera, bear some considerable resemblance to
an abundant West African black-and-white Danaine—<Amauris niavius. Similar
relationships occur in the South-East, where Acrzeas, such as Planema esebria
(white form of female) and P. aganice bear considerable resemblance to the abun-
dant black-and-white Danaines—Amauris ochlea and A. dominicanus.

It was of great interest to prove that the members of these convergent groups
oceur, not only in the same place, but at the same time. Mr. Guy A. K. Marshall
has kindly done this work, sending me several groups captured at one place in a
single day. At Malvern, near Durban, Natal, on March 6, and again on March
30, 1897, he captured Limnas chrysippus and several species of Acrea, with the
black-and-white tip to the wing. On March 27 he captured, in the same locality,
the black-and-white Planemas ( Acreine) P. esebria and P. aganice, together with
an abundant black-and-white Neptis (NV. agatha) and a closely similar day-flying
moth, Nyctimeris apicalis. It is very probable that these latter forms do not
tuimick in the Batesian sense, but are themselves specially defended and fall into a
Miillerian group. Mr. Marshall did not, on that day, capture any of the black-
and-white Danaine. Mr. D. Chaplin, however, on April 5, 1896, obtained at
Berea, a suburb of Durban, Amauris ochlea and Planema aganice, as well as
Limnas chrysippus, with two species of convergent Acreeas (A. encedon and A.
petrea). Mr. ¥. D. Godman and Mr. O. Salvin have kindly presented these
specimens to the Hope Collection at Oxford.

I think it must be admitted that there is now strong evidence for the same
convergence between specially protected abundant African species from the same
locality as that which is already well known in the tropical East and in tropical
America. Various degrees of perfection exist, and it is in every way probable
that the resemblance of some members to the standard of their group is not of
long standing, and will improve in the future.

Other facts in the colouring of African Lepidoptera also support this interpret-
ation. Thus certain Lycenide of the genera Pentila and Alena are known to
fly very slowly, and in the case of the latter to feign death when captured—cha-
racteristics of unpalatable forms. While they thus differ in habits from Lycenids
generally, they also differ entirely in their appearance, which rather suggests that
of an Acrea. The same is true of moths belonging to many groups, and perhaps
of the abundant butterflies of the genus Bybla. Similarly the large group of
Lepidoptera, which has for its centre the abundant day-flying moths of the genus
Aletis, appears to be moulded upon the colouring and pattern of Limnas chry-
sippus, differing only in an even greater conspicuousness, due to the white spots or
rings on the black body, and the highly developed black-and-white border to the
hind wing. It is probable that the common species of the genus Huphedra, which
form some of the most conspicuous members of this group, are themselves specially
protected, To take one more example, certain species of the Pierine genus My/othris

3

are rendered specially conspicuous by the interrupted black border to the hind
wings, the interruptions extending along the hind margin of the fore wings. A
white butterfly with such a border becomes an extremely conspicuous object, and
this appearance of Mylothris is mimicked, more or less perfectly, by species from
a number of Pierine genera, such as Nepheronia, Belenois, Callosune, &c. This is
usually explained as an example of true Batesian mimicry, but it is, perhaps, more
probable that the Pierine are very largely a specially protected group, many of
the genera of which, so to speak, combine their advertisements, and thus share
between them the loss of life which must necessarily ensue during the education
of each generation of their enemies.

I think sufficient evidence has been brought forward to show that the theory
of mimicry, or rather of common warning (synaposematic) colours, which will
always be associated with the name of Fritz Miller, may claim abundant examples
in Africa as well as in the other parts of the world in which it has already been
proved to hold.

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Protective Mimicry as Evidence for the Validity of the Theory of Natural
Selection. By Epwarp B, Pouuron, W.4., F.RS., Hope Professor of
Zoology, Oxford. ie

(An abstract of the paper read before Section D of the BRITISH ASSOCIATION at
Toronto on Monday, August 23, 1897. Reprinted from the ‘Report’ of the
Meeting, pages 692-4.)

Several suggestions have been put forward to account for the superficial
resemblances between animals, especially insects, occupying the same geographical
area. It has been suggested, and indeed strongly maintained, that food, climate,
or some other chemical or physical influence of the locality may have supplied
the cause. On the other hand, many naturalists consider that the facts cannot be
interpreted by any of these suggested causes, and only receive an intelligible and
probable explanation in the theory of natural selection. This theory supposes
that the resemblance is advantageous in the struggle for existence, the weaker
forms being shielded by their resemblance to the strong and well-defended species
(mimicry of H. W. Bates), or the latter gaining by a resemblance which enables
their local enemies more easily—and thus with a smaller waste of life—to recognise
and avoid them (mimicry of Fritz Miiller). The present paper directs attention
to certain facts commonly associated with mimetic resemblance which receive a
ready explanation upon the theory of natural selection as the efficient cause, but,
on the other hand, constitute a serious difficulty in the way of any other theories
as yet brought forward.

Natural selection, as is well known, acts upon any variations, whatever they
may be, which are in the advantageous direction, and are at the same time not
injurious in themselves. When the end to be gained (in this case the attainment
of a superficial resemblance) is common to a variety of distantly related species
possessing entirely different constitutional tendencies, we may feel confident that
an approach brought about by natural selection will be by extremely diverse paths
of variation. Under natural selection we might predict that such a common end
would be reached by great diversity of means, while under the other hypotheses
mentioned above a result of the kind is inexplicable. Hence the facts of the case
should act as a convenient test between these rival suggestions.

First as to colour. We know but little of the chemical nature of the pigments
made use of in mimetic resemblance. One case, however, has been investigated
by Gowland Hopkins—viz., the bright tints by which certain S. American Prerine
have come to resemble Heliconine and Ithomiine in the same locality. Gowland
Hopkins has shown that these close resemblances in colour and pattern are
produced by pigments which are characteristic of the Pierine, and of an entirely
different chemical nature from those of their models.

Another very interesting case is that of resemblance to ants. Ants are
. mimicked more or less closely by a great variety of insects and by spiders. In
some cases we find the resemblance brought about by actual alterations in the
shape of the body (spiders and many insects), which is modified into a superficial
resemblance to the Hymenopteron. In an Acridian—Myrmecophana fallax—the
shape of an ant is, as it were, painted in black pigment upon the body of the
insect, which is elsewhere light in colour and, as it is believed, inconspicuous in
the natural environment. In a certain group of Homoptera—the Membracide—
some of the 8. American species closely resemble ants. The Membracide are
characterised by an enormous growth from the dorsal part of the first thoracic
segment (pronotum), which spreads backwards and covers the insect like a shield.
In these insects the form of an ant is moulded in the shield beneath which the
unmodified body of the insect is concealed. These facts are only explicable by
Supposing that some great advantage is to be gained by resembling an ant, and

6

that very different species have attained this end, each by the accumulation of
those variations which were rendered possible by its peculiar ancestral history and
present constitution—in other words. by the theory of natural selection.

A more elaborate case, which I have recently investigated, is afforded by a
large group of tropical American Lepidoptera—moths as well as butterflies—which
closely resemble certain common wide-spread species of the Ithomiine genera
Methona and Thyridia. The appearance thus produced consists of a transparent
ground with a black border to both wings, the fore wing being also divided by
black transverse bars into three transparent areas—the hind wing usually into
two. From a comparison with other species of the various families, &c., not
altered in this direction, we know that the transparent wings are not ancestral.
When we investigate the manner in which transparency has been attained, it is
found to be by different methods in the different constituents of the group. Among
the numerous genera of Ithomiinze (Methona, Thyridia, Dircenna, Eutresis, Ithomia,
&c.) the result has been attained by the reduction of the scales to a very minute
size, so that they hardly interfere with the passage of light. This reduction
affects the two kinds of scales which alternate with each other in the rows upon
the wings of this sub-family, a common result being (e.g., in Methona and Thyridia)
the alteration of the more slender scales into hairs, and of the broader ones into
minute bifid structures, still retaining scale-like proportions in spite of their
extremely small size. In others, again, the two kinds of scales are reduced
respectively to simple and Y-shaped hairs, which regularly alternate along the
rows. In the Danaine proper, represented by the genus Ztuna, the transparency
is chiefly due to the great diminution in the number of the scales, and those
which remain are neither much reduced in size nor altered in shape. In the
Pierine, represented in this group by only a single species, Dismorphia orise, the
scales are greatly reduced in size, but are neither greatly altered in shape nor
diminished in numbers.

Hence in these three sub-families of butterflies transparency is attained in
three different ways, viz. (1) by reduction in size and simplification in shape;
(2) by reduction in number ; and (8) by reduction in size alone.

When we examine the moths which fall into the group, we find a much
greater difference in the methods, corresponding to the wider divergence in
affinity. In the several species of the genus Castnia the scales lose their pigment,
although undiminished in size, while they are at the same time set vertically upon
the wing, so that light can freely pass between their rows. In the widely sepa-
rated genus Hyelosia the arrangement is nearly the same, except that the vertical
scales are much attenuated. In the genus Anthomyza, which furnishes the group
with many species, the scales retain the normal size, shape, and overlap, but
become so completely transparent that the light freely passes through them.

In all the numerous constituents of this large group of Lepidoptera a very close
resemblance has been produced by: entirely different methods; a result which, it
has been argued above, is only consistent with the view that natural selection
alone, among all the explanations which have been suggested, has been the cause
of the observed phenomena.

I owe to the kindness of Mr. Godman and Mr. Salvin the opportunity of
studying all the butterflies of this large transparent-winged group, while Mr.
Herbert Druce kindly lent me those moths which are not represented in the Hope
Collection in the Oxford University Museum.

[Extracted from the Linnean Socrety’s JourNAL—Zoowoey,
vol. xxvi. |

Natural Selection the Cause of Mimetic
Resemblance and Common Warning

Colours.

BY

EDWARD B. POULTON, M.A., F.RS.,

HOPE PROFESSOR OF ZOOLOGY IN THE UNIVERSITY OF OXFORD,

558 PROF. E. B. POULTON: NATURAL SELECTION

Narurat SELEctTIoN the Cause of Mimetie Resemblance and

a

Common Warning Colours. By Epwarp B. Povutron,
M.A., F.R.S., Hope Professor of Zoology in the University

of Oxford.
[Read 17th March, 1898. ]

(Piates 40-44. )

ConreENTS.

Sblistoricalmlintroduction..s enuraseasessecesmecstmreeceeen cere: seer ee aee eet 558
. The various Hypotheses which have been proposed as Substitutes

for Natural Selection as the explanation of Mimicry and Common
Warning! OGlours: sosso.- sae emeeseee cece eseaene sme a se neers e eee eee eee eens 563

. The Relation of the Resemblances under Discussion to other Resem-

blances in Organic/ Nature: focseces. ac seeenuen= scene: peuscrb ease aneeee serene 565

. Mimetic Resemblance and Common Warning Colours between different

Arthropod Classes and between various Insect Orders, and their
Relation to Similar Resemblances within the Limits of a Single Order 557

. Resemblances even within the Limits of an Order are entirely in-

dependentioivAdh nity ir -n..sssococtserceseaece sacs: seer eecer eset 572

. The Resemblances in question are not accompanied by any changes in

the direction of another Species, except such as assist in the pro-
duction or strengthening of a Superficial Likeness ..................... 574

. Essential Nature of these Resemblances: their Analysis into the

several kimdsior Mitectiproduced sacs.c rss sdeterececeetes nent eh eea at eneee 576

. Conditions of a Species in any Locality are chiefly determined by its

iMabitsand WWite-history, oecdeucosceccosscresesessecesceeneete ecco eee aeeenere 578

. Mimetic Resemblance and Common Warning Colours more charac-

teristic of the Female than the Male Sex ...............ce.ssesccsesseceee 580

. The Space and Time Relationships of the Resemblances in Question... 582
. The Resemblances which Insects of various Orders bear to those of

another Order are produced in the most Diverse Ways ............... 585

. The Resemblances within the Limits of the Order are also produced

INbhe most) DiversewWV AS: cen-s-cedeacs-cseeceorsseee secon ences eaeeeeee 595

. The supposed Direct Effect of Local Forces implies the Hereditary

Mransmission of Acquired (Charactersiisss.-sece-e-sosn cesses eee eeeeee 601

. General Conclusions: Natural Selection as the Cause of Mimetic

Resemblance and Common Warning Colours ...........seeeceeeeneeeees 601

(1) Historical Introduction.

SUPERFICIAL resemblances between animals, especially numerous
in Insecta, were known long before H. W. Bates’s paper “ Uon-
tributions to an Insect Fauna of the Amazon Valley ” was read
before the Linnean Society on November 21st, 1861, and published
in the ‘Transactions ’ the following year (vol. xxi. p. 495). Some

THE CAUSE OF MIMETIC RESEMBLANCE ETC. 559
of the principal records of such observations are to be found in
the ‘ Transactions’ of this Society.

W. S. MacLeay, in his ‘ Hore Entomologice’ (London, 1819
and 1821), alluded to certain cases which are now included under
Mimiery, viz. the likeness of some Diptera to Hymenoptera
(Pt. IL. p. 365), and interpreted them, together with many other
resemblances of structure and life-history, by the principle of
Analogy as distinct from Affinity in Nature. These views were
adopted by Macleay’s immediate successors.

The Rev. William Kirby read “ A Description of some Insects
which appear to exemplify Mr. William S. MacLeay’s Doctrine
of Affinity and Analogy,” before this Society on Dec. 17th, 1822,
and the paper was published in the ‘ Transactions’ (vol. xiv.
p- 93).

Boisduval, in the ‘Spécies Général des Lépidopteres,’ pub-
lished in 1836, remarked (pp. 372, 373) on the resemblance
between certain West African butterflies belonging to very
different groups.

Prof. Westwood read ‘ Jllustrations of the Relationships ex-
isting among Natural Objects, usually termed Affinity and
Analogy, selected from the Class of Insects,” before this Society,
on Jan. 17th and May 2nd, 1837, the paper appearing in the
‘Transactions’ (vol. xvii. p. 409). In the paper many new
examples were published and figured, while MacLeay’s views
were criticised and expanded in an interesting manner.

The same recognition of Mimicry is equally well seen in the
names with the termination -formis given to so many of our
moths, indicating their resemblance to wasps, bees, and other
insects. In spite, however, of the knowledge of a large number
of instances, the subject made no real progress until the appear-
ance of H. W. Bates’s paper. The view then set forth that the
resemblances are in themselves beneficial to the possessor was, as
far as [am aware, only once suggested before,—in the well-known
‘ Introduction’ by Kirby and Spence. In Vol. II. p. 228 of the
second edition (1817) the authors write as follows :—‘ Some
singular larve with a radiated anus live in the nests of humble-
bees, and are the offspring of a particular genus of flies (Volw-
cella, Geoffr., Pterocera, Meigen), many of the species of which
strikingly resemble those bees in shape, clothing, and colour.
Thus has the Author of nature provided that they may enter
those nests and deposit their eggs undiscovered. Did these

40*

550 PROF. E. B. POULTON : NATURAL SELECTION

intruders venture themselves amongst the humble-bees in a less
kindred form, their lives would probably pay the forfeit of their
presumption.” This interesting paragraph, although fully re-
cognizing the utility of mimetic resemblance in species which
were then believed to have been separately created and to have
come into existence fully formed and complete, sustains a posi-
tion which is the very antithesis of that taken up by Bates. The
contention that the utility of the resemblance has been the
cause of its persistence, and, by the selection of variations going
further in the same direction, of its improvement, would have
been rejected, probably with indignation, by the distinguished
authors of the ‘ Introduction,’

Bates’s great paper dealt with the fauna of tropical America,
and the generalization was manifestly incomplete until it had
been extended to other parts of the world. This confirmation
was not long in coming, being supplied for the tropical East by
A. R. Wallace’s paper published:in the ‘Transactions’ of this
Society (1866, vol. xxv.), and for Africa by Roland Trimen’s
paper, also to be found in our ‘ Transactions ’ (1870, vol. xxxi.).

Tt is remarkable how completely the Linnean Society has been
the medium for the publication of classical memoirs upon
Mimicry. Up to the year 1870 it contained them all; while in
1858 it served as the channel through which the parent theory
of Natural Selection was first given to the scientific world.
The next great advance did not take place until 1879, and was
published elsewhere.

Bates had called attention to certain resemblances which could
not be interpreted under his theory of mimicry, viz. the frequent
similarity between the specially defended forms themselves.
Forms which are themselves the models for mimicry nevertheless
mimie or at least resemble other models. For such cases Bates
could only suggest the direct action of some unknown local force
or forces, and Wallace followed him in this.

In May 1879 Fritz Miller published a paper in ‘ Kosmos’ *
which for the first time offered an explanation, based on the
theory of natural selection, of these mysterious resemblances.
He suggested that such likeness between dominant forms was
advantageous to them, inasmuch as it facilitated the education of
their enemies, reducing the amount of destruction which must
be wrought during the time in which young birds and other

* “ Ttuna and Thyridia: a remarkable case of Mimicry in Butterflies.”

THE CAUSE OF MIMETIC RESEMBLANCE KETC, 56L

animals are learning what to eat with impunity and what to
reject. The paper was translated by Prof. R. Meldola and pub-
lished in this country almost immediately after its appearance
(Proce. Ent. Soe. Lond. 1879, p. xx).

The facts which the Millerian theory sought to explain con-
cerned the fauna of tropical America; the naturalist who ex-
plained them was a resident in the same part of the world. A
few years later, however, I’. Moore (Proc. Zool. Soe. 1883, p. 201)
showed that there is the same resemblance between the dominant
buttertlies of the tropical East; and last year I pointed out that
the same facts hold in Africa (Report of the British Association
at ‘Toronto, 1897, pp. 683-691). See also Roland Trimen’s
Presidential Addiess to the Hutomological Society, Jan. 19th,
1898 (Proceedings, 1897, p. Ixvi). In the ‘ Proceedings’ of
the Entomological Society (1897, p. xxix), as well as in the
former paper (p. 691), I argued that such resemblance is not true
Mimicry at all, but rather an example of Common Warning
Colour, and with the assistance of Mr. Arthur Sidgwick I sug-
gested the term Synaposematic as descriptive of it; the term
Aposematic having been previously suggested for ordinary
Warning Colours (Poulton, Colours of Animals, Internat. Sci.
Ser., London, 1890, p. 337).

I have now given a brief account of the leading phases in
the history of Mimicry. Even before the appearance of Fritz
Miller’s paper a great effect had been produced. This immediate
stimulus to the investigation of new examples and fresh aspects
of Mimicry which followed Bates’s memoir, must be ascribed to
the fact that then for the first time was offered a good working
hypothesis—a hypothesis which seemed to afford an adequate ex-
planation of one class of known facts, which challenged its critics
to find insuperable difficulties among facts as yet unknown. In
the thirty-six years which have elapsed since the appearance of
this great memoir an immeuse number of facts bearing upon the
subject have been discovered, and many naturalists consider that
Bates’s theory of Mimicry as due to natural selection (sup-
plemented and completed by the kindred theory which we owe
to Fritz Muller) has stood the test with complete success and is
in afar stronger position than in 1862. This opinion is more
generally held among the students of other groups of the animal
kingdom than among those who are specially devoted to entomo-
logy, but a considerable proportion of the latter also hold it firmly.

562 PROF. E. B. POULTON : NATURAL SELECTION

In the summer of last year (1897) an excellent opportunity of
gauging the opinion of entomologists upon this question was
presented by a discussion carried on at two successive meetings
of the Entomological Society of London. From the printed
report in the ‘ Proceedings’ of the Society, I select the following
adverse expressions of opinion as regards the adequacy of
natural selection to produce mimetic resemblance, and the state-
ments of any alternative theories or suggestions to account for
the facts. The quotations are given in the order in which they
are printed.

Mr. W. F. H. Blandford said “it was conceivable that the
causes, in most cases unknown, which brought about modifica-
tions in the colour and markings of a species in association with
its geographical range, might have produced identical results
in two species of the same genus, with a common facies, under
common conditions.” This sentence seems to suggest cautiously
the direct influence of climate or some other cause connected
with locality. Mr. Blandford, although not disputing the theories
of Bates and Fritz Miiller, considered that ‘they rested very
largely on hypothesis and were in want of further support from
observation and experiment,” and that they failed to account
for the perfection of the resemblance.

Mr. H. J. Elwes thought “there was too much assumption
about either the Batesian or Miillerian theories of mimicry,’ and
he doubted the inedibility of the so-called models, and of the
members of the Miillerian groups. Some examples were quoted
by him as suggestive of similar effects wrought by similar con-
ditions of environment.

Mr. Jacoby considered that the evidence of special protection
was insufhcient.

Sir George Hampson “ thought the cause demanded by these
theories was inadequate to produce the results assigned to it.”
In 8. India he had found it ‘‘ quite an exceptional thing to see
birds catch, or even attempt to catch, butterflies.”

The Hon. Walter Rothschild considered ‘‘it was much more
conceivable that certain climatic influences, &c., had played a
part in bringing about these resemblances, and he thought that
these groups assumed the same appearance because one given
influence was at work on them.”

Canon W. W. Fowler thought “there was too much assump-
tion about the current theories.”

HE CAUSE OF MIMETIC RESEMBLANCE ETC. 563

Mr. McLachlan suggested “the possibility that two species
might go on independently and yet apparently mimic each other
by arriving at the same results in their modifications.”

Mr. J. J. Walker and Col. Yerbury argued that birds are not
“ effective agents in causing mimetic resemblances.”

Mr. Blandford, replying at the close of the debate, maintained
that extremely close and exact resemblances are ‘“ hypertelic,”—
going beyond the limits of the useful, and therefore inexplicable
by natural selection. He restated and brought further support in
favour of the opinion that birds rarely attack butterflies. He
made the final suggestion that possibly “ sexual selection, or the
segregation of forms might take place as a direct act of perception
on the part of the insects themselves”; and quoted the observation
of H. W. Bates that the [¢éhomie when pairing tend to select none
but their exact counterparts (Trans. Linn. Soe. vol. xxxiii. 1862,
p: 501).

Only four speakers gave full support to the theories of Bates
and Fritz Miller, as explaining the facts more satisfactorily than
any other theory hitherto proposed and as likely to receive
further confirmation in the future. These four were the President
Mr. Roland Trimen, Dr. Dixey, Colonel Swinhoe, and the present
writer.

(2) The various Hypotheses which have been proposed as Sub-
stitutes for Natural Selection as the Explanation of
Mimiery and Common Warning Colours.

In the course of the discussion thus briefly epitomized, almost
every alternative theory which has been proposed to account for
the facts was brought forward.

These theories are as follows :—

(1) The direct effect of some physical or chemical cause or
causes connected with locality, such as climate, food, &e. This
hypothesis received much support. It may be called the Theory
of External Causes.

(2) The independent evolution of a similar appearance in
distinct species. This hypothesis was suggested by Mr.
McLachlan, and I have heard a similar opinion expressed by
Prof. Patrick Geddes. This appears to be the view sustained
in G. H. Eimer’s just published work ‘ Orthogenesis der Schmet-
terlinge’ (Leipzig, 1898). I have not yet had the opportunity
of studying the essay, but my friend Prof. Weldon has done so

564 PROF. E. B. POULTON: NATURAL SELECTION

and tells me that it supports a theory of mimicry as due to
internal developmental causes, compelling different species to pass
through similar phases. The hypothesis that “laws of growth”
may cause these resemblances also falls into this category. The
suggestion that such laws may account for certain phenomena
which are usually explained by the theories of natural and sexual
selection was made by Prof. D’Arcy Thompson at the Oxford
meeting of the British Association in 1894. This may be called
the Theory of Internal Causes.

(8) The operation of Sexual Selection, cautiously suggested as
a possibility by Mr. Blandford, who did not, on this occasion,
develop the details of the manner in which the principle may be
supposed to act. Fritz Miller had suggested this idea in a
letter to Darwin, who wrote not unfavourably of it to Prof.
Meldola, on Jan. 23rd, 1872. ‘ You will also see in this letter
a strange speculation, which I should not dare to publish,
about the appreciation of certain colours being developed in
those species which frequently behold other forms similarly
ornamented. I do not feel at all sure that this view is as in-
credible as it may at first appear. Similar ideas have passed
through my mind when considering the dull colours of all the
organisms which inhabit dull-coloured regions, such as Patagonia
and the Galapagos Islands” (‘ Charles Darwin and the Theory of
Natural Selection,’ Poulton (London, 1896), p. 202.)

In the present paper the attempt will be made to show that
many of the known facts of mimetic resemblance do not admit
of interpretation by any of the three theories mentioned above,
while they do receive a ready explanation on the supposition
that the resemblances are useful and have been produced by
natural selection. Certain new observations upon the details of
the resemblances in a large group of insects, undertaken specially
to test these rival theories, will be found to point strongly in the
same direction (see Section 12).

In order to render the argument as complete as possible,
various considerations which have been urged before will be
included as well as those which now appear for the first time.

The objection may be raised that such detailed treatment is
unnecessary, and that biologists generally agree in attributing
these resemblances to natural selection. To this contention the
discussion at the Entomological Society is sufficient answer.

THE CAUSE OF MIMETIC RESEMBLANCE ETC.

(3) The Relation of the Resemblances under Discussion to other
Resemblances in Organic Nature.

To those who accept natural selection as the explanation of
mimicry, the facts under discussion fall into their place as part
of the much wider group of Protective Resemblances in general.
Mimicry becomes merely “an exceptional form of protective
resemblance’ (Wallace, ‘ Darwinism,’ London, 1889, p. 265).
The following classification was suggested by the present writer,
with the assistance of Mr. Arthur Sidgwick, in 1890 (‘ The
Colours of Animals,’ Internat. Sci. Ser., London, 1890, p. 338):—

III. Epigamic
colours. — Co-
lours dis-
played in

II. Sematic colours.—
Warning and signal-
ling colours,

I. Apatetic colours.— Colours resem-
bling some part of the environment or
the appearance of another species.

A. Cryptic colours.
Protective and Ag-

B. Pseudosematice
colours. — False

courtship.

gressive Resem- | warning and sig-
blances. nalling colours.

i Procryptic 1. Pseudapose- 1. Aposematic
colours, — Protec- | matic — colours.— } colows. — Warning
tive Resemblances. | Protective Mimi- } colours.

cry.

2. Pseudepisema-
| tic colours. — Ag-
gressive Mimicry
and Alluring Co-
loration.

2. Anticryptic
colours. — Ageres-
sive Resemblances.

2. Eptsematie co-
lours. — Recognition
markings.

Thus the facts of mimicry fit into a broad system which includes
the other resemblances in organic nature. The relation between
protective resemblance (I. A.1) and protective mimicry (I. B. 1)
is as follows :—In the former an animal resembles some object
which is of no interest to its enemy, and in doing so is concealed ;
in the latter an animal resembles an object which is well known
and avoided by its enemy, and in doing so becomes conspicuous.
Thus mimicry as interpreted by H. W. Bates finds its place in
I. B. 1, while the resemblance between protected conspicuous
forms (sometimes, but, as I think, erroneously, called mimicry),
as interpreted by Fritz Miller (‘ Kosmos,’ May 1879, p. 100,
translated by Meldola in Proc. Ent. Soc. Lond. 1879, p. xx),
falls into II. 1. Such cases only differ from ordinary warning
colours in that they are common to two or more species : hence,

5€6 PROF. E. B. POULTON: NATURAL SELECTION

as has been already pointed out, the term synaposematic colours,
or common warning colours, may be conveniently applied to
them.

The arguments in favour of natural selection as the explanation
of protective resemblance run entirely parallel with those which
favour it as the interpretation of mimetic resemblance and
common warning colours. By modifying the examples and, in
some cases, the form of the argument, nearly every section of
this paper might be converted into a defence of the former, and
the arguments which are strongest in support of the one are the
strongest in support of the other, viz. those contained in Sections
4, 5, 6, 7, 11, and 12.

Under the theory of natural selection all the resemblances
among animals, mimetic and other, show the clearest relationship,
and (with the exception of the debated Epigamic colours) are to
be explained by the working of a common principle, viz., the
selection of variations which are useful in the struggle for
existence. Under the other theories mentioned above no such
grouping can be readily brought about, and mimetic resemblance
becomes due to one set of principles and the other resemblances
to another set. The majority of those who look on mimicry as
due to external or to internal causes, or to sexual selection, would
probably agree in explaining protective resemblance by natural
selection. And yet these latter cases, while far more common,
are often as detailed and as remarkable as those of mimicry.
Those who adopt the most extreme form of the theory of external
causes might perhaps maintain that the resemblance to twigs,
leaves, and bark is to be explained in the same manner, and would
thus bring protective and mimetic resemblance under the operation
of the same set of forces ; but few will be prepared to carry the
theory so far. Under the theory of internal causes it is impossible
to bring the two kinds of resemblance together ; for while it is held
by some that two or more animals may independently and with-
out selection arrive at corresponding points in their evolutionary
history, which are such as to involve mimetic resemblance, no
one could believe that the similarity to bark or earth has been
produced in the same manner. ‘Those who are inclined to accept
sexual selection as the explanation can only bring the two classes
of facts together by supposing that the appearance of some
minute portion of the total vegetable or mineral environment
has acted as a stimulus and has led one sex to select the other

THE CAUSE OF MIMETIC RESEMBLANCE ETC. 567

according as it resembled the object in question ; just as a similar
selection has been supposed to take place from the stimulus
supplied by the appearance of another species. Probably no one
is prepared to adopt this view as regards the former class of
facts, although Darwin had, as the above-quoted letter shows,
considered the possibility of the general tints of the environment
influencing the trend of sexual selection in this way.

A fatal objection to any explanation based on the theory of
sexual selection is the fact that protective resemblances are so
extremely common and perfect in the immature stages of
insects. The same objection holds, although with less force,
against its use as an explanation of mimetic resemblance.

The conclusion appears inevitable that under no theory except
natural selection do the various resemblances of animals for their
organic and inorganic environments fall together into a natural
arrangement and receive a common explanation. On any theory
except natural selection this cin only be brought about by the
adoption of extreme views as to the area over which the alternative
theory is to be applied,—views which, at any rate, the great
majority of those who are disposed thus to explain mimetic
resemblance are not prepared to adopt.

(4) Mimetice Resemblance and Common Warning Colours between
different Arthropod Classes and between various Insect Orders,
and their Relation to Similar Resemblances within the Limits

of a Single Order.

The discussion at the Entomological Society was based almost
exclusively upon the phenomena presented by mimetic resemblance
and common warning colours affecting the species of a single
Order of insects (Lepidoptera), and generally the species of a
single Family (Nymphalide). I cannot but think that this
limitation of the survey to one small part of the field over which
the resemblances commonly occur is, in large part, the cause of
the rejection of natural selection aud the substitution of alter-
native suggestions. There is something attractive and plausible
in the suggestion that the strong mutual resemblances within a
group of butterflies of different genera and subfamilies, inhabiting
a single locality, are due to the direct action of peculiar local
physical or chemical influences; but the suggestion loses all its
attractiveness when it is applied to the resemblance between a

568 PROF. E. B. POULTON: NATURAL SELECTION

spider and an ant, or a moth and a wasp. And yet few could
bring themselves to believe that the resemblances which are here
contrasted have been built up by two entirely different sets of
forces. Mr. Blandford alluded to this relationship and gave it as
his main reason for accepting Buates’s theory, although he rejected
the Mullerian theory of Common Warning Colours. This dis-
crimination in favour of the former theory is not justified by the
facts. The resemblance between different insect orders has not
been as yet sufficiently regarded from the Miillerian standpoint ;
but there are, and have been for many years, the strongest indi-
cations that here also much of the ground formerly believed to be
covered by the older theory will be found to be oceupied by the
newer.

The Miullerian theory by no means demands that the methods
of defence in the members of a convergent group should be
uniform.

So long ago as 1887 (‘‘ The Experimental Proof of the Pro-
tective Value of Colour and Markings in Insects in reference to
their Vertebrate Enemies,” Proc. Zool. Soc. 1887, pp. 191-274)
I tabulated the colours and markings of all insects which up to
that time had been experimentally proved to be specially defended,
and was enabled to apply to the whole group of conspicuous
insects the explanation offered by Fritz Miller (/.¢. p. 227).
This general conclusion will be found to be supported by many
facts and considerations in the paper referred to.

In alluding to the resemblance which the black-and-yellow-
ringed unpalatable larva of Huchelia jacobee bears to a wasp, I
wrote (1. ¢. pp. 235, 288) :—“ Thus it is more than probable (as
has been previously suggested by other observers) that the species
reudered conspicuous by alternate rings of black and yellow gain
great advantages from the justly respected appearance of Hornets
and Wasps. It must not be forgotten, however, that the latter
forms also probably gain to some extent by the greater publicity
which follows from the resemblance.”

I may here digress for a moment and invite the attention of
those who dwell on the excessive amount of assumption in the
theories of mimicry, to the numerous tables in the paper mentioned
above. In these will be found recorded the whole of the results
of actual experiments made, up to 1887, upon the palatability
and unpalatability of conspicuous and inconspicuous insects. It
may be safely asserted that the theories in question are not nearly

THE CAUSE OF MIMETIC RESEMBLANCE ETC. 569

so devoid of support from the results of experiment and observa-
tion as has been represented.

Since 1887 further evidence has been forthcoming in support
of the Miillerian explanation ; for it has been shown in many cases
that insects which resemble specially defended members of another
order, themselves belong to a specially defended group within
their own order. Thus Haase (‘ Researches on Mimicry,’ part ii.
London, 1896, English translation by C. M. Child, p. 70) points
out that the South-American moths which resemble “immune”
Coleoptera—the Lycine, “ belong to the immune families of the
Glaucopide (Mimica, Lyeomorpha) and Arctiide (Pionia),” and
also that (7. ¢. p. 73) the South-American Glauwcopide furnish
numerous cases of resemblance to Aculeate Hymenoptera. Ina
note to the same page Haase adduces some little direct evidence
for the inedibility of a Glaucopid.

During the past year Dr. L. O. Howard, of Washington, has
kindly presented to the Hope Department, Oxford University
Museum, a pair of specimens which prove that the protected moth
Lycomorpha latercula (Edw.) occurs in the same locality and at
the same time of the year as the beetle Lygistopterus rubripennis
(Lec.), which it closely resembles, the former having been captured
on June 18, the latter on June 5, 1897, in the Chiricahua Moun-
tains, Arizona, by Mr. H. G. Hubbard.

Furthermore, the resemblance between the species of the two
great sections of the order Lepidoptera—the Rhopalocera and
Heterocera—is frequently of the Miillerian rather than the
Batesian kind. Thus Sir George Hampson has pointed out that
the moth Abrawxas etridoides, resembling the butterfly TLeracolus
etrida, belongs to a specially protected genus, and that similarly
three genera of the Chalcosia group of Zygenide, which are said
to resemble Danaine and Papilionine butterflies, are also extremely
distasteful to insect-eating animals (‘ Nature,’ 1898, Feb. 7,
p. 364). Similarly, Mr. Roland Trimen, in his Presidential
Address to the Entomological Society (Jan. 19, 1898), points out
that the “abundant and extremely conspicuous, slow-flying,
diurnal Lithosiid moth Alctis helcita,” together with its “ appa-
rently protected analogues '....... the closely similar Lithosiid
Pheagarista helcitoides, and Agaristid Eusemia falkensteinit,”
show great similarity to the group which is headed by Danais
chrysippus,— so that from the aspect of warning of distasteful-
ness to enemies the two sets may be regarded as practically but

570 PROF. E. B. POULTON: NATURAL SELECTION

one.” Similar facts will probably be found in numerous other
examples of moths which resemble the butterflies.

It may be safely asserted that, even with our present limited
knowledge, Miilerian resemblance, no less than Batesian mimicry,
can be found in the species of groups with all degrees of affinity,
and that there is no ground for Mr. Blandford’s contention that
the latter alone derives support from the facts presented by the
groups which include species from different orders.

Under natural selection the interpretation of the whole series of
facts is perfectly valid. The domimant forms which in each locality
move towards each other and towards which less dominant forms
also move, are in some way specially defended. The principles
are the same when the approximation is between the species of
different orders or suborders, or between those which are much
more closely related. The Millerian theory explains the resem-
blance of such large numbers of stinging insects to each other
and of other specially defended forms to them, whether they be
closely or distantly related: it also explains the resemblance of
the dominant Heliconine and Ithomiine im each locality in South
and Central America and of other forms to them. Batesian
mimicry explains the cases in which the attracted forms are not
specially defended.

The conclusion which emerges most clearly is the entire inde-
pendence of zoological affinity exhibited by these resemblances ;
and one of the rare cases in which Darwin’s insight into a bio-
logical problem did not lead him right was when he suggested
that a former closer relationship may help us to understand the
origin of mimicry. Further confirmation of this conclusion will
be found in the additional details supplied in the succeeding
section.

When we look at the phenomena of mimicry and common
warning colours as a whole, it is found that the theory of natural
selection is equally applicable throughout ; while the theories of
external causes and internal causes cannot be applicd to some of
the most striking. 1esemblances, those of moths, beetles, and
Diptera to stinging Hymenoptera. ‘The theory of sexual selec-
tion is less logically assailable on these grounds; but with the
other two suggested substitutes for natural selection, it entirely
fails to account for the attractive force exercised by specially
protected forms. Under any of these three theories it is a mere
coincidence that the insects which are resembled by species of all

THE CAUSE OF MIMETIC RESEMBLANCE ETC. orl

kinds happen to possess stings—that the central types in the
groups of butterflies belong to subfamilies which are more
abundant and even more unpalatable than the generality of their
order. It is, furthermore, a mere coincidence that such groups
are formed round the Danaine and Acreine, wherever they oceur
in all the warmer regions of the world, aud in trepical America
also round the Ithomiine (Neotropine), which are closely related
to the former, and the Heliconine, which are closely related to
the latter.

No theory except natural selection explains why the number
of colours and patterns in the dominant groups of butterflies
mentioned above are so few in relation to the number of species,
as was pointed out by Prof. Meldola (Ann. & Mag. Nat. Hist.,
Dec. 1882). These colours and patterns have been recently
studied very carefully, especially in the Ithomiinag, by A. G.
Mayer (Bulletin of the Mus. of Comp. Zool. at Harvard Coll.,
Feb. 1897, p. 169). Mayer shows that “the 200 species of
Papilio in South America display 36 distinct colors, while the 450
species of Danaoid Heliconide |Ithomiine| exhibit only 15.
Hence the numbers of the species and of the colors are almost in
inverse ratio in the two groups. This may be explained by the
fact, that the Danaoid Heliconide mimic one another, while the
Papilios do not...... There is no lack of individual variability
among the species of the Danaoid Heliconide ; yet the species
as a whole vary but little from the two great types of color-
pattern represented by MMelinea and Ithomia. In order to
account for this remarkable fact, I am forced to resort to Fritz
Miiller’s theory of mimicry ” (J. ¢. p. 229). Again on page 225
Mayer remarks: “It is difficult to account for the remarkable
conservatism in respect to color-variations among the Heliconide
[here used, as in Bates’s original paper, to include Danaine, Itho-
miine, and Heliconine|, unless we resort to the explanation
afforded by the theory of mimicry; for, while there is such
remarkable simplicity and uniformity of color-pattern throughout
the whole group of the Heliconide, individual variations are very
common.”

It is not from any predisposition or bias in favour of natural
selection that these conclusions are reached, but simply because
natural selection offers an explanation of so many remarkable
facts which are utterly meaningless under any other theory yet
brought forward.

HT 2 PROF. E. B. POULTON: NATURAL SELECTION

(5) Resemblances even within the Limits of an Order are
entirely independent of Affinity.

The entire independence of affinity is specially well seen in the
groups of convergent moths and butterflies which are found in
different localities in South America. Although the resemblance
is clear enough in all the members of a large group, it is far closer
in certain species than in others. When these are examined they
are found not to be more nearly related than other members of
the group, but frequently the reverse. Thus it is very common
for a species of Heliconius to resemble with the most remarkable
precision a species of Melinea or some other Ithomiine genus in
its locality. Such resemblance is in these eases far closer than
that of the former to the species of the other genus (Hwetdes) in
its subfamily, and than that of the latter to any species in the
numerous related Ithomiine genera. Thus, to illustrate this
conclusion from some examples in the Hope Collection, in Hon-
duras by far the strongest resemblance is to be found between a
Heliconius and a Melinea; and this is also the case (both
species being different) inSurinam. In Trinidad the resemblance
is closest between a Heliconius and a Tithorea.

Under the theory of natural selection this association is readily
explicable. The pairs which thus form the centres of local
groups are probably the dominant forms, relying more com-
pletely than the other members upon the defence afforded
by their warning colours and the associated unpalatability.
As a matter of fact there is some evidence for their exceptional
abundance as compared with the other members of their groups.
Again, they are usually more nearly of the same size than the
other members, so that the perfection of the resemblance in colour
and pattern is aided by resemblance in another quality.

The theory of external causes entirely fails to account for
these facts. Uniform local conditions, if they can produce
any effect at all, must be expected to produce the closest likeness
where there is the closest constitutional similarity—in other
words, in the more nearly related forms, in preference to the less
nearly related, in each locality. With the theory of internal
causes we should also expect the facts to be the reverse of those
which exist. At the best it is unable to account for the observed

phenomena.
Any theory of selection (natural, artificial, or sexual) affords a

THE CAUSE OF MIMETIC RESEMBLANCE. 573

logical explanation of the facts, in the sense that it is quite con-
ceivable that the observed results might be thus obtained. Hence
the objection to sexual selection as a suggested cause is not as
strong as the objection to the other causes which have been
brought forward. Nevertheless, I believe that very few will be
found to support the former suggestion.

The conclusions here arrived at by a consideration of the facts
presented by the Lepidoptera are entirely confirmed by those
already known in the Coleoptera; although as yet but little
attention has been paid to the latter Order in this respect.
Mr. C. J. Gahan, in an interesting paper (Trans. Ent. Soc. Lond.
1891, pp. 867-374), clearly shows that the Phytophagous genus
Diabrotica is in the same position as the large protected groups
of butterflies already mentioned (Danaine, Ithomiine, Helico-
nine, Acreine). The individuals of its species swarm in the
localities where they occur ; they are conspicuously coloured, and
many of them are known to feign death when captured and to
discharge a yellow fluid from various parts. The facts at present
ascertained justify the conclusion that these Coleoptera form
centres of Miillerian resemblance, in that “some of the species
belonging to one section io this genus are, in colour and marking,
extremely like certain species of the other section which come
from the same localities” (J. ec. p. 872). Mr. Jacoby has also
“recorded that many of the species of his genus DWeobrotica
exhibit most striking resemblances to species of the closely
related genus Diabrotica.” The latter species are also mimicked
by those of the allied genus Dircema. Mr. Gahan shows,
furthermore, that 18 species of the genus Lema, belonging to a
different sub-family, closely resemble the species of Diabrotica (in
one case the allied genus Cerotoma) found in the same localities
in tropical America. In three cases species of Neobrotica, and in
one a species of Dircema, fall into the groups thus formed.

Mr. Gahan is disposed to regard the resemblance of the
species of Lema, together with that of the Longicorn Ozy-
lymma gibbicollis, for a species of Diabrotica, as an example of
Batesian mimicry. Future observation and experiment must
decide upon this as upon so many other cases concerning which
we are uncertain whether to adopt the Batesian or the Millerian
interpretation. The tendency of recent observation, however,
strongly favours the opinion that the latter theory will explain
a much larger number of resemblances than the former.

LINN. JOURN.-—ZOOLOGY, VOL. XXV2. 41

574 PROF. E. B. POULTON: NATURAL SELECTION

But whichever interpretation be ultimately adopted, the fact
remains the same—that the resemblances in the Coleoptera are
of the same character as those in the Lepidoptera, and are, like
the latter, independent of affinity. They are readily to be
explained by the operation of a theory of selection, but present
the same difficulties as those presented by the Lepidoptera to an
interpretation by any other theory as yet brought forward.

(6) The Resemblances in question are not accompanied by any
changes in the direction of another Species except such as
assist in the production or strengthening of « Superficial
Likeness.

This argument is fatal to any theory as yet advanced except
one based upon the principle of selection directed to a definite
end, viz. the production of resemblance. It is impossible to
explain why external forces or internal forces should thus act
upon a certain set of characters whose only relationship is that
they tend to produce a superficial likeness to another species—
that they should act upon these alone to the exclusion of all
other sets. No assistance can be obtained’from the conclusion
that the results are recent and therefore superficial, and that a
resemblance in deeper characters will follow in time. In the
first place, the examples of more perfect (and presumably older)
resemblance show no more tendency towards approximation in
characters which do not help to produce likeness, than the
examples in which the resemblance is comparatively rude (and
presumably recent in origin). In the second place, deep-seated
parts of the organism are affected when the superficial resemblance
is thereby increased, but not otherwise. To take a single
example, the common British Longicorn Clytus arietis strongly
suggests the appearance of a wasp, partly because of its black and
yellow banding, but even more because of its alert and wasp-like
movements. ‘This implies, of course, appropriate changes in its
nervous and muscular systems.

There are many cases, like that of Clytws,in which the changes
in deep-seated structures are of more importance than any-
thing else in determining the resemblance. I know of no more
striking example of this than the movements and attitudes of the
young (Lepidopterous) larve of Endromis versicolor, which
render them extremely like the larvee of saw-flies (Phytophagous
Hymenoptera). Numerous experiments have convinced me that

THE CAUSE OF MIMETIC RESEMBLANCE, 575

the latter are almost invariably distasteful. I exhibited the
former larve in the second stage at the Eutomological Society on
June 3, 1891. The following account is given in the Proc. Ent.
Soc. 1891, p. xv:— At this period [viz. the early stages of
growth] the larve arrange themselves in small groups upon the
leaves and leaf-stalks of the birch, and when disturbed they
raise the anterior part, bending the head over the dorsal surface
of the posterior part of the body. In this attitude they strongly
remind the observer of those Tenthredo larve which, when
irritated, bend the tail forwards over the anterior part of the
body. The fact that the head is raised in the one, and the tail
in the other, does not cause any conspicuous difference when the
larvee are seen from a little distance. The common Tenthredo
larva, Crasus septentrionalis, is about the same size as these
small Lepidopterous larvee, feeds in similar small groups when
large (when small the groups contain far more individuals), and
also frequents the birch.’ In my experience, however, the
Cresus feeds much later in the year. Mr. W. Hollaud also
noted the same resemblance in the ‘ Entomologist’s Record’
for Oct. 15th, 1891 (see vol. ii. p. 228). Mr. Holland has also
kindly lent me his notes made at the time, and I see that he
observed the saw-fly-larva-like movements which follow dis-
turbance. The groups have never been figured before, so far ag
I am aware. In May 1896 we reared some larve from the
egg in the Hope Department, and I was able to get some
excellent coloured drawings by Mr. P. J. Bayzand. These
drawings, reproduced on Plate 40. figs. 2 & 3, show the attitude
taken up on disturbance better than any description. They also
prove that, like the saw-fly larvee, the groups contain far more
individuals in the younger stages than later. The conspicuous
orange-coloured true legs suggest the appearance of the orange
ventral glands of the Crasus, which are everted when the larva
is irritated.

Thus the causes of the resemblance we are discussing may be
deep-seated or may be superficial, or, more generally, may be due
to several kinds of causes in each category. It is in the latter
extremely complex cases, and these are far the commonest, that
the argument for natural selection becomes irresistible. This
will be more thoroughly dealt with in the succeeding Section ;
but even in the case of the simplest element in the resemblance,

viz. the similarity in colour and pattern taken alone, the theories
41*

576 PROF. E. B. POULTON: NATURAL SELECTION

of external and internal causes are unable to offer an adequate
explanation of certain facts which are clearly explicable by
natural selection. In the males of the Pierine group Dismor-
whina, the long-and-narrow-winged appearance of an Ithomiine
butterfly is largely produced by the excessive overlap of the
upper upon the under wings. This results in the concealment of
a large part of the upperside of the under and of the underside
of the upper wing; and it will be found that the mimetic
patterns are withheld from these hidden surfaces, which often
retain some distinct trace of the old Pierine character, viz. an
opaque white appearance.

The male of Dismorphia praxinoe is shown on Plate 40. fig. 4,
the underside in fig. 5; the upper and under sides of the female,
for comparison, in figs. 6 and 7. This fact holds true even for
such a specialized and perfect mimic as Dismorphia orise (see
Plate 40. figs. 8 & 9, for the appearance of the male).

F. D. Godman and O. Salvin, in the ‘ Biologia Centrali- A meri-
cana’ (Rhopalocera, p. 173), speak of these hidden chalky
patches, surrounded by a silky area covered by peculiar scales,
as a character of the Dismorphina. The interpretation of the
patch as a sexual brand perhaps with the nature of a scent-
producing organ, does not in any way disprove the suggestion
here adopted—that the white pigment in the scales is a survival
from an ancestral condition still found over the greater part of
the wing-surface in so many non-mimetic Pierine, as well as in
the males of many mimetic species (e.g. in the genus Mylothris).
It should be noted, however, that the patch is not white in
certain species of the Dismorphina. Mr. Belt (‘ Naturalist in
Nicaragua,’ pp. 384, 385) states that the white patch is usually
concealed by the males, as indeed may be inferred from the
change in character of the surface, which indicates the normal
amount of overlap of the fore upon the hind wing. Mr. Belt
also suggested that the white appearance is ancestral and has
been retained by the operation of sexual selection.

The restriction of the effects to those parts of the surface
which can be seen tells very strongly against any theory which
is not based on the principle of selection.

(7) Essential Nature of these Resemblances: their Analysis
into the several kinds of Effect produced.
The resemblances under discussion are made up of elements

THE CAUSE OF MIMETIC RESEMBLANCE, Sha

of very different kinds combined in single individuals; but the
essentially composite nature of the effect easily yields to analysis.
Some of these complex combinations only require to be stated in
order to show the inadequacy of the theory which is most usually
substituted for natural selection, viz. the theory of external
causes.

A mimetic appearance is commonly made up of (1) colour,
including (a) structural as well as (0) pigment colours; (2)
pattern ; (3) form; (4) attitude; (5) movement.

It may be plausible to hold that direct local influences deter-
mine colour, but the case becomes much more difficult when
structural tints are included, as they frequently are. Thus it
might well be held that the dark pigment of a female Hypo-
limnas and of the Luplea which it resembles are alike the
direct effect of the locality they both inhabit. But the most
convinced advocate of direct local causes would probably hesitate
to explain, by the operation of the same forces, the structurally
caused blue sheen which overspreads the dark pigment in some of
these mimetic pairs. Similarly with pattern, it is much moie
difficult to understand the appropriate arrangement of the
colours by direct forces than the production of the tints them-
selves; still more difficult to uuderstand how such forces could
modity shape, and again, more difficult to mcdify the nervous
and muscular systems so as to produce appropriate attitudes and
movements. Most difficult of all to understand, except on a
theory of selection, how the several elements in the complex set
of changes could be kept in their proper relationship and guided
to a definite end, viz., the production of a superficial resemblance
to another species.

The objection to the theory of internal causes is not that it is
inadequate to produce each of the effects, but that it is in the
highest degree improbable that so complete and barmonious an
effect could be frequently produced accidentally by the com-
bination of such diverse elements.

When, therefore, it is argued that these resemblances are the
uniform result of uniform forces peculiar to the locality, investi-
gation proves that the results are very far from uniform. They
appear at almost any point in the structure of the body, super-
ficial or deep-seated, generally at many points in a single
individual both superficial and deep-seated, and the only common
bond which can be established between the various elements

578 PROF. E. B. POULTON: NATURAL SELECTION

which make up the common effect is that they all co-operate in
producing superficial resemblance to some other species.

It is here shown that the changes wrought in a single species
are far from uniform. It will be shown later on (see Sections
11 and 12, pages 585 to 602) that there is frequently no uniformity
in the methods made use of by mimic and model, nor any
uniformity between the various mimics of the same model, nor
between the different members of a synaposematic group. These,
too, often have only one thing in common, and that inexplicable
except on a theory of selection, viz. the subordination of all
these divergent methods to a single end—the attainment of
a superficial resemblance.

The arguments in this and the preceding Section are equally
powerful in support of the interpretation of protective resem-
blanees as due to natural selection.

Again, mimetic resemblances are comparatively rarely seen
in more than one stage of insect life, and are, in the great
majority of cases, restricted to the final stage. In all such
species the total appearances presented by the final stage, includ-
ing mimetic resemblances, are prepared for in the earlier stages,
and especially the larval. Not only are the changes in question
confined, as has been pointed out in the earlier parts of this
Section, to the parts, tissues, and organs which affect the super-
ficial appearance, but they are also generally confined to the
final stage of insect life. During larval life the foods peculiar
to the locality are devoured and the material for the mimetic
stage is stored up. The larval and pupal stages are together, in
the great majority of cases, far longer than the imaginal stage,
and are no less, and, as regards food, far more, subject to the
direct action or the forces peculiar to the locality. On what
theory except natural selection is it possible to explain the rigid
limitation of these changes, in so large a proportion of cases,
to the final stage, and their entire exclusion from the stages in
which they are, in the history of the individual, predetermined ?

(8) Conditions of a Species in any Locality are chiefly
determined by its Habits and Life-history.

In the last Section it was shown that there is no uniformity
in the effects produced in any locality. In this Section it will be
made clear that there is no uniformity in the forces which, by

THE CAUSE OF MIMETIC RESEMBLANCE. 579

their uniformity, are supposed to produce the effects. When we
are told that common food, common climate, &c. produce a
common effect, we have the means for proof or disproof in, at
any rate, some striking examples ; for we know the food and con-
ditions of certain species which exhibit mimetic or common
warning associations. There are many examples of Longicorn
beetles mimicking Lycide (Malacoderm beetles) in the same
locality ; but during the earlier stages, im which the appearance
of the final stage is determined, the former lives in a burrow,
feeding upon wood or the tissue of plant-stems, and sheltered
from many of the climatic influences and changes, while the
other lives in the open, freely exposed to them all, and sustained
by an exclusively carnivorous diet. I owe this suggestive com-
parison and the Section which arose out of it to a conversation
with Mr. C. J. Gahan, of the British Museum. Similarly in the
case of S. American moths belonging to the Castniide, which
resemble Ithomiine butterflies (see Section 12, page 598), the
larve of the former burrow in plants, while the latter are freely
exposed on the leaves which form their food.

It is hardly necessary to insist on the importance of the larval
stages in this respect. When the imago emerges from the pupa
and its expanded wings have dried, it has assumed its permanent
appearance, and nothing that it will eat or endure henceforward
produces any further effect upon its colours or patterns, &c.
Henee identity of food and conditions during the final stage cannot
be of any assistance in the interpretation of mimicry. It is neces-
sary to point this out clearly, inasmuch as Beddard (‘ Animal
Coloration,’ London, 1892, p. 232, footnote) has said, speaking of
the resemblance between Fvistalis, the drone-fly, and the hive-
bee, “‘ It is an interesting fact, in connexion with the resemblance
between this fly and a hive-bee, that it feeds upon pollen and
honey. This fact may have some significance in relation to the
effects of food upon form and coloration.’ But the larva of
Eristalis stores up nutriment, out of which the final form is
built, by feeding on putrefying animal matter, a food as different
as possible from that provided for the larval bee. The peculiar
conditions under which the larve of stinging Hymenoptera
obtain their food invariably contrast strongly with the larval
condition of their numerous mimics. We find in this Section,
as in the others, that the suggested interpretation of these
resemblances as the common effect of a common cause or set of

580 PROF. E. B. POULTON: NATURAL SELECTION

causes breaks down the moment it is analysed. The view is
a superficial one, and cannot be sustained when the slightest
attempt is made to understand the nature of the phenomena it
professes to explain.

(9) Mimetic Resemblance and Common Warning Colours more
characteristic of the Female than the Male Sex.

These resemblances are far commoner in females than males,
and when the two sexes differ in the closeness with which a
likeness to some other form is brought about, it is the female
which always attains the greater perfection. Examples of
mimetic females with non-mimetic males are extremely abundant,
being in fact a high proportion of all the cases which occur ;
examples of the converse relationship are very nearly unknown.
These general statements hold with common warning colours as
well as with truly mimetic species; they are equally true in all
the warmer parts of the world where examples of mimicry are
well known and abundant.

In the numberless cases in which a non-mimetic male is
attended by a mimetic female, the former bears the ancestral
appearance, so that when we pass to a land where both sexes of
the representative species are non-mimetic, both resemble the
non-mimetic male of the former species. In a long series of
related species, moreover, the males are found to be nearly alike,
while the females diverge in all directions after the species
which serve them for models. Furthermore, the females, by
reversion, are in rare instances brought back towards the ances-
tral type represented by the males.

It is hardly necessary to pomt to examples, for these general
principles will probably be at once conceded by any who have
made a study of the subject. I may, however, allude to the
non-mimetic Papilio meriones of Madagascar and the related
forms with similar males but widely different mimetic females
on various parts of the mainland of Africa; to the general
resemblance between the males of so many forms of Hypolimnas
of the bolina group to each other and to those of H. misippus,
&c.; to the varying degrees of reversion towards the appearance
of the male presented by occasional females of Hypolimnas
bolina.

These relationships are the reverse of those which usually

THE CAUSE OF MIMETIC RESEMBLANCE. 581

obtain. Outside these resemblances it is the rule, when any
difference between the sexes exists, for the female to show us
the ancestral type, the male the more modern development ; and
the male in growth from youth to maturity generally passes
through the condition permanently retained by the female.

No probable interpretation of these unusual relationships ean
be offered by any theory except natural selection. The theory
of external causes demands the improbable hypothesis, for which
no evidence can be found, that the female of mimetic species
(but not of others) is constitutionally more ready to respond to
the direct action of external forces than the male, and that the
difference is commonly great enough for the female to have
given a complete and detailed response, when the male, subject
to the same direct forces, does not exhibit the faintest trace of
the operation of any such influence.

The facts are equally inexplicable by the theory of internal
causes—and not inexplicable only, but the reverse of what we
should expect; for, as I have already stated, it is the female
which, outside these resemblances, tends to retain the ancestral
form.

The theory of sexual selection also fails to account for the
facts. If it were valid, the selection would be that of the male,
for these recent developments are specially characteristic of the
other sex. In other cases in which male rather than female
selection is supposed to have acted in the production of colour
or pattern in butterflies, there is some direct evidence derived
from the observation of courtship; but here no such support is
forthcoming.

Under the theory of natural selection the facts at once receive
an explanation. Wallace suggested long ago that the slower
flight of the females “ when laden with eggs, and their exposure
to attack while in the act of depositing their eggs upon the
leaves, render it specially advantageous for them to have some
additional protection.” In animals which are hidden by
protective resemblance, similar causes explain why the female
is so often better concealed than the male. In birds the dangers
of incubation balance the dangers of egg-laying in insects.
But protective resemblances are less special than cases of
mimicry in the sense that the models (bark, twigs, leaves, &c.)
are more generally alike throughout all countries, and less
rapidly change their distribution than the models of mimicry and

582 PROF. E. B. POULTON : NATURAL SELECTION

the dominant types of common warning colours. These and
other reasons, such as the great number and wide geographical
range of species belonging to the same genus and adopting
a single method of concealment, compel the belief that examples
of protective resemblance are extremely ancient in the past
history of the species as compared with examples of mimicry, so
that we ean well understand how it is that in the former, when
the female differs it is ancestral as compared with its male, while
in the latter the converse relationship obtains, and the appear-
ance presented by the male is comparatively ancestral.

The main conclusion which emerges is that the advantageous is
the thing that is attained. If an ancestral appearance is advan-
tageous it is retained, especially in the sex that needs it most ;
if a new appearance is advantageous it is attained, especially by
the sex that needs it most. The female sex becomes conserva-
tive or progressive according to the needs of the species, and
natural selection is limited by no bounds of constitutional dif-
ference between the sexes as regards the preservation of the old
or the initiation of the new.

(10) Lhe Space and Time Relationships of the Resemblances
in Question.

A mimetic group is found in the same locality, or at least the
mimic (Batesian) is not found beyond the range of its model.
The types of Common Warning Colours are remarkably local,
although probabiy certain members of the group (being ex
hypothesi all specially protected) may sometimes have a wider
range than others. Whensuch a mimic as Hypolimnas misippus
can invade and thrive in South America and the Antilles in the
absence of its model (Limnas chrysippus), we probably have to do
with a Miillerian rather than a Batesian association (see also
a paper on Mimicry in the Genus Hypolimnas by E. B. Poulton
in Report Amer. Assoc. for Adv. of Sci., Detroit Meeting, 1897,
where other arguments in support of this conclusion are urged).

Looking at the examples broadly the phenomena are charac-
teristically local. This, although harmonizing with the other
suggested explanations, is quite unintelligible if the theory of
internal causes be adopted. Why should these results if
attained independently in the evolution of various forms be
attained in the same locality P The number of patterns and the
number of forms is so vast that we must expect a certain amount

TILE CAUSE OF MIMETIC RESEMBLANCE. 583

of accidental resemblance due to internal causes, as has been
suggested by Beddard (‘Animal Coloration, London 1892, p. 252);
but such resemblances will differ from those under discussion in
this among other things—that they will not be characteristically
loeal. The theory of internal causes offers us a valid interpretation
of such cases, which are, as a rule, readily distinguished from those
which are here considered.

There is another aspect of locality which only receives an
explanation on the theory of natural selection. Why should
examples of mimicry and common warning colours be so much
more abundant and perfect in one country than another? The
physico-chemical influences, the effects of luxuriant vegetation,
as Wallace has pointed out, are very similar in_ tropical
S. America, Malaya, and W. Africa, and yet the first-named
country is pre-eminent in affording examples of the resemblances
under consideration. This is not only true of likenesses within
the Order of Lepidoptera, it appears to be equally true within the
Coleoptera; it is true of the resemblances of moths to wasps.
It is even more marked in Mullerian resemblance between pro-
tected forms than in the examples of Batesian mimicry. If the
direct action of forces connected with locality cannot explain the
immense predominance of tropical South America in this respect,
we are driven to enquire whether insect-life is especially luxuriant
and remarkable in this part of the world, and whether it 3s not
probable that the struggle for existence is especially keen.
There is no doubt about the answer to the former question ;
the variety, peculiarity, and abundance of insect life is far greater
than in any other part of the world, and it is a fair inference
that the conditions are in an equally marked degree favourable
for rapid and complete modifications under the operation of
natural selection.

We have not as yet sufficient evidence that mimetic groups
and groups with a common warning coloration appear at the
same time of the year. Such evidence as we have points in this
direction. The rarity of the mimetic species is usually stated to
be due to their being lost in the swarms of the abundant model.
There are a large number of cases in which the forms have been
caught together by a collector who has passed a limited time in
a given locality.

We are collecting at Oxford as many examples as possible of
such species, captured upon the same day in the same place.

584 PROF. E. B. POULTON : NATURAL SELECTION

This series, which we intend to keep separate, when more com-
plete, will afford very valuable evidence on this point. Already
it proves that the members of the groups which converge round
Limnas chrysippus, Amauris echeria, and the black and white
species of Amauris in Natal are upon the wing together. This
evidence has been very kindly supplied me by Mr. Guy A. K.
Marshall. I have similar, but less complete, evidence as regards
some of the Central and 8. American groups.

It will probably be conceded that the phenomena generally
are likely to exhibit the same relationship in time which has
been already proved to exist for many of them. This conclusion,
however, is a considerable difficulty in the way of the theory of
external causes as well as a further difficulty to the theory
of internal causes. As regards the latter, the time relationship
is an entirely unexplained coincidence ; as regards the former, it
is a coincidence which leaves much to be explained. It is diffi-
cult enough to believe that local forces could produce local
resemblance; it is a further difficulty that the resemblances are
contemporaneous. If, as is probable, the forces are supposed to
act during larval life, they must include in their effects an
influence on the rate of growth and development, an adjustment of
the duration of stages delicate enough to bring the various species
into the phases in which the resemblance is shown at similar
times of the year. But such effects are entirely different from
those which are manifest in the resemblance itself, and add
a further complexity to a result already shown to be so complex
that the theory of external causes fails to supply an interpreta-
tion (see Section 7, p. 576).

But it has been shown in many cases, and is probably true in
all, that the time relationships between the species which exhibit
these resemblances are not confined to their appearance at the
same season of the year. They are such that they fly together
under those conditions of light which render the resemblance
visible to enemies. When moths resemble butterflies, they are
mostly species which are as truly day-flying as the butterflies
themselves; in other cases they are species which fly readily by
day when disturbed. Similarly with the species of various orders
which resemble Hymenoptera. The case of Coleoptera recently
suggested to me by Mr. Gahan is peculiarly interesting. It is
known in so many cases that beetles which are about by day
possess finely facetted eyes as compared with the larger fewer

THE CAUSE OF MIMETIC RESEMBLANCE. 585

facets of the nocturnal species, that it is possible to infer the
habits from the structure of the eyes. Thus the species of the
Longicorn genus Doliops (family Lamiide), which closely resemble
weevils (see p. 596), are, judged by this standard, diurnal in their
habits. The case is all the more interesting, inasmuch as such
an eye-structure, such habits, and such mimicry is quite ex-
ceptional in the group, the Miphonine, to which the genus
belongs.

The facts recorded above imply such a resemblance between
the nervous systems and sense-organs of the various species as
will make them diurnal (or in some cases semi-diurnal) in their
habits. This constitutes a further grave difficulty in the way of
any explanation based on external or internal causes. If the
diurnal habits are supposed to be due to such causes, the vastly
increased complexity of the result is the difficulty. If the
resemblances are supposed to be thus produced only in the
species which are already diurnal, it is impossible to explain why
the external or internal forces are thus restricted in their
operation.

It is hardly necessary to point out that the time and space
relationships, which are such a difficulty in the way of the other
two theories, are entirely necessary to the explanation based on
the theory of natural selection. If they did not exist it would
be overthrown.

(11) The Resemblances which Insects of various Orders bear to
those of another Order are produced in the most Diverse Ways.

The most common types for mimetic resemblance are those of
the wasp and ant. These aggressive, abundant, and successful
forms are resembled by insects of various orders. Still more
interesting is the fact that the resemblance is produced in the
most varied ways.

A superficial resemblance to stinging Hymenoptera is probably
more general and is brought about by smaller changes in Diptera
than in any other Order of insects. A fly which gains alternate
black and yellow rings on its body is at once suggestive of an
appearance presented by many common wasps. In more extreme
cases, the body gains a constriction presenting a strong likeness to
the slender petiole of the wasp’s abdomen, there are changes inthe
manner of folding and sometimes in the colour of the wings, in

586 PROF. E. B. POULTON: NATURAL SELECTION

the buzz, in the movements of the body (the latter being such as
to suggest the power of stinging), and (in the mimicry of hairy
humble-bees) in the acquisition of an abundant hairy covering.
A good example of a fairly perfect resemblance is shown in
Plate 41. fig. 5a, an Australian fly, the models from the same
part of the world being shown in figs. 5 B and 5.

A Hemipterous insect requires the most profound modification
in the shape of its flattened un-wasp-like body, and in the dis-
play and characteristics of its wings. Corresponding to these
much greater initial differences, the resemblance is much rarer
than in Diptera. A beautiful instance is shown in Plate 41.
fig. 1, a Hemipterous insect—a species of A/yocoris, probably
M. braconiformis (Burm.)—which is extraordinarily like fig. 2,
not, in this instance, a wasp but an ichneumon, using the term
in the broad sense ; for the species belongs to the Braconide and
is probably an Jphiaulax. Although the two specimens in the
British Museum, which I have been kindly permitted to figure,
did not come from the same part of S. America (the Bug was from
Para, the Ichneumon from Ega), the resemblance is so close in
every way and the modification of the Hemipteron so extreme
and remarkable, that there can be little doubt as to this being
a genuine case of mimicry and as to the geographical coincidence
of the model and mimic: both may have a far wider range than
is shown by the specimens which have been figured *.

A Lepidopterous insect requires above all to gain trans-
parent wings, and this in the most striking cases that have been
studied is produced by the loose attachment of the scales, so that

* Since the above sentence was written I have searched the collections in the
British Museum for further examples, with the help of Sir George Hampson,
Mr. C. J. Gahan, and Mr. W. F. Kirby. We found that the type of appear-
ance represented in Plate 41. figs. 1 and 2 iscommon and widely distributed in
S. America, forming a group which includes many species of Lraconide and of
Hemiptera which resemble them. Many figures of such insects presenting
this type of appearance are to be seen in the plates of the ‘ Biologia Centrali-
Americana.’ In the distasteful group of moths, the Syntomide, two species are
obviously similar, viz., Leucotmemis varipes (Walk.) from Para, and L. tenthre-
doides (Walk.) from Kga. Myocoris braconiformis is recorded from Surinam as
well as from Para.

The co-existence of species of Aculeate Hymenoptera, Hemiptera, and
distasteful Heterocera, suggests that the group as a whole is distasteful or
specially protected, and is an example cf common warning colours. Many
other members of the group will probably be recognized

THE CAUSE OF MIMETIC RESEMBLANCE. 587

they easily and rapidly fall off and leave the wing bare except for
a marginal line and along the veins (Hemaris, Trochilium). In
other cases again (certain Sestide) the scales may remain on the
wing, but themselves become transparent. In the numerous
more perfect instances the body is banded, and may gain a
marked “ waist,” while the scales upon it may be lost or moditied,
so that the appearance of the hard shining body of the model is
suggested with extraordinary exactness.

The means adopted among Coleoptera, even in closely related
genera, are so curiously different that a longer description,
accompanied by several illustrations, is necessary.

The following examples are all selected from the Longicorns.
The simplest resemblance to a wasp is that attained by the
common beetles of the genus Clytus, such as the British C. arietis.
A better example is the Mediterranean species Plagionotus
scalaris, shown on Plate 41. fig. 3. In these cases there is
nothing visible to represent the wings of a wasp; but the elytra
and thorax are black banded with yellow, there is a far more
pronounced ‘“‘ waist” than is usual in Coleoptera, the legs are
slender and wasp-like, and are moved with wasp-like activity and
jerkiness. In spite of the apparent want of wings, the effect
produced is very considerable, and many persons hesitate to
touch the insect. This, then, is the method adopted in the group
of Clytine ; but in various other allied tribes, such as the Necy-
daline, the Rhinotragine, the Esthesine, the Callichromine,
and others, the elytra, which form by far the largest part of the
visible dorsal surface in the Clytine, become greatly reduced so
as to show the under wings, which, folded over the back or
expanded in flight, in either case strongly suggest the wings of
a wasp, or in some cases anichneumon. Furthermore, the elytra
are reduced in two different ways—in some genera to linear
rudiments more or less broadened at their bases; in others to
small subquadrate or oval structures representing the bases
alone. Good examples of the latter are seen in Esthesis ferru-
gineus (Hsthesing) (Plate 41. fig. 5) from Australia, and in
Nothopeus hemipterus (Callichromine) from the Oriental Region
(see fig. 4). Examples of the former are shown in Isthmiade
braconotdes (Rhinotragine), fig.6, from Brazil, and in a Hephes-
tion sp. (Necydaline), fig. 7, from Chili.

The examples figured are merely a few selected from a large
number, but they serve to show the different degrees to which a

588 PROF. E. B. POULTON : NATURAL SELECTION

mimetic resemblance is carried, and the very different methods
employed, in a nearly related set of insects.

The Hymenoptera, which form the models of these insects,
have not been worked out in many of the instances. In the case
of Esthesis ferrugineus, however, two Aculeate Hymenoptera
of very different form, although belonging to the same large
group—the Humenide, which inhabit the same region (Australia),
have a colourmg and pattern which are superficially very
similar to those which the Longicorn has attained; while a
Dipterous insect, an Australian species of Dasypogon (Asilide),
also appears to fall into the group. The appearance of the
three latter insects is represented in Plate 41, the dipterous insect
being shown in fig. 5a, the wasps in fig. 5 B, Abispa australis
(Smith), and fig. 5 c, Humenes Latreillei (de Sauss.). These three
figures were drawn from specimens in the British Museum, kind
permission having been accorded 1o me to figure them for the
purpose of this memoir. Future study will probably add many
species to this very characteristic group.

We thus find that wasps and allied forms are resembled by
species of many groups of insects, and the resemblance is attained
in all kinds of different ways.

The numerous mimetic resemblances to the aggressive, abundant,
and well-defended ants supply an even better illustration. In
the majority of cases the whole body of the mimetic form is
moulded from the ancestral shape, which is still exhibited by
its non-mimetie allies, into that which is characteristic of an ant.
In some groups this means a large amount of alteration, in
others less. In this case, too, the resemblance extends to species
which are altogether outside the Insecta ; for many small species
of spiders closely mimic ants. In the family of Attide alone,
and such resemblances occur in several other families of spiders,
George W. and Elizabeth G. Peckham state (Occasional Papers
of the Natural History Society of Wisconsin, vol. 11., 1892,
Milwaukee; see also a paper by the same authors in vol. i.,
1889) that about a hundred ant-like species are known from
various parts of the world, and that they are ‘‘ very much more
numerous in South America and in the Malay Archipelago than
in any other countries,” viz. in the very countries in which other
examples of mimicry are especially abundant. The spider with
its two-fold division of body is often made to assume the appear-
ance of an ant, with its three-fold division, by a constriction which

THE CAUSE OF MIMETIC RESEMBLANCE. 589

sometimes crosses the cephalothorax, sometimes the abdomen.
The absence of antenne in the spider is known to be compen-
sated in some of the species, which have been studied in the
living state, by the habit of holding up one pair of legs, while the
walking legs are thus reduced to the ant-like number of six. Of
two well-known North American species, Synageles picata holds
up the second pair, and Synemosyna formica the first. The habits
of seizing and dealing with prey, and the movements generally
are extremely un-spider-like and most suggestive of ants; so
that the nervous and muscular systems, as well as the body-
form, have been modified. The remarkably ant-like appear-
ance of these two species is shown in the adjoining fig. 1 (A

and B).

Fig. 2.

Fig. 1.—Two North-American Attid spiders which resemble ants. A is Synageles
picata; B, Synemosyna formica. (From G. W. and E. G. Peckham,
Occasional Papers of the Nat. Hist. Soc. of Wisconsin, vol. i. 1889,
pp. 110 & 112.)

Fig. 2 (x 3).—The young larva of Stauropus fagi seen from above and from
the left side.

Among the Insecta, too, there are many examples of an ant-like
appearance brought about by changes of the same kind as those
mentioned above, although less marked because the forms
to be approximated are less essentially different. Among the
Lepidoptera the young larve of a British moth, Stawropus fagi,

ave often been described as resembling ants. The likeness has
recently been analysed in much detail by Portschinski (‘‘ Colo-
ration marquante et T'aches ocellées,” V.: St. Petersburg, 1897,
p- 44). This acute observer considers that the head of the
LINN. JOURN.—ZOOLOGY, VOL. XXVI. 42

590 PROF. E. B. POULTON : NATURAL SELECTION

larva represents the globular abdomen of the ant, while the head
and antenne of the latter are suggested by the larval caudal shield
with its two appendages. He believes that the disturbed larva
represents an ant which has seized and is endeavouring to carry
off some object on the branch which it is exploring. Under
these circumstances the head of the ant, with its mandibles fixed
in the object, would be held low and remain motionless, while the
abdomen would be elevated and the legs in constant activity,
moving the posterior part of the body from side to side. Such
an appearance and such movements, he maintains, are strongly
suggested by the disturbed larva if we only identify the posterior
end of the one with the anterior end of the other, and vice versa.
I have to thank Mr. W. R. Morfill for very kindly translating
the memoir of the Russian naturalist. During the present
summer (of 1898) I have had the opportunity of studying these
larve. The young larve were thought to be ants by all the
friends to whom they were shown. One lady considered that
they were “double ants’””—an interpretation evidently due to
their disproportionate length and to the head-like appearance of
the caudal shield. Drawings of the larve at this stage were
made by Mr. P. J. Bayzand and are reproduced in fig. 2 (p. 589),
but they fail to convey the ant-like appearance which depends so
largely on movement. A better effect is produced by Mr. Bay-
zand’s coloured drawing which is reproduced on Plate 40. fig. 1.
I should add that I did not observe any attitudes which
support the details of Portschinski’s interpretation, nor did I
witness the appearances which he figures (/. ¢. p. 45, fig. 21).
His comparison of the caudal appendage with a head seemed, on
the other band, to be entirely confirmed.

Turning to other Orders which supply examples of the
mimicry of ants, the Hemiptera have perhaps the farthest
distance to travel in the modification of their flattened bodies.
A beautiful example from East Africa, viz. that of Wyrmoplasta
myra (Gerstaecker), is shown in fig. 38. Gerstaecker states
that a single specimen of this insect was sent from Rosako,
Usaramo, Aug. 1888, as “an ant,” together with two undoubted
species of these Hymenoptera (Polyrhachis gagates and Ponera
tarsata). ‘The resemblance between the former species of ant
aud the Hemipteron, Gerstaecker describes as strong enough
to be deceptive. It is brought about, he states, by the short
globular abdomen extremely constricted towards the thoiax

THE CAUSE OF MIMETIC RESEMBLANCE. 591

Fig. 3 (Xx 3).—An ant-like East-African Hemipterous insect, Myrmoplasta myra
(Gerst.), seen from above and from the left side. (From Gerstaecker,
Article 6, Hemiptera, p. 9, in Fr. Stuhlmann’s Zool, Ergeb. 1888-
1890, Berlin 1893.)

Fig. 4.—An ant-like N.-American beetle, Huderces picipes (Fab.), seen from
above and from the right side.

(Zool. Ergeb. einer Reise in Ost-Afrika, Fr. Stuhlmann, Bd. I.
no. ix. 2; Article 6, Hemiptera, p. 9: Dietrich Reimer, Berlin,
1893).

Among Coleoptera the resemblance to ants is very common. I
select as an example a little Longicorn (Huderces picipes, Fab.),
which I found very abundantly upon the heads of Umbelliferous
plants at Pine Lake, Hartland, Wisconsin, in July and August
1897, when visiting Dr. C. A. Leuthstrom. Ants were also very
common on the same flower-heads. The appearance and move-
ments of the beetles were extremely ant-like, the suggestion of
a stalked abdomen being conveyed by an oblique white line
crossing the elytra in a very shallow depression in which the
dark ground-colour of the insect appeared to be of a more intense
black than elsewhere. The increased darkness was in reality

4.2*

592 PROF. £. B. POULTON: NATURAL SELECTION

due to the shadow in the depression combined with the effect of
a difference in the texture of the surface. This combination of
characters produced a strong superficial resemblance between
the elytra of the beetle and the abdomen and thorax of the ant,
while the head of the latter was represented by the beetle’s head
and thorax together. These resemblances are indicated in fig. 4 ;
but the living insect is required in order to recognize them
fully.

In all the cases alluded to above, the resemblance is attained
by a modification in the form of body and limbs, accompanied
by changes in those more deep-seated structures which affect
the habits and movements.

There are, however, other very different means by which the
same end is attained. One of the most interesting of these is the
case of a Locustid (Phaneropterides), Myrmecophana fallax from
the Sudan, described by Brunner von Wattenwyl (Verhandl.
der k.-k. zool.-botan. Ges. in Wien, 1883, p. 247). This insect
was erroneously described as an Acridian in my paper before
the British Association at Toronto (Report, 1897, p. 693, 1. 3).
Brunner’s two figures are reproduced as the adjacent fig. 5.
Upon the stout body of this insect the slender-waisted form of
an ant is represented in black pigment, the remainder of the
body being light in colour and probably invisible against a
similar background. Of the habits of the insect nothing is
known, but the method is of great interest, being so entirely
different from that adopted by any other insect as yet described.
In a more recent work (‘Observations on the Coloration of
Insects,’ English translation by E. J. Bles, Leipsic, 1897, p. 11),
Brunner von Wattenwy] again alludes to this example, and states
that the form of the species ‘“‘leads to the conclusion that it
lives on the ground,” viz. in the position which gives a meaning to
the resemblance. In spite ot this he asks, “Is this imitation an
accidental freak of nature?” If Myrmecophana were the ouly
example of such resemblance the question might fairly be asked,
but in view of the numerous other equally close resemblances to
ants, produced in various ways, it is quite unnecessary. The
suggestion of an “‘ accidental freak ” can never explain such close
likeness, in appearance, in movements and habits (so far as they
are known), in locality—a likeness, furthermore, not to any
insect in the environment, but to insects of a specially successful
and aggressive group—a likeness not produced in one way but

en

THE CAUSE OF MIMETIC RESEMBLANCE. 593

in many different ways. To suggest an “accidental freak” as
the explanation shows an amazing credulity, only to be explained
by the bias which is ready to accept any interpretation except
that afforded by the theory of natural selection.

Fig. 5.—-A Locustid from the Sudan, Myrmecophana fallax (Brunner), seen from
above and from the left side. On this insect the appearance of an ant
is represented in black pigment, all other parts being light in colour
and presumably concealed. (From Brunner von Wattenwyl, Verhandl.
d. k.-k. zool.-botan. Ges. in Wien, Bd. xxxiii, 1883, pl. xv. figs. 1a
& 1.)

Fig. 6.—A Central-American Membracid (Rhynchota Homoptera) in which the
prothoracic shield resembles an ant. Thus the body of the insect
which is not like an ant is concealed by an ant-like shield. The
species is Heferonotus trinodosus as seen from above and the left side.
The upper of the two figures seen from the side represents a female,
the two other figures, males. (From W. W. Fowler, Biol. Centr.-
Am., Rynch. Homopt. vol. ii. pl. 6. figs. 16, 16 a, & 17.)

Another and equally interesting method is adopted by certain
tropical American Rhynchota Homoptera belonging to the family
Membracide. In this remarkable group the dorsal region of the
first thoracic segment (the pronotum) is of enormous. size,
extending upwards and backwards so as completely to cover the
insect with the exception of the head, limbs, and wings. What
natural selection effects in the general body-form of other
insects, must here be effected, if it is to be of any value, in the
shield which is seen, and not in the body which is concealed.

594, PROF. E. B. POULTON : NATURAL SELECTION

This change has been brought about, and certain species of the
group have their un-ant-like bodies concealed under an ant-like
shield. In other species the prothoracie shield is modified into a
resemblance to other objects, such as seeds, thorns, &c.—an
excellent example of the parallelism between mimetic and pro-
tective resemblance insisted on on page 565. The effects pro-
duced in the shield are at least as exact and detailed as those
which in other cases are wrought in the form of the whole
body ; for, as Mr. W. F. H. Blandford has pointed out to me, a
peculiar characteristic of certain tropical American ants (viz. the
bead-like dilatation in the stalk of the abdomen) is reproduced
in the shield of the membracid. This is well seen in Heteronotus
trinodosus shown in fig. 6, copied from Canon W. W. Fowler’s
Monograph of the group in the ‘ Biologia Centrali-Americana.’
Finally, in the same group of Membracide we meet with another
example which is also incapable of interpretation by any theory
as yet brought forward except natural selection. An immature
form of membracid, with the prothoracic shield not yet formed,
found by W.1L. Sclater in British Guiana, strongly resembles
one of the leaf-carrying ants which are so common in that part
of the world; but the resemblance includes the leaf as well as

Fig. 7.

Fig. 7.—About three times the natural size. On the right is represented an
immature Membracid (Rhynchota Homoptera) from British Guiana,
which resembles an ant together with the leaf it is carrying. The
latter is shown on the left, and represents the species (codoma cepha-
lotes from the same locality. (From Poulton, Proc. Zool. Soe. 1891
pl. xxxvi. fig. 2.)

2

the ant! The dorsal region of the membracid is flat and
compressed, so that it is as thin as a leaf; its border (the dorsal
surface, which formed a sharp edge) is jagged like a gnawed leaf,
and during life it was green in colour. Beneath this leaf-like

— = -

a

THE CAUSE OF MIMETIC RESEMBLANCE. 595

expanse the brown head and legs of the insect were visible just as,
in the case of the ant, they appear beneath the piece of leaf which
is carried vertically between the mandibles and thrown over the
back. This example was deseribed by the present writer in Proc.
Zool. Soe. 1891, p. 462. The appearance of the membracid and
leaf-carrying ant is seen in fig. 7, which is reproduced from the
paper just mentioned. It is possible that certain species of the
Orthopterous genus TZettix (Acridiide) also resemble ants
carrying leaves.

It would of course be ridiculous to ascribe this last resem-
blance to any direct external forces connected with locality, or
to any internal forces, independently producing a like result,
and, as the species was in an immature condition, it is equally
impossible to invoke the aid of sexual selection.

Natural selection remains as the only feasible interpretation.

Even more striking than this remarkable example is the con-
templation of all these various methods and their relation to
each other. The means by which the resemblance to ants are
brought about are diverse, the end—the resemblance itself—is
uniform. Furthermore, the likeness is almost always detailed and
remarkable, however it is attained, while the methods made use
of differ absolutely. Such a result, it would seem, is the most
complete proof of the operation of natural selection that can be
attained, short of the actual demonstration of its action by
observation and statistics. If this argument be confirmed by a
study and comparison of the foregoing figs. 1 to 7, I venture to
think that it will meet with general acceptance.

When one insect resembles an ant by the superficial alteration
of its whole body-form, another by the modification of a shield-
like structure which conceals its unaltered body, another by
having the shape of an ant painted, as it were, in black pigment
upon its body while all other parts are concealed; another
by a further modification of its body, so that it represents not
an ant only, but the object which the ant is almost always
carryiug,—when the effect of all these results is heightened by
appropriate habits and movements, we are compelled to believe
that there is something advantageous in the resemblance to an
ant, and that natural selection has been at work. The phenomena
do not merely disprove all other suggested causes of change, but
they constitute the most powerful indirect proof of the operation
of natural selection.

596 PROF. E. B. POULTON: NATURAL SELECTION

(12) The Resemblances within the Limits of the Order are also
produced in the most Diverse Ways.

Illogical as I believe the position to be, it is quite possible that
many observers may concede the force of the argument con-
eluded above, and yet continue to hold that the resemblances
within the Order are produced by external or perhaps by
internal causes. It can however be shown that the same con-
dition is true in the more restricted group as was found to
apply to the wider. Even within the Order itself resemblances
are produced in very diverse ways, although minute examination
is sometimes necessary before the essential difference which
separates them can be revealed.

Certain Longicorn beetles resemble weevils, the Curculionide
being, so it is believed, specially detended by their extremely
hard chitinous covering. ‘The weevils closely resembled by such
Longicorns as Doliops curculionoides and D. geometrica have
short antenne ending in a knob. The antenne of the Doliops
are nearly three times as long; the resemblance to the weevil
being produced by a dilatation of the third joint, which repre-
sents the knob, while all the joimts beyond are of such excessive
fineness that they are almost invisible. The strong resemblance
of the Longicorn Estigmenida variabilis to Estigmena chinensis,
belonging to the Hispide (Phytophaga), is brought about im a
similar manner, as was pointed out by C. J. Gahan. In this
case about one third of the length of the Longicorn’s antenna
is concealed by its extreme fineness, while the apparent terminal
thickening is produced by hairs at the end of the thicker
section.

In these examples, kindly shown me by Mr. Gahan, neither
the theory of external nor that of internal causes is of any avail.
It is impossible to believe that the resemblance is a direct effect
of climatic or other forces connected with locality, when the
results are in reality so utterly different and yet superficially so
entirely alike. he fact which requires explanation is the extra-
ordinary likeness in spite of the essential difference, and this,
when it is repeated again and again, cannot be interpreted by
any theory unless based upon the principle of selection.

Many other examples of the same kind could easily be
brought forward: in fact, it may be admitted as a general
principle that in protective mimicry and common warning colours

THE CAUSE OF MIMETIC RESEMBLANCE. 597

the resemblance is never attained by precisely similar methods,
and generally by methods which are extremely unlike. I propose,
in concluding this Section, to discuss a few examples from the
Lepidoptera, inasmuch as the resemblances in question have
been chiefly studied in this group, and because an explanation
based on the theory of external or on that of internal causes
has been sought more often and pressed more strongly in the
case of the Lepidoptera than in that of any other Order.

The Pierine are specially lable to take on these resem-
blances. In tropical America they chiefly resemble the I¢tho-
miine, Heliconine, and Papilionine, affording some of the best
and earliest recorded examples of mimicry (although Dr. F. A.
Dixey has now shown that they are more probably to be inter-
preted as common warning colours). The chemical nature of
the wing-pigments of the Pierine has recently formed the
subject of an interesting paper by F. Gowland Hopkins (Proc.
Roy. Soc. 1894, p. 5, and Phil. Trans. 1895, B. p. 661). The
author shows that the white pigment so common in the group
is an impure uric acid probably uncombined, that the yellow and
orange pigment is a derivative of uric acid to which he gives the
name ‘‘lepidotic acid,” while a much rarer red pigment, less
fully investigated, is probably of a closely similar constitution.
These three pigments, with black, which is apparently intimately
associated with the cuticle of the scales, with a pigment placed
between the two lamine of the wing and with superadded
optical effects due to structure, account for the whole of the
colouring of Pierine butterflies. No pigment of similar consti-
tution to the Pierine white, yellow, and red, was found by
Mr. Hopkius in any other butterfly—not even in the allied
Papilionine. The Pierime butterflies which resemble the J¢tho-
miine or other butterilies were found by Mr. Hopkins to achieve
this end, not by gaining the true pigments of their models,
but by means of the characteristic Pierine pigments. The
bands of warm red-brown, the spots of white and yellow, which
so closely resemble the same tints in the IJthomiine, are in
reality caused by pigments of an entirely different nature—the
resemblance, even between the pigments themselves, is wholly
superficial.

The argument that the resemblances we are discussing are
due to the common result of common forces, is simply an
improbable assumption. It has been proved again and again

598 PROF. E. B. POULTON: NATURAL SELECTION

that the results are not common, the resemblance merely
deceptive. And now this has even been shown for the colours
themselves, in some of the best known and most striking
examples of mimicry.

The last example of similarity in appearance produced
through diversity of method is one which occurred to me a few
years ago when lecturing on mimicry in the Hope Department
at Oxford. It was worked out in detail during the summer of
1897, and the general results were communicated to Section D
of the British Association at Toronto, on Monday, August 28rd,
1897. A brief abstract is printed in the Report of the Meeting
(pp. 692-694).

Although the Lepidoptera are characterized as an Order by
the clothing of scales upon the wings, cases are very frequent
in which this covering has been in part, or almost wholly, lost.
By comparison with kindred unmodified forms, as well as by
microscopic examination of the transparent areas themselves, it
is possible to show that this loss is recent, and to trace the
steps by which it has been reached from a condition in which
normal scales were present.

In the very large convergent group of tropical American
Lepidoptera, which has sprung up round the best known species
of the Ithomtine genera Methona and Thyridia, transparency of
a large part of the wings is a characteristic feature. Through-
out the group the ground-colour of the wings is transparent,
with a black border which generally passes inwards as two trans-
verse bands in the fore wing, separating the transparent part
into three areas, and as one band in the hind wing, separating
it into two areas. The group consists of [thomiine (Neotropine)
of many genera, of Danaine, of Pierine, and of moths of the
genera Anthomyza and Hyelosia, belonging to the Pericopide
(Hypside), and the widely separated genus Castnia (Castniide).

Under the theory of external causes we should expect that the
transparency would be attained in a similar manner throughout,
by the reduction of scales to hairs, by the complete loss of scales,
or by some other uniform method. Under the theory of natural
selection we should expect that the methods would be different
in the different groups. There are many ways in which
transparency can be attained, and we should expect that one
group would submit one set of variations making towards the
resemblance, another a different set, to the operation of

THE CAUSE OF MIMETIC RESEMBLANCE. 599

natural selection. It will be shown below that this prediction
is abundantly justified.

Although H. W. Bates did not know all the species we
recognize now, he knew of all the chief sections of this group,
and described them (figuring certain of the species) in his paper
published in the Transactions of this Society for 1862. A repre-
sentation of the group as he describes it is shown in Pl. 42.
fig. 1; but he also mentioned another set of rather smaller insects
which is really continuous with the other group and should
be considered with it. This included the Ithomiine species
Dircenna epidero, D. dero, and D. rhoeo, and the moth Hyelosia
tiresia.

Owing to the kindness of Mr. Godman, Mr. Salvin, and.
Mr. Herbert Druce, I am able to figure the group as it is now
known with over thirty species (Pl. 42. fig. 2). The simple
combination of transparency and black which makes up the
whole of the appearance of these species renders an uncoloured
representation entirely sufficient. The strong general resem-
blance which runs through this large group is well seen in fig. 2,
which is taken from a beautiful photograph of the actual insects
made by Mr. Alfred Robinson, of the Oxford University Museur.
The arrangement of the insects for the camera upon small
pieces of cork glued on to a sheet of glass, was a work involving
great care and skill, and in this, as in so many other sides of my
work, L wish warmly to thank Mr. W. Holland, of the Hope
Department, for his most efficient help and ready kindness.

Microscopic examination of the Ithomiine (Neotropine)
showed that the scales on the wings are of two kinds, broad
and narrow, which alternate more or less regularly. In the
transparent parts both kinds of scale can still be detected, the
narrow being frequently reduced to fine simple hairs, and, in
the most extreme cases, the broad scales being reduced to
Y-shaped hairs. The two commonest species of the whole
group, probably forming the centre towards which the others
have converged, are shown in PI. 43. fig. 1. These species,
belonging to very different genera, afford a good example of the
closeness of resemblance which may be attained by common
warning colours. The two forms are constantly found intermixed
in collections, and superficially are almost exactly alike. Never-
theless, a study of the transparent part of the wing under the
microscope reveals the fact that the degeneration of the scales

600 PROF. E. B. POULTON: NATURAL SELECTION

has reached a very different level in the two species. In this,
as in all other instances, the transparent patch at the apex of the
left fore wing was selected for study and for photomicrography.
Dr. Gustav Mann, of the Oxford University Physiological
Laboratory, kindly assisted me in the difficult details of this
latter work, and such success as I have been able to attain I owe
mainly to him. Fig. 2 shows the structure of the transparent
patch in Methona confusa; fig. 3 its structure in Vhyridia
psidit. In both the broad scales are reduced to small bifid
structures, of a distinctly different form in the two species ;
while the narrow scales are reduced to long extremely fine hairs
in the Methona, and to much broader ones in the Thyridia.
In both the transition into the scales of the opaque part of the
wing is well seen on the upper part of the figures. The
comparison between the almost identical appearance of the two
insects as shown in fig. 1, and their real differences which are
seen to be very marked in figs. 2 and 3, affords another good
example of the principle which has been already so abundantly
illustrated—that when a close superficial resemblance has
been attained no further similarity in the details of structure
is produced. This is apparent enough even in these two genera
belonging to the same sub-family. It is of course illustrated in
a far more striking manner when the affinity is much more
remote.

We now pass to the Danaine which fall mto the group.
The Danaine supply it with two species, both belonging to
the genus J¢una, shown in Pl. 43. fig. 4. The Ithomiine were
formerly classed with the Danaine, and in any case form the
sub-family which stands next to them. But fig. 5 shows that
transparency is attained in an entirely different manner, the
scales being neither greatly reduced in size nor much altered
in appearance. It is the great reduction in numbers which
is here the cause of the transparency. The scales retain their
dark pigment and produce the effect of a grey dusting over
the transparent areas. The transition into the opaque black
part of the wing is well seen on the upper part of the figure.

Finally, there is a single species belonging to the distant
family Papilionide and sub-family Pierine—Dismorphia orise.
This species is shown in Pl. 44. fig. 1 together with dethona
confusa. The upper two figures of the Jethona and of the
Dismorphia show a male and female of the type form of each

THE CAUSE OF MIMETIC RESEMBLANCE. 601

species, while the lowest figure shows a form which is found in
Keuador. In certain parts of Ecuador the black borders and
bands of the Methona (recently distinguished as JZ. psamathe,
Godm. & Salv.) are much narrower than in the type form, and
it is seen that the Dismorphia from the same locality has
followed in the same direction. This interesting parallelism
was pointed out to me by Mr. Salvin, and is here illustrated
from lithographs of specimens in the Godman-Salvin Collection.
Fig. 2 shows that, in Dismorphia, the transparency is attained,
unlike the Jtwna, by a reduction in size of the scales; while,
unlike the Jthomiing, the normal shape and outline are pre-
served, almost unchanged.

Tt is a little remarkable that in this large and dominant group
no member of the Helicunine has yet been shown to find a
place.

The moths which join the group are shown in PI. 44. fig. 3.
I believe that all the convergent species of Anthomyza and
Castnia are represented in the figure; these are, at any rate, all
that are to be found in the collection of Mr. Herbert Druce
and the Hope Collection. There are, however, other species
of Hyelosia which I was unable to obtain at the time the photo-
graphs made use of in preparing these Plates were taken.

Microscopic examination proves that the moths become trans-
parent in two ways, which differ entirely from each other, and
from any of the methods already described in the butterflies.
A representation of the scales of the transparent part of the
Castnias is shown in PI. 44. fig. 6. The scales are not reduced
in size, but they have lost their pigmeut and are transparent ;
they are furthermore set up on edge so that the light freely
passes between them. The scales further from the opaque border
of the transparent patches are much more upright than those at
the margin which have been selected for figuring. The arrange-
ment in Hyelosia is shown in fig. 5, and it is seen to be closely
similar to that of Castnia, save that the scales are much reduced
in number. The lithograph of the scales of Anthomyza (fig. 4)
proves that the scales are normal in size and arrangement.
They lie flat on the wing-membrane with the usual overlap,
but are so transparent that the light freely passes through
them. Although transparent. they retain a more or less faint
yellow or greenish-yellow tinge, but this is also to be noticed in
the transparent part of the wings of Methona, Dismorphia, &c.

602 PROF. E. B. POULTON: NATURAL SELECTION

The comparison of these details is almost a demonstration of
the operation of natural selection. We cannot conceive of
natural sclection acting other than along some such lines as
those which have here been shown to exist ; for it is impossible
to believe that very different species with very different natures
would present anything but very different variations for its
action. On the other hand, we cannot conceive of any theory,
not dependent upon the principle of selection, which could
produce such extraordinary superficial resemblances among
numbers of species by methods which are eutirely unlike in all
their details.

13. The supposed direct Effect of Local Forces implies the
Hereditary Transmission of Acquired Characters.

Finally, the hypothesis which is more commonly than any
other substituted for natural selection, has the further dis-
advantage that it implies the unproved improbable hypothesis of
the hereditary transmission of acquired characters.

A discussion of this latter hypothesis cannot be attempted here.
It will be sufficient to observe that after years of search no
particle of evidence in its favour, which can stand the test of
examination, has been forthcoming.

14. Genera, Conciustons: Natural Selection as the Cause of
Mimetie Resemblance and Common Warning Colours.

I think it is not too much to claim that, even if the theories
which have been proposed as substitutes for Natural Selection,
have not been destroyed in single Sections of this memoir,—and
I confidently believe that they have been thus destroyed over and
over again,—their most convinced supporter will admit that they
must yield to the accumulated pressure of all the arguments
here brought forward.

The resemblances of mimicry and common warning colours
have certain salient features in common, certain peculiarities
which are apt to manifest themselves repeatedly ; they also bear
certain general relationships to other resemblances in organic
nature. In this paper I have attempted to set down all the
general statements which can be made as to the phenomena
under discussion. These general statements represent an

THE CAUSE OF MIMETIC RESEMBLANCE. 603

enormous number of observed facts in all parts of the world. I
believe that the generalizations will be admitted to be sound
and to be well warranted by the facts. Under any theory which
is not based upon selection, the whole of the facts on which
the generalizations rest become mere coincidences and receive
no explanation of any kind. Under natural selection this vast
body of facts becomes at once intelligible. Here the accu-
mulated facts of the most diverse kind, which receive an intelli-
gible explanation by the theory in question, yield a firm support
to the theory. There are many theories which are held upon
indirect evidence of precisely the same nature. We believe in
evolution, not because we see it taking place, but because
of the immense number of observed facts which it renders
intelligible.

In the case of natural selection in relation to mimicry and
common warning colours it is to be confidently hoped that direct
evidence may yet be added ; indeed, a considerable amount is even
now forthcoming. Prof. Lloyd Morgan’s recent work upon the
activities and instincts of young birds of many species (‘ Habit
and Instinct,’ London, 1896) proves that their education is
actually of the kind which is presupposed in the theories of
H. W. Bates and Fritz Miller. He shows that they have no
instinctive knowledge of things which are good for food, but
examine and test everything. On the other hand, they have very
retentive memories, and retain a firm impression of the appear-
ance of objects which have given them an unpleasant experience.
Furthermore, there was evidence that they are influenced in
their behaviour towards other objects resembling the one
which has proved objectionable to them. As to the aggressive
Hymenoptera, the evidence of their special methods of defence
is obvious to everyone. With regard to specially protected
groups of butterflies there is a large amount of evidence from
observation and experiment, but more is to be desired. As
this paper is going through the press I have received the last of
Mr. Frank Finn’s interesting and important set of papers on this
question (Journal, Asiatic Society of Bengal, Ixiv. pt. 1. 1895,
p. 344; lxv. pt. i. 1896, p. 42; lxvi. pt. 1. 1897, p. 528; Isvii.
pt. i. 1897, p. 614). After a long series of experiments begun
and conducted with a perfectly open mind, upon Indian insect-
eating vertebrates, Mr. Finn concludes in favour of the theories
based upon natural selection in the following words :—

604 PROF. E. B. POULTON: NATURAL SELECTION

* T conclude from these experiments :—

“1, That there is a general appetite for butterflies among
insectivorous birds, even though they are rarely seen when wild
to attack them.

“2, That many, probably most species, dislike, if not intensely,
at any rate in comparison with other butterflies, the ‘ warningly-
coloured’ Danaine, Acrea viole, Delias eucharis, and Papilio
aristolochie, of these the last being the most distasteful, and the
Danaine the least so.

“3. That the mimics of these are at any rate relatively'palatable;
and that the mimicry is commonly effectual under natural con-
ditions.

“4. That each bird has to separately acquire its experience,
and well remembers what it has learned.

“That therefure on the whole, the theory of Wallace and
Bates is supported by the facts detailed in this and my former
papers, so far as they deal with Birds (and with the one Mammal
used). Professor Poulton’s suggestion that animals may be
forced by hunger to eat unpalatable forms is also more than
confirmed, as the unpalatable forms were commonly eaten with-
out the stimulus of actual hunger—generally, also, I may add,
without signs of dislike.”

Mr. Finn concludes with some very valuable suggestions as to
the conduct of future experiments.

The chief objection that has been raised against the theories
of Bates and Fritz Miller, and which was often insisted on in
the discussion at the Entomological Society, is the want of
evidence that birds are in any important degree the enemies of
butterflies. Many excellent observers have rarely seen butter-
flles attacked by birds. On the other hand, the Miillerian
theory only presupposes that the young birds test the palatability
of a few members of each convergent group in their locality and
henceforward avoid all the members, so that the recent tendency
to explain so many of these resemblances on Miillerian rather
than on Batesian lines is in harmony with the conclusion that
the members of such groups are not greatly attacked.

As regards butterflies which do not exhibit these resem-
blances, | may point out that it is impossible to exhaust the
details of the struggle for existence even as regards a single
species, in the intervals of the time devoted to collecting. Such
an investigation would demand the whole time of a first-rate

a ee

+. Se ES

>

THE CAUSE OF MIMETIC RESEMBLANCE. 605

observer, and, so far as I am aware, the inquiry has never been
approached in so thorough a manner. LHven if collectors would
pay attention to the worst specimens instead of the best, some
evidence of the nature and amount of attack would be forth-
coming. During the visit of the British Association to Canada
last year (1897) I made a point of capturing butterflies which
had evidently been pecked by birds. In this way, although I
did not witness a single attack, I obtained the proof that butter-
flies are not nearly so immune as has been asserted. On
another occasion I hope to deal with this evidence in detail.
Similar observations have been made by Fritz Miiller.

The review of the whole subject during the past 36 years
increases our confidence in the theories of Bates and Fritz
Miiller, while it disposes of all alternative hypotheses. Even more
than this,-—it will, I believe, be claimed by all who take a broad
view over the whole field of evidence, that the explanation of
these deeply interesting facts, which form so fascinating and
important a department of natural history in the tropics, is one
of the most notable triumphs ever won by the great theory of
natural selection.

EXPLANATION OF THE PLATES.

Prats 40.
Certain aspects of Mimetic Resemblance in Lepidoptera.

Fig. 1. Natural size. This figure represents the appearance of the young larve
of Stawropus fagi in the first stage, upon a twig of birch. The drawing
was made from living specimens in the Hope Department at Oxford
by P. J. Bayzand. The ant-like appearance is produced by the
immensely elongated second and third pairs of thoracic legs, which are
kept ina state of active movement upon the slightest disturbance.
Tbe caudal shield with its two processes resembles the head and
antenne of an ant. This figure should be studied in connexion with
figures | to 7 in the text (Section 11), in which the resemblance to
ants is shown to be brought about in many very different ways, thus
supporting an interpretation based on the theory of natural selection.

Fig. 2. Natural size. A group of the young larvae of Hndromis versicolor upon
a twig of birch. Drawn from life in the Hope Department by P. J.
Bayzand. The larve are represented in the attitude they assume
when disturbed, the anterior part of the body being thrown back and
the orange-coloured thoracic legs exposed to view. In this position
they strongly suggest the appearance of a group of young larve of

LINN. JOURN.—ZOOLOGY, VOL. XXVI. 43

606 PROF. E. B. POULTON : NATURAL SELECTION

Tenthredinide, in which the posterior part of the body is raised on
disturbance, while the orange ventral glands are everted and rendered
conspicuous. The larve of Exdromis afford a good example of mimetic
resemblance almost entirely brought about by appropriate instincts
(the gregarious instinct, the movements and attitudes assumed on
disturbance). Thus the results are due to internal changes (in the
nervous system) with only a minimum of superficial modification.

Fig. 3. Natural size. A group of larger larve of Hndromis versicolor, also
drawn upon birch by P. J. Bayzand. The figure shows that as the
larvee increase in size their instincts undergo appropriate changes
leading them to form much smaller groups like the smaller groups of
the more advanced larvie of Tenthredinide, The attitude represented
is that which follows disturbance.

Figs. 4-9 were drawn from specimens in the Hope Department, and are intended
to show the retention of ancestral features upon parts which are
normally concealed and the restriction of mimetic modification of
colour and paitern to the parts which can be seen. They incidentally
show examples of the more complete development of mimetic resem-
blance in the female sex.

Fig. 4. About ? natural size. The male of Dismorphia praxinoe, showing the
manner in which the long-and-narrow-winged appearance of an
Ithomiine is brought about in this broad-hind-winged butterfly. A
comparison of the right and left sides of the figure clearly indicates
the manner in which this appearance is produced by the excessive
amount of overlap which occurs in the natural position of the wings
(left side). Hence the greater part of the surface of the upper side of
the hind wing and the under side of the fore wing is hidden ina natural
position, and upon these concealed parts a large patch of the old
Pierine white appearance is retained, seen, in the case of the hind
wing, on the right side of the figure. 7

Fig. 5. About 3 natural size. The male of Dismorphia praxinoe, showing the
under side of the wings in the natural position (left side), and with
the wings separated on the right side so as to expose that part of the
under surface of the fore wing which is normally concealed. Here,
too, a large white patch is to be seen.

Fig. 6, About 3 natural size. The female of Dismorphia prasxinoe, figured in
the same manner as the male in fig. 4, and showing that the wings are
really long and narrow with only the amount of overlap which is
usual in butterflies. No white is to be seen upon the narrow band of
the hind wing, which is concealed in the normal position.

Fig. 7. About 3 natural size. The female of Dismorphia praxinoe, figured to show
the under side of the wings similarly to that of the male in fig. 5.
Here, too, it is seen that no white patch like that of the male is
retained.

Fig. 8. About 3 natural size. The male of Dismorphia orise represented in the
same position as that of the male of D. prazxinoe in fig. 4. Here, too,
in this transparent, black-barred species, immensely modified by
mimetic resemblance, the white patch is still retained (in the male), as

THE CAUSE OF MIMETIC RESEMBLANCE. 607

seen on the right side of the figure. The female of the species is
shown in Group 1 on Plate 42 and two forms of it in fig. 1 on
Plate 44. The female of D. ovise lacks, as do all the females of the
genus, the chalky patches which are present in the males of many of
the species,

Fig. 9. About $ natural size. The male of Dismorphia orise represented in the
same position as that of the male of D. praxinoe in fig. 5, so as to
show the under side of the wings. The white patch is distinctly seen
on the normally concealed part of the fore wing (right side).

Prater 41,

The diverse methods by which Mimetic Resemblance is attained, often in
closely related species. I have been kindly permitted to make use of
specimens in the Natural History Museum for obtaining figures 1, 2,
54,5B,and5c, The remaining figures represent specimens in the Hope
Department.

Fig. 1. Natural size. A South-American Hemipterous insect belonging to the
genus Myocoris, probably M. braconiformis. The usual appearance of
a Hemipteron has been profoundly modified so as to produce a super-
ficial resemblance to the Aculeate Hymenopteron shown in the next
figure.

Fig. 2. Natural size. A South-American species of Braconide, probably
belonging to the genus Zphiaulax, closely resembled by the bug
shown in fig. 1. A large assemblage of tropical American insects
probably converges round this type of appearance. Already many
species of Braconide and of Hemiptera, together with two species of
moths (Syntomide), can be recognized. So far, all the members of
the assemblage belong to distasteful or specially defended groups, and
probably supply an example of Common Warning Colours.

Figs. 3, 4. 5, 6, & 7 show the different methods by which the resemblance to
stinging insects (Aculeate Hymenoptera) has been brought about in
nearly related Longicorn beetles,

Fig. 3. Natural size. A Mediterranean species of Clytine, Plagionotus
scalaris ; one of many species of the group in which the resemblance
to a wasp is produced by the black and yellow bands on the closed
elytra, as well as by the slender yellow legs and active movements, &c.
In this case the functional wings are concealed, and there is nothing
to suggest the wings of the model. In spite of this shortcoming the
living species of this group are very wasp-like, The resemblance is
carried to a still higher level by very different means in allied tribes
of Longicorns.

Fig. 4. Natural size. A species of Callichromine from the Oriental Region,
viz. Nothopeus hemipterus. In this beetle the general resemblance to
a dark-coloured slender wasp is carried much further than that of
Plagionotus (fig. 3) to a black and yellow wasp such as the species of
the genus Vespa. Furthermore the likeness is heightened by the
size and conspicuousness of the hind wings of the beetle, which are

608 PROF. E. B. POULTON: NATURAL SELECTION

freely exposed at rest as well as in flight. This is brought about by
a reduction of the elytra to small oval scales. Hence the resemblance
which is so largely due to the colouring of these parts in Plagio-
notus is here in an allied form due to the appearance of parts
which are concealed, except during flight, in the former. It is also
to be noted that the exposed under wings have the dark tint, which is
so common in the species of stinging Hymenoptera, and that such
exposure of the functional wings is very unusual in Coleoptera and
is almost, if not entirely, confined to the species in which their display
is made part of a mimetic resemblance.

Fig. 5. Natural size. An Australian species of Esthesine, Esthesis ferrugineus.
In this beetle the elytra are reduced to squarish scales and the under
wings freely exposed at rest. The bold and characteristic colouring
is found in many Australian insects of which three other examples
are figured here.

Fig. 5a. Natural size. An Australian fly, a species of Dasypogon (Asilide),
with the same type of colour and pattern as that of the last and the
two succeeding figures. This Dipterous insect resembles the wasp
represented in fig, 5 B with especial closeness.

Fig. 538. Natural size. An Australian wasp, Abispa australis (Eumenide),
which is in all probability the central type of this characteristic
Australian group.

Fig. 5c. Natural size. Another Australian wasp, Hwmenes Latreillei (Eumenide),
which falls into this group. Other species of Australian wasps are
also included, and probably additional insects of other Orders from
the same region will be found to possess a similar type of colouring.

Fig. 6. Natural size. A Brazilian species of Rhinotragine, Isthmiade braco-
noides, in which the very slender form strongly suggests the ap-
pearance of a Hymenopterous insect. The colouring and pattern of
the wings is very rare in beetles, and is in all probability mimetic of
some stinging insect from the same part of the world. The elytra
are here reduced to linear vestiges somewhat broadened at their
bases; they leave the under wings freely exposed.

Fig. 7. Natural size. A species of Hephestion sp. (Necydaline) from Chili.
The appearance is extremely wasp-like, and is brought about by a
reduction of the elytra very similar to that indicated in the last
figure. ‘The under wings are dark like those of so many wasps, and

the whole appearance most suggestive of a stinging insect and unlike
that of a beetle,

Thus in this Plate five species of nearly related Longicorn beetles are
represented. The appearance of a wasp is chiefly suggested by the colouring
of the fully developed elytra of one species (fig. 3), by the reduction of the
elytra and form and colour of the parts thus displayed, in all the rest (figs. 4,
5, 6, & 7). Furthermore the elytra are reduced to oval remnants in one (fig. 4),
squarish in another (fig. 5), and linear (with broader bases) in the remaining
two (figs. 6 & 7). Stout-bodied black and yellow wasps, such as those of the
genus Vespa (or the Adispa shown in fig. 5), are resembled by two (figs. 3 & 5),

en ea

|
|

A a iii lia

THE CAUSE OF MIMETIC RESEMBLANCE. 609

while the remaining species (figs. 4, 6, & 7) are mimetic of slender Aculeata
with opaque, iridescent, or even particoloured wings.

Hence in this group of allied species of beetles there is great diversity in
the methods by which the mimetic resemblance is attained and in the form of
the model which is imitated. One thing is common to the whole, viz. the
production of a strong superficial likeness to a stinging insect. Such com-
munity of end through diversity of means strongly supports an interpretation
based on the theory of natural selection.

Puare 42.

The resemblances between the members of a large and characteristic group
of transparent and black-barred South-American Lepidoptera.

Fig. 1. About § natural size. The group as described by Bates in 1862 (Trans.
Linn. Sce. xxiii. pp. 495-566). The names of the species are engraved
on the Plate. The species belong to very different divisions, families,
and sub-families of Lepidoptera. Thus the Ithomiine are represented
by two species in different genera, the Danaine by two species, the
Pierine by one, while the moths belong to the very different families
of the Castniide and Pericopide, the latter including Anthomyza.
Bates also described a group of smaller Ithomiine species, Dircenna
epidero, D. dero, and D. rhoeo, as well as another smaller moth
(Hyelosia tiresia) belonging to the Pericopide. Of these he made a
second group, but they are in reality continuous with that shown in
fig. 1, having a very similar arrangement of black bars and border on
their transparent wings. The group has in all probability converged
around the two common and widespread Ithomiine species of the
genera Methona and Thyridia. All members of the group, including
these latter, are to be looked upon as primitively opaque-winged. If
selection has been at work in the alteration of the scales so as to
render them transparent, we should expect that this common end
would haye been reached by very different methods in the case of
species belonging to such widely separated divisions.

Fig. 2. About 7 natural size. By the kindness of Messrs. Godman and Salvin I
have been enabled to figure, from their great collection, all the species
of butterflies which are now known to fall into the two groups described
oy Bates 37 years ago. The Danaine and Pierine remain unchanged,
but the /thomiine have been immensely enlarged, containing 20 species
in seven genera. Many of these are extremely rare, and so complete
w set could not have been got together from any other collection.
I also owe to their kindness the loan of butterflies shown in fig. 1 and
of some in Plates 43 and 44.

Many species of Castniide and of Pericopide have now been added
to the group. ‘Those which are here shown, as well as in fig. 1, and
those in Plate 44 were in part obtained from the Hope Oollection and
in part from the very fine collection of Mr. Herbert Druce, who kindly
lent them for the purpose of this memoir. ‘The set is probably
complete, or nearly so, except in the genus Hyelosia.

610 PROF, E. B. POULTON : NATURAL SELECTION

PLATE 43.

- The divergent methods by which the transpareney—an important element in
the resemblance between the members of the group shown on Plate 42—
has been attained in Ithomiine and Danaine.

Fig. 1. About } natural size. The two central Ithomiine species round which
the group has in all probability converged. The extraordinarily close
resemblance between these butterflies belonging to entirely different
genera is a very fine example of common warning colours. The
next two figures will show that so close a resemblance is quite super-
ficial ; for there is a very wide difference between the shapes of the
scales on the transparent areas of the two species.

Fig. 2. x about 15. In this and all the other figures of scales on Plates 43
and 44, a portion of the transparent patch at the apex of the left
fore wing has been figured, including a small part of the opaque
margin, so asto show the manner in which the transition is effected.
The scales of butterflies and moths overlap from the base of the wing
towards its apex, and therefore in the arrangement here adopted the
uppermost part of each figure lies in the direction of the apex of the
wing. On this Plate a portion of the opaque border on the outer side
of the transparent patch, viz. towards the apex, is represented, and is
therefore uppermost in each of the three figures. In Plate 44 a
portion of the border towards the base has been drawn, and therefore
occupies the lowest part of each of the four figures,

The magnification was as nearly as possible equal in all the seven
figures representing the scales. All the specimens which were thus
drawn belong to the Hope Department. Fig. 2 represents the scales
of Methona confusa, and it is seen in the upper part of the circle that
the opaque surface of the wing is covered with broad forked scales
regularly alternating with long and narrow ones. On the transparent
part the latter are reduced to extremely fine hairs, the former to
minute bifid structures.

Fig. 3. Xabout 15. The scales of Thyridia psidii. The scales of the border in
this species are seen to differ somewhat in form from those of Methona,
while the difference on the transparent part of the wing is very marked,
the narrow scales being far broader and the bifid ones of a different
shape. Thé method by which transparency is attained is however
the same although the details differ. A method similar in all
essential respects was found in all the Ithomiine species shown on
Plate 42. fig. 2. In certain species, however, the process was carried
further: thus in Dircenna the broader scales were reduced to ¥Y-shaped
hairs, as well as the others to simple hairs.

Fig. 4. About 4 natural size. The two species of the Danaine genus J¢wna,
I. ilione and I. phenarete, which enter the group. The Ithomiine
were formerly classed with the Danaine, but the important differences
between them are now generally recognized as justifying their separa-
tion into two distinct sub-families.

Fig. 5. xabout 15. The method by which transparency is attained in the
Danaine genus Jtwna is seen to be entirely different from that followed

THE CAUSE OF MIMETIC RESEMBLANCE. 611

in the Ithomiine (figs. 2 & 8). The scales on the opaque border of
the transparent area are almost uniform in appearance, forming the
sharpest contrast with those of the Jthomiine. In the transparent
part the individual scales are but little changed in appearance or
size, but they are immensely reduced in number, so that the light
freely passes between them. The scales which remain merely cause
the appearance of a grey dusting over the transparent surface. Jtuna
ilione was drawn for this figure, the other species, Z. phenarete, only
differing in this respect by its even more complete transparency in
consequence of a still greater suppression of the scales.

Thus the course of development leading to superficial resemblance
has been entirely different in the species of these two sub-families—
the Ithomiine and Danaine.

Puate 44,

Further divergent methods by which the transparency necessary for the

likeness has been attained in other members of the group shown on
Plate 42, viz. in the Piering, and among moths in the Pericopide and
Castniide.

Fig. 1, About 4 natural size. On the right side is seen the single Pierine

Fig. 2.

species which enters into this group, viz. Dismorphia orise. The
two upper figures are male and female of the type form found in
many parts of tropical South America. Opposite them on the left
is a male and female of the type form of Methona confusa. The
specimen represented in the lowest position on each side is a form
which occurs in Ecuador, in which the transparency is increased at
the expense of the black bars and border. It is very interesting that
this form of Methona should thus have been followed by the Pierine
butterfly in the same area. The former has recently been distin-
guished as Methona psamathe (Godm. & Salv.).

xabout 15. The scales on the transparent part of the left apical patch
on the fore wing of Dismorphia orise, the opaque border being repre-
sented at the lower part of the circle. In this case a third method
has been adopted. In Ithomiine the widely different scales were
yeduced in size and simplified in shape: in Danaine the uniform
scales were reduced in numbers rather than in size: while here in the
Pierine they have been reduced in size rather than in numbers, but
remain almost unchanged in shape. The amount of reduction in size
in this species is better seen by comparison with other parts of the
opaque surface upon which the scales are far larger than those of the
border which are shown in this figure.

Fig. 3. About } natural size. A set of day-flying tropical South-American

moths which fall into this group. The two left-hand rows are
five species of the genus Anthomyza. The upper pair are A. brotes
(Druce), male left, female right; the next 4. Buckleyi (Druce), male
left, female right ; the next 4. Swainsoni (Druce), both female ; the last
pair being A. praailla (Druce) on the left, and A. tiresta (Cr.) on the
right. The two lowest moths of the figure are Hyelosia tiresia (Cr.).

612

PROF. E. B. POULTON ON MIMETIC RESEMBLANCE.

These and the species of Anthomyza belong to the Pericopide
(Hypside). The row of moths on the right hand are widely separated
from the others, being the species of -Castniide which belong to the
group. The uppermost moth is Castnia linus (Cr.), the next C. dodona
(Druce), the next C. heliconioides (Herr.-Sch.), and the lowest C. micha
(Druce).

Fig. 4, xabout 15. The method by which transparency has been gained in

the genus Anthomyza. Although A. Buckleyt has been drawn for this
figure, the method is identical in all the species. The scales lie flat on
the wing with an entirely normal arrangement and overlap. Their
size and shape are but little altered, but the scales themselves become
quite transparent (compare the border scales seen in the lower part
of the circle). There is generally a little yellowish or faintly greenish-
yellow pigment retained, giving a tint which closely resembles that of
Methona and other members of the group, none of which are entirely
colourless. This method contrasts in a most interesting manner with
all the others made use of in attaining transparency as an element in
superficial resemblance.

Fig. 5. Xaboutl5. The scales on the transparent part of the wing of Hyelosia

tiresia, part of the border being shown on the lower part of the
figure. The scales are greatly reduced in number, they become trans-
parent and are set in a far more upright position on the wing-mem-
brane. In these ways they interfere but little with the passage of light.
The wide contrast with Anthomyza, belonging to the same family, is
of high interest.

Fig. 6. Xabout 15. The method adopted by the species of Castnia has much

resemblance to that of Hyelosia, although the forms are so distantly
related. All the species examined have the same method, the
scales of C. dinus being actually drawn for the figure. The
scales are not reduced in size and but little, if at all, in number,.
The shape is somewhat simplified, and the pigment is lost. At the
same time the scales are set much more steeply on the wing-membrane,
and the light freely passes between their rows. This is seen to be the
case in fig. 6, but further from the border of the transparent patch
they become practically upright. Even where the slope becomes less
the light still passes through (although less perfectly), because of the
transparency of the scales.

Hence in this large group of superficially similar South-American
Lepidoptera figured and studied in some of its minute details in Plates
42, 43, and 44, the transparency which is essential to mutual resem-
blance has been produced by widely different methods:—the same
end, viz., the production of a close superficial likeness, has been
reached from very divergent paths. Such a result, while harmonizing
with an explanation based on the principle of selection, and especially
on the theory of natural selection, is quite inexplicable on any other
hypothesis which has as yet been suggested,

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RESEMBLANCES IN PIERINAZ ano MOTHS.

An Experimental Enquiry into the Struggle for Existence in Certain
Common Insects. By Epwarp B. Pourron, M.A. PRS. Hope
Professor of Zoology, Oxford, and Cora B. SANDERS.

[An abstract of a paper read before Section D of the British Association at Bristol,
on Monday, September 13, 1898. Reprinted from the ‘ Report’ (pp. 906-909) of the
meeting. |

Many Lepidoptera have been proved to possess the power of adjusting the
larval or pupal colours to those of the immediate surroundings. This power can
only be exercised once in the case of the pupa (viz. at the end of larval life), and
rarely, if ever, more than once or twice in the case of the larva. Many naturalists
consider that the power is protective, and has been produced by the operation of
natural selection. Others have doubted this conclusion, and W. Bateson! has
attempted to cut away the foundation of such an interpretation as regards the
pupa of Vanessa urtice by arguing that there is no struggle for existence during
this brief stage. This argument was opposed, and the lines of an experimental
enquiry were suggested in the same year by one of us.” In the discussion which
followed a paper on mimicry, read before the Linnean Society on March 17, 1898,°
it was strongly urged, especially by Professor Weldon, that such an experimental
enquiry should be conducted. Our present work is the outcome of that discussion,
and we desire to express our thanks to the Government Grant Committee of the
Royal Society for assistance in carrying on the investigation.

We determined to concentrate our attention on the pupz of certain butterflies,
this stage being especially suitable because the chrysalis is motionless, and, there-
fore, remains in any position in which it has been fixed until it is seized by an
enemy or emerges as an imago. Our object was to decide :—(1) whether there is
a struggle for existence durimg the pupal stage; (2) whether the struggle, if it
takes place, is decided by the conspicuousness of the pupa.

The enquiry was almost confined to the pupa of Vanessa urtice, which ranges
from a brilliant golden appearance through increasingly dark varieties up to black.
Seven degrees of colour variation were distinguished, as in the researches into the
sensitiveness of this pupa to its environment.’ Captured larvee were placed in
boxes lined with gilt, black, and yellow paper, &c., so as to produce pup with
different degrees of colour, the aim being to obtain the most contrasted results.
The pup were then fixed to the surfaces upon which they are known to occur
in nature, and others upon which they may be supposed to occur—viz. the food
plant (nettle), tree trunks, fences, stone walls, and rocks—while a few were placed
on the ground. They were attached by small nails driven through the silken web, in
which the caudal hooks were entangled, or (in the case of the food plant) by sewing
the web on to the leaves or stem with green silk; in other cases the hooks were
entangled in the outer part of a little plug of cotton wool, which was forced into a
erack in bark, wood, or stone. Careful notes were taken of the degree of colour,
method, and height of attachment, character of surface, and the date of all visits
until the pupa either disappeared, emerged, or died. With very few exceptions,
visits were made every twenty-four hours, and in many cases at much shorter
intervals. Over 600 pupz of this species were thus fixed, and of these about

550 disappeared or emerged. The experiments were conducted in three different

1 Trans. Ent. Soc. Lond. 1892, pp. 212, 213.

2 Poulton, Trans. Ent. Soc. Lond. 1892, pp. 471-477.

3 Poulton, Natural Selection the Cause of Mimetie Resemblance and Common
Warning Colours. Not yet published.

* Poulton, Phil. Trans. Roy. Soc., vol. clxxviil. (1887), B. p. 320; and 7rans. Ent.
Soc, Lond. 1892, p. 362.

d 4

2

localities—Oxford, Switzerland, and the Isle of Wight—with very divergent
results, as will be seen from the following table :-—

Oxford Miirren Visp i ee :

Surfaces to which [aces eee Saga Rae
| Pupe were fixed June 25—-July 23 July 11—July 17 July 21-July 25) Aug. 2-Sept. 3

Taken | Emerged| Taken | Left | Taken | Left | Taken | Emerged
Bark ; : 3) 28 0 = — 5 16 135 84
iencesi: : : 4 0) 0 18 = == 90 8
Rock = = ae ih soHl 7 25 = =
Walls : 4h 0 a = 1 5 14 12
Nettle . AE ALT 4 = | = — — 20 15
Ground . : Salaepelal! 0 — _ — — = _
Totals. : 5 55 4 4 | 49 13) | 936 259 119

It should be noted that the Swiss pupze which were not taken are marked
‘left’ because many of them did not emerge, but were removed at the close of the
visit and used over again. The results thus tabulated leave no doubt about the
existence of an immense amount of extermination at Oxford, and, although much
less, a large amount in the Isle of Wight, while there was comparative immunity
in the two Swiss localities. This strong contrast is probably to be explained by
the scarcity of smail birds in the latter, and their abundance in England, and
especially in Oxford, together with the fact that the Oxford experiments were con-
ducted earlier in the year Other considerations point to the same conclusion
viz, that birds are the enemies in question ;—the inaccessible position of the pupe,
which were nearly always fixed at a height of about four feet from the ground ;
the fact that the hard caudal extremity was frequently left attached after the pupa
itself had been taken ; the excessive variation in mortality in neighbouring locali-
ties, corresponding to well-known facts in the distribution of birds. A watch was
kept in localities where the pupze were known to disappear rapidly, and, in a single
instance, a great tit was seen to creep over the bark of a tree, from which the
pupa was then found to have gone.

So far as the observations extended, we inferred that the comparative freedom
of the Swiss pupze from the attacks of Vertebrate enemies is compensated by the
far greater destruction of the larvee by insect parasites. It is probable that the
birds which attack the English pupz benefit the larvee by keeping down the
number of parasites.

In the course of the enquiry the possibility was suggested that perhaps the
birds actually see the pupz being suspended, and afterwards search the spot.
A large number of pupz were, therefore, fixed at night by the light of a lantern,
but, so far as we can judge from general impressions (the analysis of the results
being unfinished), no difference was caused. It was also thought that, when
several pupze were suspended in close proximity, the birds, after finding one or
more, might search the neighbourhood with especial keenness, so that the chances
of successful concealment would be smaller than those of isolated pups. In order
to test this suggestion, a number of pupze were scattered over a large area, succes-
sive individuals being separated by a distance of about 100 yards or more. Here,
too, our present impression is that little, if any, difference was produced. Both
these modifications of the usual conditions of experiment were made in the Isle of
Wight. |

The question whether there is a struggle for existence during the pupal period
of Vanessa urtice is answered with certainty in the affirmative as regards those
localities where small birds are abundant. In such places it is now proved that
there is a tremendous struggle with an immense mortality, in spite of the brevity of
the pupal stage (from ten days to three weeks in length).

Q
o

The attempt was also made to answer the second question whether the struggle
is decided by the conspicuousness of the pupa. First, as to conspicuousness in
form, pup were fixed to surfaces which unequally concealed them ; thus the rough
surfaces of stone and bark (rough-barked trees being almost invariably selected),
and the shelter afforded by overhanging leaves of nettle, concealed their rough
angular forms far more than the comparatively smooth surface of fences, Looking
at the table on page 907, it 1s seen that at Oxford butterflies only emerged from
pup fixed to nettles, while in the Isle of Wight the mortality on fences
(90 taken to 8 emerged), was enormously greater than on bark (135 to 84), walls
(14 to 12), and nettle (20 to 15). When therefore the pupa is suspended from a
surface against which it stands out conspicuously, it is in far greater danger than
when it is fixed to one upon which it is concealed. This result is inexplicable,
except on the theory that the sense of sight is important to enemies in the dis-
covery of the pupe.

Secondly, as to conspicuousness in colour. In Nature the golden forms of the
pupa of this species are produced upon nettle, the darker forms on walls, rocks,
fences, and probably bark ; furthermore the darkest varieties are produced on the
darkest surfaces. In fixing the pupe, part were distributed as they are in Nature,
while part were given a reverse arrangement—dark forms being fixed to nettle, and
golden forms to black fences and dark surfaces of bark, &c. Some of the experi-
ments gave extremely positive results, in that the mortality among the latter pups
was far higher than among the former; other experiments were negative. Until
the whole of the experiments have been analysed in far more detail than has as yet
been possible, we cannot make any statement as to the general bearing of the
inquiry upon the danger or otherwise of conspicuousness in colour, although the
danger of conspicuousness of form has been shown to be conclusively proved.

It may be supposed that the experiments were vitiated by the accidental loss
of the pup. Many considerations, however, indicate that no serious error has
been introduced in this way. In the absence of enemies, in Switzerland, the pup
remained suspended until they emerged or until we removed them, and this was
also the case in the places of small mortality in the Isle of Wight ; in many places
the ground was bare, and a fallen pupa (always searched for) would have been
easily seen; the hard caudal extremity was frequently left fixed to the supporting
surface; the different results obtained on fences and on bark, &c., are manifestly in-
explicable on this ground, the means of fixation being the same. The act of
emergence was obviously a much severer test of our method of suspension than
that supplied by the motionless pupa; and yet in a large proportion of cases the
empty pupal shell was found hanging to the support, and thus remained for days,
in spite of the fact that its lightness enabled each breath of wind to blow it about.

In future inquiries we trust that a pupa with a wider colour variation than
V. urtice may be available. All attempts to obtain the larva of Vanessa io were
unsuccessful, but next year we hope to get it and test the bright green and dark
forms which its pupa assumes, Other excellent examples would be the bright
green, bone-coloured, and dark forms of the pup of Pieris napi and P. rape,
which have the further advantage of being available for experiment during the
winter. Through the kindness of Mr. F. Merrifield we shall be able to carry on
the inquiry with these latter and some other species during the winter of 1898-9,
when we hope again to appeal to the kindness of the President and Fellows of
Magdalen College for the opportunity of continuing the investigation begun during
the past summer in the College grounds.

We also desire to acknowledge the kind assistance we have received from Miss
Drummond and Miss Sidgwick in taking notes of the Oxford pups during our
absence in Switzerland, and from Miss Notley for much kind help in the Isle of
Wight. Professor H. F. Osborn, Professor F. O. Bower, and Mr. Arthur J.
Evans also witnessed the experiments in the Isle of Wight, and offered valuable
suggestions and criticism.

The investigation, of which this is a brief epitome, was manifestly a preliminary
inquiry: it has nevertheless yielded far more definite results than we ventured to
hope for when we undertook it.

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Section D,—Bradford, 1900. |

(2. Observations on Mimicry in South African Insects.
By Guy A. K. MarsHAtt,

[Arranged and Communicated by EDWARD B. Poutton, M.A., F.R.S. Fellow of
Jesus College, Oxford, and Hope Professor of Zoology in the University. ]

The following paper is an abstract of the results obtained by Mr. Guy A. K.
Marshall in South Africa. When no locality is mentioned it is to be understood
that the observation was made at Salisbury, Mashonaland (5,000 feet). The
observations here briefly recorded have added in a most important manner to our
knowledge of the natural history (bionomics) of South African insects, a subject of
which the foundations were laid by Roland Trimen. Groups of mimetic Lepido-
ptera captured on the same day as their models have been obtained both from Natal
and Mashonaland (Salisbury), thus demonstrating more fully than has been done
hitherto the fact that model and mimic fly at the same time as well as in the same
place. The groups bring out the extraordinary power of Danaine butterflies in, so
to speak, moulding the species of other sub-families into a superficial likeness to
themselves. There are only four or five species of Danaine in the region under
consideration, and each one of them is the centre of a group of forms superficially
similar, but remote in affinity. The abundant and widespread Limnas chrysippus
was largely resembled both in Natal and Mashonaland. In experiments conducted
upon insect-eating animals, this butterfly appeared to be less unpalatable than the
Acrea (A. encedon) which resembles it. The explanation may be found in the far
wider range“of the Danaine model, which would render it familiar to enemies
passing from an area in which the Acrea does not, to one in which it does exist.

In the case of two forms of Euralia (E.mima and E. wahlbergi) mimicking two
very different species of Amauris (A. echeria and A. dominicanus) there is good
reason to believe that a single species has become dimorphic. Photographs of
four mima (two male, two female) and four wahlbergi (three male, one female) were
shown, the whole set having been part of a company of twelve individuals going to
rest together on a small clump of fern under a steep kraantz, Umbilo River,
Malvern, near Durban, Natal (June 28, 1897). The two forms have also been
taken in coitu, and have been found together freshly emerged from the pupa on the
same tree. Intermediate varieties are also known. Ifspecific identity be established,
the case will constitute a new form of mimetic dimorpbism in the Lepidoptera,
similar to that of the Dipterous genus Volucella. Allcases of dimorphism in mimetic
Lepidoptera hitherto described are either sexual or confined to a single sex. In one
sex, indeed, a mimetic species may be polymorphic, as in the female of Papilio
cenea or Hypolimnas misippus.

In the case of the distasteful sub-family Acreine, the two very different
species A. natalica and A. anemosa were shown captured at Salisbury on the same
day. Although entirely different in detail, the pattern of the two species is broadly
the same, and during flight would probably appear to be identical. Another even
more striking example of Miillerian or synaposematic resemblance was afforded by
a set of eighteen specimens belonging to five species of small Acrzeas captured on
the same day (December 31, 1898) in the same locality. The whole group pre-
sented a wonderfully uniform appearance in size, shape, colour, and the general
distribution of markings.

An example of a Hesperid (Baoris netopha) mimic of an Acrea (A. double-
dayt), the two captured on the same day, added another to the rare instances of
mimicry in this family. The resemblance is only seen in the attitude of rest,
and is confined to the undersides of the wings.

d 12

2

The aberrant Lyceenid (Alena amazoula) has a general Acreine aspect, and
is very unlike the well-known appearance of its true family. It is probably dis-
tasteful, and is resembled with tolerable closeness, especially upon the under-
sides of the wings, by a day-flying Geometrid moth (Petovia dichroaria), They
were captured together at Malvern, near Durban, on September 26, 1897.

A most interesting series of injured specimens of butterflies showed the
probable attacks of birds or lizards, observations in the field affording strong sup-
port to this interpretation. Members of the specially protected conspicuous
groups were, as Fritz Miller showed in the case of South America, also subject to
attack. Comparatively few of the injuries were inflicted at the junction of the
fore and hind wings, or indeed anywhere except at the apex of the fore wing, or,
more commonly still, at the anal angle of the hind wing. In many cases the
injury was symmetrical, indicating that a piece had been bitten out of both right
and left wings during the usual attitude of rest, or as they came together momen-
tarily in flight. The two points of special attack are commonly rendered con-
spicuous by special marks, and, in the case of the hind wing, structures such as
‘tails,’ ‘ eye-spots,’ &c. In the Lycenide, where the ‘tails’ are rendered still fur-
ther prominent by movement, many specimens were captured with these parts
bitten out from one or both of the wings.

Large additions were also made to our knowledge of mimicry and warning
colours in Coleoptera. In the Carabid genus Anthia, a probable warning character
common to different species consisted in a large, white patch, placed in some
species on the sides of the thorax, in others on the anterior surface of the elytra.
The general effect was the same, but the anatomical relations entirely alien. The
characteristic banded pattern of the Cantharide was shown to be resembled by
beetles of widely different groups, the spotted pattern of the Coccinellide by a
Hemipterous insect (Steganocerus multipunctatus), while the appearance of many
8. African species of Lycide, light-brown anteriorly and black posteriorly, was
reproduced in beetles of many groups, many species of Hymenoptera, a Hemi-
pteron, two moths from remote sections of the order, anda fly. In the vast
majority of these cases of likeness to and among Coleoptera, it is probable that the
resemblance is Miillerian (synaposematic) rather than Batesian (pseudosematic).
The interpretation of the remarkable likeness borne by a species of Longicorn
(Phantasia gigantea) for certain Curculionide is more uncertain, although it is
clear that some general principle is at work, inasmuch as resemblances between
other species of the same groups are well known in many parts of the world. An
interesting group of superficially similar insects from three different orders con-
sisted of a Bracon, a Reduviid bug, and a Longicorn beetle.

Evidence of the struggle for existence in Coleoptera was supplied by a group
of five beetles taken from the crop of a Guinea-fowl (Numida coronata). The
four species belonged to the Buprestids, Curculios, Longicorns, and Phytophaga.
All the beetles had been swallowed whole and were almost uninjured, even as
regards limbs and antenne.

Among the other orders of Insecta the Hemiptera afforded a wonderful
example of mimicry or common warning colours from Malvern, near Durban, the
Reduviid bug Phonoctonus nigro-fasciatus bearing the most remarkable likeness to
the somewhat smaller Lygeid Dysdercus superstitiosus. The mimetic resem-
blance of Diptera to Aculeate Hymenoptera was illustrated by many examples,
model and mimic having been captured in the same place and within the same
month. The most remarkable of these was a splendid new species of Hyperechia,
closely resembling the black, reddish-brown banded, African species of Xylocopa,
such as X. flavo-rufa. Instances of common warning (synaposematic) colours in
Hymenoptera were also illustrated by a group of three species with a general
resemblance to each other: the Aculeata being represented by a species of
Myzine and one of Ceropales, the Terebrantia by a species of Ichnewmon. All
were captured at Salisbury in January 1899.

The whole of the material here briefly described may be seen in the Hope
Department of Zoology, Oxford University Museum.

3. Observations on Mimicry in Bornean Insects.
By R. Suerrorp, B.A., Curator of the Sarawak Museum.
oF

[Arranged and communicated by EDWARD B. PouLTon, M.A., F.R.S., Fellow of
Jesus College, Oxford, and Hope Professor of Zoology in the University. ]

The following paper is an abstract of results obtained by Mr. R. Shelford,
B.A., Curator of the Sarawak Museum, British North Borneo. The vast majority
of his observations were made at or near Kuching, the capital of Sarawak ; a few,
however, in Singapore. When no locality is mentioned, Sarawak is to be under-
stood. The observations form a very important addition to our knowledge of
mimicry in Malayan insects, especially the Coleoptera.

Among Lepidoptera an Llymnias, believed to be a new species from Mount
Penrissen, is a tolerable mimic of the well-known Euplea, Tronga crameri.
Among the Chaleosid moths, three species of Isbarta mimicked two of Euplea and
one of Pierine (Delias cathara). ‘The latter is of considerable interest, inasmuch
as the Pierme model appears to be excessively rare. There can be little doubt,
however, as to the true relationship, for another species of the same genus,
I. pandemia, is a magnificent mimic of another species of Delias (D. pandemia),
both coming from Mount Kina Balu in North Borneo.

In the Neuroptera the Mantispides are shown to be mimics, a splendid new
species (M. stmulatrix, McLachlan) resembling a common Bracon flying with it
on Mount Matang, near Kuching, while a small species from Singapore (JZ. ? cora)
exactly mimicked an ichneumon flying with it.

It is in the Bornean Coleoptera, and especially the Longicornia, that by far
the largest additions to the subject of mimicry have been made. Many Longicorn
species, chiefly of the genus Oberea, were excellent mimics of the Braconide,
and perhaps other Hymenoptera. The long narrow form of the beetle resembled
the Bracon at rest with wings folded. As seenfrom the side, certain species of Oberea,
notwithstanding their uniform diameter, were apparently ‘waisted’ like a Hymeno-
pterous insect, the effect being due to a conspicuous white patch on the side of the
anterior abdominal segments. The part of the body thus covered is obliterated, while
the outline of the patch is such that the uncovered, and therefore conspicuous, part
of the body conforms to the shape of a slender ‘ waist,’ from the posteriar end of
which the abdomen gradually swells. ‘The effect in one species is as perfect as if
an artist had deliberately painted the profile of a Hymenopterous abdomen upon
that of a beetle. Among other examples of the same form of mimicry was a
magnificent Cerambycid from Mount Penrissen (Nothopeus or n. gen., n. sp.), a
beautiful mimic of the abundant wasp, Saliws sericosoma, which flew with it.
The common Dammar Bee (Trigona apicalis), which does not sting, but is
formidable because of its bite, is the centre of a group of three species with the
most remote affinities. Not only is there a Longicorn, Epania singaporensis, but
a Bracon and a Reduviid bug. The mimicry is probably Miillerian in most, if not
all, of the species of this group.

Another important set of Longicorns, species of Entelopes. Tropimetopa,
Chreonoma,and Astathes were extremely perfect mimics of Phytophaga(Galerucide).
In one large group both models and mimics were reddish brown, in another
iridescent blue-black, in a third anteriorly blue-black, posteriorly reddish brown.
Another species of Entelopes (FE. glauca) resembled a common Coccinellid (Caria
dilatata), a Cassid also falling into the group.

The Lyctde were models for Longicorns and other insects in Borneo no
less than in South Africa. Species belonging to the Longicorn genera Erythrus

4.

Ephies, Xyaste, and Eurycephalus mimicked Lycids with remarkable accuracy. In
the last-named genus one species, &. lundz, was a mimic, while another closely
related (4. cardinahs) exhibited a warning coloration of the most startling
character, an indication that the genus is distasteful and the mimicry Millerian.
In addition to these, the Lycids were mimicked by a Clerid beetle, by numerous
Hemiptera and a Zygzenid moth, the latter from Singapore.

The resemblance of certain Longicorns to the Rhynchophora was far more
evident than in South Africa, for not only was there a mimic ( 7rachystola granulata)
of a Curculionid (Sipalus granulatus), but there were species belonging to no less
than four genera mimetic of the Brenthide. These latter mimics hold their long
antenne extended forwards side by side, the tips only, or in some species the
anterior halves, diverging. Thus the rostrum of the Brenthid; together with its
usually short antenne, are represented by the long antennz of the Longicorn.
The Anthribide were mimicked by Longicorns of the genera Hreis and Cacia.

A feature of both Rhynchophorous models and their mimics, and one very
unusual in mimicry, is the inconspicuous mottled colourmg and the absence of
strongly contrasted tints.

A very interesting Longicorn mimic of an Endomychid beetle (Spathomeles sp.
near turritus) was a rare species of Zelota as yet undescribed.. The curved
spine on the elytron of the model was represented by a brush of hairs on that
of its mimic. Experiments indicated that the ndomychideé as a group were
distasteful, and large synaposematic sets of purplish black, yellow or orange
spotted species were found near Kuching together with several species of Hrotyhde
aud a Pentatomid bug with the same general appearance. Another group of dark
Endomychids was rendered conspicuous by numerous spines (Amphisternus).

Two groups of Longicorns were mimicked by other Longicorns belonging to
entirely ditlerent sections. The iridescent green Cerambycide of the genus
Chloridolum were closely resembled by two Lamude (Saperdine? genus, and
Chiorisanis viridis) and by the Cerambycid genera Xystrocera, Psalanta, and
Leptura, Many genera and species of the banded Cerambycid Clytine were
very closely mimicked by Lamizde and other Cerambycide. This last case is of
peculiar interest, imasmuch as the Clytime are themselves perhaps the most con-
spicuous mimics of Hymenoptera to be found in the whole of the Longicornia.
All over the world their numerous species commonly present a black yellow-
banded appearance bearing a general resemblance to wasps, while mimicry of
Mutilide, Cicindehde, and, in the allied T2llomorphine, of ants is also found.
When, therefore, we also find that this group itself furnishes numerous models to
other Longicorns we are driven to conclude that it is in some way specially
defended, and that its resemblance to Hymenoptera is Mullerian rather than
Batesian.

The mimetic resemblance to the aggressive and active Crcindehde was very
marked, examples being afforded not only by Longicorn beetles of the genera
Selethrus and Collyrodes, but also by a Dipterous insect found flying together with
its model (Collyris emarginata) on Mount Seramba, December 1898. ‘This is the
first example of the mimetic resemblance of a fly to a tiger beetle. The remark-
able Locustid mimic Condylodera tricondyloides (or a closely allied species)
described by Professor Westwood from Java was also rediscovered in Borneo, and
its habits for the first time observed.

Indirect evidence that the mimicry of Cleride is Mullerian rather than
Batesian is similar to that which pointed to the same conclusion in the Longicorn
Clytine. One Bornean species of a Clerid genus (Thanasimus) resembled a
Mutillid, another (genus near Tenerus) a Lycid, while a third, a species of Lemidia,
was mimicked by the Longicorn Daphisia pulchella.

Among the Diptera a splendid black Hyperechia (H. fera) was a beautiful
mimic of the abundant Xylocopa latipes, another example of parallelism with South
African bionomics. An allied species, Laphria sp. near Terminalis, was an
excellent mimic of Salus aurosertceus. Dipterous mimics of Hymenoptera are
extremely abundant in Borneo: remarkable among them was a species which
mimicked an ichneumon of the genus Mesosternus. The short antenne of the

5

fly in no way resembled the very long black and white ones of the ichneumon.
The fly, however, held up its black and white legs, applying their bases to its
head and moving them so that they closely resembled the antenne of the
Hymenopterous insect in movement as well as in colouring and proportions.
Another species of fly possessed true antennze which were remarkably long for
this order and thus closely resembled those of an ichneumon.

With few exceptions, the whole of the material here briefly described may be
seen in the Hope Department of Zoology, Oxford University Museum.

PRINTED BY
SPOTTISWOODE AND CO., NEW-STREET SQUARE
LONDON

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Extracted from the PROCEEDINGS OF THE ENTOMOLOGICAL
Society oF LonpDon of October 5th, 1898.

Prof. E. B. Poutron exhibited a series of Precis octavia,
(Cram.) var. natalensis Staud., and Precis sesamus, Trim.,
both captured and bred by Mr. G. A. K. Marshall at
Salisbury, Mashunaland (5000 ft.) and read the following
notes thereon :—

“The specimens exhibited prove conclusively (as Mr.
Marshall has shown in the July number of ‘The Annals and
Magazine of Natural History,’ p. 30) that these two butter-
flies, which differ so entirely in appearance and even in shape,
size and habits, are seasonal forms of a single species. Mr.
Marshall deserves the thanks of all naturalists for this dis-
covery of the most wonderful example of seasonal dimorphism
as yet known, and for the generous manner in which he has
despatched to this country the invaluable material constitut-
ing his evidence, so as to make it accessible for all time
in the British Museum and Hope Collection.

“Entomology is deeply indebted to Mr. Marshall for his
liberality now and at other times in placing large quantities
of valuable material at the disposal of investigators, and for
his readiness in undertaking biological observations ; and I
desire specially to thank him for constant help, and for
the valuable information contained in his letters, some of
which I hope to lay before the Society.

“The relationship between Precis octavia-natalensis and
P. sesamus has been described by Mr. Trimen (8S. Afr. Butt.,
i, and iii, app.), who connected the two forms by a number | of
comparatively rare intermediate varieties, which, having
regard to the fact that the forms themselves had been captured
in coitu, he regarded as probably hybrids. The conclusion
that the two forms were examples of seasonal dimorphism
was reached by Mr. C. N. Barker and independently by Mr.
Marshall (Trans. Ent. Soc. Lond., 1896, p, 557).

“Of this conclusion the specimens exhibited furnish
absolute proof. They include in the first place specimens
captured by Mr. Marshall towards the end of the wet (summer)
and beginning of the dry (winter) seasons.

a)
_

The red form (natalensis) was captured on Feb. 13, 1898,
(a pair in coitw) and Feb. 20, the blue form (sesamus), on
March 2, 12 and 16, and April 2. The latter form was
noted by Mr. Marshall to become dominant about the middle
of March.

“ Larve previously captured also produced sesamus at this
period, whereas an exactly similar individual that had been
placed ina damp jar on March 23 and pupated April 1,
produced on April 13 the typical natalensis now exhibited.
It would be of the utmost value to have many more ex-
periments of the kind.

“Of far greater importance is the fact that Mr. Marshall
has succeeded in breeding sesamus from natalensis in three
cases (‘ Ann. and Mag. Nat. Hist.,’ /.c.) ; two are exhibited,
the third is in the British Museum. Of the former, one
parent laid three eggs on Feb. 27, of which two produced
butterflies, one a typical sesamus (larva, March 5-31; pupa
March 31 to April 15), the other a red natalensis tending
slightly towards sesamus in its heavy black markings and
blue-marked border (larva March 5 to April 5; pupa
April 5-20). The second parent laid one egg on March 6
which produced a typical sesamus (larva March 12 to April 7 ;
pupa April 7—30). This larva was placed in a damp jar
from March 30 to April 5, but the ‘dry form’ of imago
emerged. The third example, in the British Museum, is a
sesamus bred from an egg laid by the summer form on Feb.
13 (larva, Feb. 19 to March 20 ; pupa, March 20 to April 4).

“As to the meaning of this seasonal dimorphism, I agree
with Mr. Marshall’s suggestion that the red natalensis,
appearing as it does in the keener struggle! of the warm

1 Mr. Blandford has pointed out, and the objection occurred to me
independently, that the press of life in the warm damp summer may
not, and probably does not, mean greater chance of destruction for
any individual insect, the larger needs of enemies being more than
compensated for by an increased supply of food. The question can-
probably be decided on the spot, and I am writing to Mr. Marshall
on the subject. In this climate the argument certainly holds good,
the danger to individuals of the comparatively few winter species
being probably far greater than to those of the numerous summer
forms. Granted that somewhat similar conditions exist in S. Africa,
it might appear that the mimetic form would be found at the time

3

damp summer, is probably an incipient mimic of an
Acrean type of colour and markings. Mr. Marshall has
suggested that natalensis presents many similarities to Acrea
acara, Hew., especially upon the under side of the wings,
The resemblance is probably closer when the insects are flying,
the most noticeable difference being in the shape of the
wings.

“Tt is in favour of this interpretation that a comparison
with allied species of Precis tends to show that the blue form
is ancestral and the red a recent departure ; furthermore, the
latter differs from the former and all its allies in the general
similarity of the upper and under sides of the wings—a
characteristic of the Acrainz, as of all specially protected
groups of butterflies.

* Mr. Marshall’s suggestions command increased confidence
because of his intimate knowledge of the habits of the two
forms. His description of the wide differences therein, as
well as in their appearance, is full of interest. The red
natalensis is described as frequenting higher and more open
country, while seswmus is found in shady places and gardens,
and is distinctly warier and more difficult of approach when
not feeding.

“ Problems of the deepest interest remain to be attacked.

of greater stress, viz., the dry winter season. But in England,
although there are abundant examples of insects with warning
colouring and many which mimic them in the summer, when there
isa plentiful supply of palatable food, in winter none are to be
seen. Those, such as Coccinellide, which exist in the perfect form,
hide themselves. 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. Experiments
with hungry animals support this view. It is possible that the con-
ditions are similar in 8. Africa, 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 here-
after be shown that Precis is to some extent unpalatable, and that
its resemblance to an Acrzan 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, It should be noted,
however, that Mr. Marshall (/.c.) is strongly of opinion that the
species is not unpalatable, and that the colours of the summer form
are not aposematic.—K, B. Pourron, November, 1898,

4

Thus Mr. Trimen has described the pup of the two forms as
differing, and Mr. Marshall considers that both are certainly
procryptic in colouring, that of sesamus being gilded and con-
cealed by its harmony with the yellow winter foliage, while
that of natalensis is dull brownish-black, with black patches
on the wings, and is protected among the dark withered leaves
of the moist summer. Im one case he has bred the red form
from a gilded pupa, but never sesamus froma dark one. It
would be of great interest to ascertain whether these
pupe exhibit true seasonal differences, or whether, as seems
more probable, the mature larve are susceptible to the
colours of their immediate environment, as has been
proved to be the case with those of several of the allied
Vanessidee.

‘“‘Of still more importance is, the decision as to the cause
or causes of the seasonal differences of the imago. On this
question Mr. Marshall maintains that ‘it at once becomes
evident that the directly exciting cause is a climatic one’
(l.c., p. 37), and he argues that moisture, rather than tempera-
ture, supplies the efficient stimulus. The great want in the
matter is experimental inquiry on as extensive a scale as
possible, carried on over a large part of the year, so as to in
clude larve and pupex, that in the ordinary course certainly
produce the blue form, as well as those that produce the red,
and those that, at intermediate times, are known to produce
both forms.

“ Concerning those last-mentioned, Mr. Marshall has shown
conclusively that the offspring of a single parent, brought up
under the same conditions, may belong to both forms; in
fact, in his experiment already described, the summer form
emerged five days later (7.e., when the winter season was to
that extent advanced) than the winter form. This fact leads
us to contemplate the possibility that the change of form is
due to internal causes and independent of, although on the
whole synchronous with, the seasons.

“‘ And even if some exciting cause be proved to be necessary,
it is probable there are great differences in the internally
caused degree of Susceptibility to stimulus in the two seasons,
while a mixed susceptibility exists in the intermediate period.

5

“Tt is likely that Mr. Marshall’s observations are to be
explained-by one or other of these alternatives.”
The Presipenr discussed Prof, Poulton’s exhibit, and also
congratulated Mr. Marshall on the importance of his
discoveries,

% * % * * * * *

Mr. F, MerrirteLp read a paper, illustrated by a large
number of specimens, on “The Colouring of Pupex of P»
machaon and P. napi caused by the exposure to coloured
surroundings of the larve preparing to pupate.”

It had generally been considered that the larva of P.
machaon was insusceptible to its surroundings when at the
period of pupation, but at the instance of Prof. Poulton he
had tried experiments last summer which proved that this was
not the case. Of eleven pups of the first brood obtained in
July, six larvee were set to pupate on dark sticks; four of
these were of the bone-coloured form, but a fifth was inter-
mediate and the sixth fell to the earth, where it formed a
green misshapen pupa. Of the five larve set to pupate on
white sticks of peeled willow, four pupated on them and were
all green, the fifth pupated on the white muslin top and was
bone-coloured. Of seven other pupz that were found to have
attached themselves to the stems of the green carrot tops on
which they were fed, six were green and one bone-coloured,
This made a strong case, but experiments tried on a larger
number of the larve of the second brood in September were
conclusive. Of 16 pupated on black paper or dark sticks,
all but one were bone-coloured, and most of them dark, four
that pupated on green carrot tops were all green, and of 19
others that pupated on yellow or orange all but one were
green. Various intermediate colours gave mixed results ; abso-
lute darkness produced five bone-coloured pup but of a pale
hue. Seventy-two pupz had been obtained, and all these were
exhibited, as were the pup or pupa-cases of the eleven of the
summer brood that had pupated. Several hundred pupe of
the second brood of P. napi were also exhibited, and these
were shown .to be exceedingly sensitive. Of ten on black

6

paper all were dusky with much spotting of dark brown, so
were seven out of eight on dark sticks, the eighth being
green. Of twelve on green cabbage leaves or on the glass of
the bottle containing them the majority were green or
greenish ; seven out of eight of those on yellow paper, and all
the eleven on orange leno were green ; of seven in absolute
darkness six were green, two much spotted with dark brown,
one bone-coloured with much spotting. He also exhibited
28 pupe of P. brassice and about 40 of P. rape, with results
in accordance with those obtained and recorded by Prof.
Poulton. In the case of both these species he had some con-
fined during the whole period of susceptibility to clear glass,
with no substances near that reflected any colours or white or
black ; the results very nearly resembled some kept in absolute
darkness, except that those on the clear glass were a little
darker. One very interesting napi exhibit was that of the sheet
glass roof of a breeding cage in two compartments, one having
the interior partly orange and partly yellow, the other black; in
the orange-yellow compartment 46 had pupated on the roof on
which orange leno had been placed, transmitting orange light,
and all but four of these were green with very little dark
spotting ; in the other compartment 34 had pupated on the
glass roof which had been covered outside with opaque black
paper and all of them were bone-coloured, most of them much
spotted with dark brown. He thought it was impossible to
doubt that much of the colouring was protective.

Mr. Bateson said that he had lately bred P. nap on a large
scale, and as regards that species he could quite confirm
Mr. Merrifield’s observations. His own experiments had been
undertaken with a different object, but he had frequently
noticed that there was a fairly close correspondence between
the colour of the pupz and that of the substances to which
they were attached, though exceptions to this rule were not
uncommon. He had moreover observed that in this respect
there was considerable difference between different families of
larvee. In his experiments the offspring of each pair were
kept separate, and in some families many larve pupated on
the food plant, while in others scarcely any did so. In some
families the pupal colours conformed in nearly every case,

7

while in other families the unconforming pup were numerous
The pup were often clustered together in little groups, and
individuals composing such clusters sometimes agreed with
each other in conforming or in not conforming.

He did not propose on that occasion to discuss the general
proposition that the colour-resemblance was in these cases a
protection ; but he pointed out that if a larva had the power
of forming a green pupa when attached to a green plant the
conformity of colour would make the pupa less conspicuous
only so long as the plant remained green. In the case of the
summer brood of nap, for instance, this resemblance might
perhaps be effective, though in this case the pupal period was
very short, only two or three weeks. But in order that the
resemblance may benefit pupze which over-winter it is necessary
that the green plant to which green pupe are attached should
remain green in the winter. His own pupe had turned green
on several food-plants which wither and go brown: in these
cases the pup became conspicuous again,

To form a sound judgment on this matter it would be
necessary to know how the pup occur in nature. Pupe of
P. napi were hard to find. Had any one found the green
pupz in the wild state ?

Prof. Poutron wished to congratulate Mr. Merrifield on
the results he had obtained. He was particularly interested
to know that the pup of ?. machaon had now been proved
to be sensitive. His own experiments on the species led him
to infer the opposite, but they had been conducted with very
few individuals, and he had always hoped that the species
would be investigated on a large scale.

The results obtained by Mr. Merrifield upon Pieris napi
were very striking and showed that: the mature larve were
most sensitive to the colours of their surroundings. The
strong influence of orange and yellow backgrounds in pro-
ducing green pup was corroborated by his own experience
and confirmed the conclusion reached in his earlier experi-
ments, that the efficiency of the greens of nature in producing
green pupze (and larvee) in sensitive species was due to the
orange and yellow rays which they reflect.

He exhibited some pupze of Piers brassicw which had been

8

rendered black-spotted and green by black and yellow. sur-
roundings respectively, anda few pupex of Papilio podalirius
which seemed to indicate that that species was sensitive. In
some experiments which he had conducted during the past
summer upon the struggle for existence in pupe, very large
numbers of pup of Vanessa urtice were produced upon back-
grounds of various colours, with results which confirmed
the conclusions he had already recorded. He hoped soon to
bring a detailed account of these observations upon pupal
colour-adaptation before the Society.

Replying to Mr. Bateson’s objections to the interpretation
of the colour-susceptibility of the pupe of Pieris napi as of
protective value, he considered that it was in every way
probable that the larvee wandered before pupation (as those
of allied species were known to do) and that their power of
becoming green was of value when they were fixed to grasses
or other plants which remain green through the winter.
Even if all such green surfaces were covered up with snow in
the more northern part of the range of the species, this would
not be the case for the whole of the winter pupal period, nor
would it hold good for other parts of the range. As to the
objection that the results obtained with bred pup were not
invariably consistent, in his experience the more completely
the experiment was conducted with this sole object in view,
the more uniform and convincing was the result.

Furthermore, the conditions of even the most carefully
conducted experiments differed in many respects from those
which obtained in nature. The proof required by those who
objected to an interpretation based on natural selection was
the discovery in natwre of the exceptions noted in their
experiments—that of light green pupe of P. napi or P. rape
or the golden or green pupz of Vanesse upon grey stone, or
the dark forms of all these upon leaves.

Although some of these pupz had been often observed in
one or both situations he had not heard of any single ex-
ception to the rule; and if they occurred they must be
excessively rare.

( 369 )

XU. The Colowr-relation between the pupe of Papilio
machaon, Pieris napi and many other species, and
the surroundings of the larve preparing to pupate, ete.
By F. MERRIFIELD, F.E.S.,and Epwarp B. Pout-
TON, M.A., F.R.S.. etc., Hope Professor of Zoology
in the University of Oxford.

[Read October 5th, 1898.]

A.—INTRODUCTORY. (F. Merririevp and E. B. Povrtoy.)

B.—EXPERIMENTS UPON THE PUP OF Papilio machaon.
1. Experiments wpon the Summer Pupx of Papilio machaon.
(HM)
2, Results of the above Experiments. (E. B. P.)
3. Experiments upon the Winter Pupx of Papilio machaon.
(F. M.)
4. Resulis of the above Experiments. (E. B. P.)
5. Mr. C. V. A. Peet’s Winter Pupe of Papilio machaon
described. (EK. B. P.)
C.—EXPERIMENTS UPON THE PUP OF Papilio poda-
lirius. (Cora B. SANDERS and E, B. P.)
D.—EXPERIMENTS UPON THE PUP OF Pieris napi.
1. Experiments upon the Winter Pupe of Pierisnapi. (F. M.)
2. Results of the above Experiments. (E. B. P.)

H.—EXPERIMENTS UPON THE PUP OF Pieris brassicz.
1. Experiments wpon the Winter Pupe of Pieris brassice.
(F. M.)
2. Results of the above Experiments. (E. B. P.)
3. Experiments wpon the Winter Pupe of Pieris brassice.
(E. B. P.)
4. Experiments with conflicting colours upon the Winter Pupe
of Pieris brassice. (E. B. P.)
F.—NOTES ON THE STRUGGLE FOR EXISTENCE IN
THE LARVZ OF Pieris brassicx. (E. B. P.)

G.—EXPERIMENTS UPON THE PUP OF Pieris rape.
1. Experiments upon the Winter Pupe of Pierisrape. (F. M.)
2. Results of the above Experiments. (E. B. P.)
TRANS. ENT. SOC. LOND. 1899.—PART IV. (DEC.) 25

370 Mr. F. Merrifield and Mr. E. B. Poulton on

H.—EXPERIMENTS UPON THE PUP OF Vamesside.
1. Experiments upon the Pupx of Vanessa urtice and Pyrameis
eardui. (C. B.S. and E. B. P.)
2. Haperiments upon the Pupe of Vanessa io. (MABEL E.
Nortiey, FLorENcE A. Wricut and E. B. P.)
I.—EXPERIMENTS AND OBSERVATIONS UPON THE
SUSCEPTIBILITY OF CERTAIN LEPIDOPTEROUS
LARVA AND PUPA TO THE COLOURS OF THEIR
SURROUNDINGS. (A. H. Hamm and E. B. P.)

K.—OBSERVATIONS ON THE COLOUR-RELATION BE-
TWEEN A COLEOPTEROUS SPECIES (Cleonus sulcirostris)
AND ITS SURROUNDINGS. (W. Hotzanp and E. B. P.)

L.—APPENDIX. THE QUALITY OF LIGHT REFLECTED
FROM THE COLOURED AND OTHER BACK-GROUNDS
EMPLOYED IN THE EXPERIMENTS RECORDED IN
THE PRESENT MEMOIR. (Sir Jonn Conroy, F.R.S., and
Dig 855m)

A,—INTRODUCTORY.

In Prof. Poulton’s paper in the Philosophical Transactions
of 1887, vol. 178 B. pp. 311-441, “An Enquiry into the
Cause and Extent of a Special Colour-relation between
certain exposed Lepidopterous Pupz and the Surfaces
which immediately surround them,” he recorded some
experiments on the full-fed larvee of Papilio machaon from
which he inferred that this species was not susceptible to
the colours of its surroundings, a conclusion which sur-
prised him, having regard to the marked dimorphism of
the pupze [the larva not showing any corresponding dimor-
phism to which the different colours of the pupze could be
ascribed, as in the case of the geometrid genus Hphyra
(Phil. Trans. /. c. p. 437)], and Prof. Poulton suggested that
further experiments should be tried, more especially as he
had had only eleven larve, of which two died. At the
meeting at Cambridge in August last of the International
Congress of Zoology M. Bordage of Réunion communicated
a paper in which he expressed the opinion that the pupz
of the genus Papilio appeared to have lost any suscepti-
bility to colour which they might at one time have
possessed, but Mr. Trimen, your President, gave an instance
to the contrary, and expressed the opinion that too few
experiments had been made to warrant at present any
conclusion on the subject.

Adjustment of colour in various pupex, etc. 371

Early in July last I happened to mention to Prof.
Poulton that I had then a considerable number of larvee of
P. machaon which I had received from Germany, and at
his suggestion I experimented on some of those I had left
at this time, receiving much useful information from him
personally as well as from the very full record of his
experiments on other species, in the paper above referred
to, and in his subsequent paper in the Transactions of this
Society for 1892, pp. 293-487.

The experiment thus begun gave rise to many other
experiments in which different species were employed.
The results were in large part displayed and an account
given of them at the meeting of this Society on October
5th, 1898 (Proc. Ent. Soc. Lond. 1898, pp. xxx—xxxi).
Professor Poulton was at the meeting and also showed the
results of some further experiments he had been making
in 1898 (Proc. Ent. Soe. /. ¢. pp. xxxii—xxxuli). The same
day I suggested to him that 1t would be useful to arrange
the results of my experiments according to the standards of
colour which he had already published, and construct fresh
standards for the species with which he had experimented
but little or not at all. He approved the suggestion and
agreed to arrange the results accordingly, and also offered to
include the results of the experiments he had recently made.

This memoir accordingly appears in our joint names.
The name or initials of the worker who conducted the
investigation will appear in the heading of each descrip-
tion. Professor Poulton is responsible for the new
standards of comparison and the tabulation of all the
pupz. In arranging the pups, he allowed due weight to
the description of my results so far as I had made one.

F. MERRIFIELD.

A few brief words are all that are necessary to form my
introduction to our joint paper ; inasmuch as Mr. Merrifield
has fully explained the circumstances under which it came
to be written. I was only too pleased to act upon his
suggestion, and thus to combine the record of our investi-
gations. It is more convenient in every way that there
should be a single complete account instead of two less
complete ones. The method adopted of placing the initials
of the writer after the title of each Section he has com-

municated makes our individual responsibility perfectly
clear.

372 Mr. F. Merrifield and Mr. E. B. Poulton on

A portion of my investigations were carried on jointly
with Miss Cora B. Sanders, of Lady Margaret Hall,
Oxford. These Sections are preceded by her name or
initials as well as my own.

The experiments upon the colours of the pup of Vanessa
vo were carried on, under my direction, by Miss Mabel E.
Notley and Miss Florence A. Wright, of Lady Margaret
Hall. Their names are added to the title of the Section in
which the experiments are described.

Some interesting observations of Mr. A. H. Hamm and
Mr. W. Holland of the Hope Department of Zoology,
Oxford, form in large part or entirely the subjects of two
Sections to which their names have been added.

Sir John Conroy, F.R.S., very kindly helped me in
determining the quality of the light reflected from the
various backgrounds, and his name has been similarly
added to the Appendix in which our results are described.

I wish also warmly to thank Mr. W. Holland and Mr. A.
H. Hamm of the Hope Department for their most efficient
help in many parts of the work; Mr. C. V. A. Peel for
kindly lending specimens which have been described, and
Mr. Arthur Sidgwick and Mr. Nicholson for drawing my
attention to interesting observations made by them which
are recorded in Section I.

E. B. Poutroy.

B.—EXPERIMENTS UPON THE PUPA OF

Papilio machaon.

1, EXPERIMENTS UPON THE SUMMER PUP OF
Papilio machaon. (FE. M.)

Iam generally away from home from nine or ten o'clock
until five or later, and therefore my opportunities for
taking a larva just at the right moment, when it has
done feeding and before its sensitiveness to colour has
begun, are imperfect. But I was able to select fourteen
larvee, which seemed to have arrived at the full-fed stage,
and I prepared a few glass cylinders of about 6 inches in
height and mostly 3} inches in diameter, though some
were an inch or two wider, which I placed in flower-pots
nearly filled with earth. They were in two divisions, viz.
(1) furnished with dark sticks, and (2) furnished with hght
sticks. Division (1) had about six dark brown sticks of
from } to % of an inch in diameter taken from an old

Adjustment of colour in various pupe, etc. 373

faggot-stack, and had a roof of black net. Division (2)
had the same number of peeled and therefore nearly
white willow slips of the same size, and had a roof of
white muslin. The sticks were secured by driving them
into the earth, and the cylinders were in both cases
exposed to the same amount of light, 7. close to a
rather large window, but on most days moved from the
W.N.W. to the E.S.E. side of the house, and back, to avoid
hot sunshine. As larvee, where crowded, have been found
to affect each other’s pupal colouring (Phil. Trans. 1887,
l. c. Ent. Trans. 1892, 1. ¢.), only two or three were placed
in each cylinder.

I exhibit the eleven pupz or pupa-cases, which were
obtained under these conditions. As the dark pigment
resides in the pupa-case, it is easy to see on examining
this whether the pupa was a green one or not, as, if the
pupa was a green one, the pupa-case is practically free
from pigment, though sometimes stained in places by the
meconium, whereas if the pupa was bone-coloured the case
is much darkened by brown or black markings.

It was obviously desirable to ascertain what was the
proportion of grey * and yellow-green pupe in my stock, and
on going through the whole remainder of them, 145 in
number, I found that 76 might be described as of the
former and 69 of the latter or yellow-green form. Many
of these however were of intermediate colouring, and
it was not easy to classify them. Seven of the 145 had
pupated on the green carrot-tops on which the larvee had
been fed; of these six were yellow-green, one was grey.

This made a very strong case for susceptibility, but it
was not conclusive, and I determined to try the experiment
on a larger scale with the second brood.

Before describing in detail the results of this and other
experiments it is necessary to tabulate the colours which
the pupa of P. machaon is known to assume, so as to
have a standard of comparison. The following table and
description were made by Prof. Poulton on May 2nd, 1899,
after carefully comparing the whole of my pup together
with a number belonging to Mr. C. V. A. Peel.

2. RESULTS OF THE ABOVE EXPERIMENTS. (KE. B. P.)

There is a distinct dimorphism in the pup of Papilvo
machaon, and the intermediate forms are very rare as

* See note on page 374.

374 Mr. F. Merrifield and Mr. E. B. Poulton on

compared with the extreme. The two forms may be
classified as (1) yellow-green, and (2) grey.* The former
have a yellow ground-colour with deep green markings,
the latter a bone-coloured ground with dark purplish
brown, and in places black markings. With few exceptions
the dark markings of the grey forms correspond in position
with the green markings of the yellow-green forms, the
chief exceptions being on the surface of the pupal wings,
where the green forms a more continuous area, less
Anterrupted by the yellow ground-colour than the dark
marking is by bone-colour.

This correspondence in position becomes all the more
interesting and remarkable when it is remembered that
the two markings are entirely different in constitution,
origin, and even in the pupal layers in which they are
respectively situated, the green pigment being relatively
unstable, probably a modified form of chlorophyll derived
from the food-plant, and situated in the deeper laminated
layers of the pupal cuticle, the dark pigment being very
stable (remaining permanently in the empty pupal case),
solely due to the metabolic activity of the animal organism,
and confined to the thick layer of cuticle which lies above
the lamimated layer and forms the outermost part of the
pupal shell. (See Poulton, Proc. Roy. Soc. 1885, Vol.
Xxxvill, p. 279, in which however the laminated layer is
erroneously distinguished from the “ true cuticle.”)

We can again classify the (1) yellow-green and (2) grey
forms as (a) dark and (6) light, thus :—

; a. Light forms.
1. VYellow-green | ne

a. Light -
2. Grey te Dark

»”

2?

* I formerly spoke of the darker pupz as brown (Phil. Trans. 1887,
I. c. p. 407), a description which is clearly incorrect. Mr. Merrifield
has introduced the appropriate term “bone-coloured,” which accurately
expresses the appearance of the ground-colour of these pupe as well
as those of the corresponding forms of Pieris napi and P. rape. The
whole appearance of these pupe is however due to the combined im-
pression made by the pale ground-colour and the dark markings, and
I think that “grey” expresses this effect as a whole more truly than
any other word ; although there are pup in which the markings are
so inconspicuous that the effect seen is that of the ground-colour
alone.

HUB.

Adjustment of colour in various pupx, etc. 3875

A light yellow-green form (1a) is bright yellow over the
dorsal surface slightly mottled with green on the sides, this
colour becoming distinct round the spiracles and forming
a spiracular streak interrupted between the abdominal
seoments. At the posterior extremity of the pupa this
ereen streak is continued on to the sides of the anal spine,
and in the more strongly marked pupz of this degree the
whole of the spine is green. The green mottlings are
rather more distinct upon the abdominal segments below
the spiracular band than they are above it, and have a
more pronounced longitudinal arrangement. The dorsal
region of the thorax is also more or less mottled with green
which becomes concentrated to form a distinct apical patch
on the mesothoracic median spine. The wings, limbs,
antennee, and head are deep green, the latter being, in the
palest pupze, the most strongly coloured part of the whole
surface.

In the more deeply coloured of the light forms (1 a)
the mottled green tends to spread backwards from the
mesothoracic spine forming a broad indistinct greenish
band.

In the dark yellow-green pupe (1 5), further development
and coalescence of the mottlings transform this dorsal band
into deep green. It occupies the whole dorsal area between
the subdorsal rows of small tubercles. Below, the green
of the spiracular stripe,—continuous in pup of this degree
—spreads upwards and invades the yellow ground-colour
leaving only a yellow lateral band, sharply defined above
where it terminates at the level of the subdorsal tubercles,
while below it gradually passes into the invading green.
Anteriorly this yellow band terminates below the meso-
thoracic spine. The dorsal green band is palest (yellowest)
in its median part and in the posterior half of its length,
in front of the caudal spine, the dorsal surface of which is
green. The band is also interrupted in the region of the
metathorax by a yellow patch, traversed by a green median
line.

The light grey forms (2 a) resemble the light yellow-
green (1 a), substituting bone-colour for yellow, and dark
purplish-brown for green, allowing of course for the greater
contrast between ground-colour and marking which is thus
brought about, and for the difference over the wings which
has been already alluded to.

The dark grey forms (26), by making a similar substitu-

876 Mr. F. Merrifield and Mr. E. B. Poulton on

tion, resemble the dark yellow-green (1b). The purplish-
brown is increased in extent as is the green of the corre-
sponding form (1 0), the dark dorsal band becoming a
specially prominent feature which contrasts strongly
with the appearance presented by the light grey pupe
2 a).

: I will now apply the arrangement suggested above to
the results of Mr. Merrifield’s experiment. The pup were
compared on May 2nd, 1899. Three of the 14 larvee died
without pupating. Of the remaining 11, 6 had been
placed in the cylinders furnished with dark sticks; 5
of these pupated on the dark sticks. In comparing
these summer pupze and putting each in its place in the
scale of colour, great assistance was derived from Mr.
Merrifield’s descriptions made in 1898, when more of the
pup were alive and those which were dead had changed
less extensively.

All the pupz attached to dark sticks were clearly grey
forms (1), except one (probably dead), which was inter-
mediate between yellow-green and grey, having a bone-
coloured ground with many of the markings, especially
upon the wings, greenish instead of brown. Of the remain-
ing 4 pup, 1 had emerged in 1898, and was intermediate
between dark and light grey (2 a, b) or a lightish dark
grey (2 6); 1, still alive, was also a (2 a, 0); 2, dead, were
certainly grey, and probably dark grey (2 6).

The sixth pupa was dead and much discoloured, but from
Mr. Merrifield’s description in 1898, it had evidently been
a yellow-green form (1 a) or (1 0): “The larva had
imperfectly attached itself to a dark stick and had then,
before pupating, fallen on the earth which, being moist,
was of a dark colour. It had there formed a yellow-green
pupa, somewhat misshapen.”

Of the 5 placed in light surroundings, 4 were attached
to shaved white sticks.

Two were dead, but both were evidently yellow-green,
one probably a light and the other a dark form of this
degree, (1 a) and (1 D).

Two had emerged, but both had been yellow-green and
one certainly a light form (1 a), the other probably dark
(1 0).

The fifth pupa, attached to the white muslin top, had
also emerged. This pupa was an exception, being a distinct
light grey form (2 a).

Adjustment of colour in various pupex, etc. 377

3. EXPERIMENTS UPON THE WINTER PuPm OF Papilio
machaon. (F.M.)

In order to continue the experiments, begun upon the
summer pup, I obtained from Germany, from the 31st
of August to the 2nd of September, 1898, about 150 more
larvee, most of them young.

Apparatus.

In order to cope to some extent with the difficulty I had
in taking the larvee just at the right moment, owing to my
long daily absences from home, I provided two special
breeding-cages. These were made of ;5, inch deal,each in two
compartments. Thus there were four compartments the
external dimensions of which were 16 inches in height, 12
inches in depth from front to back, and 8 inches in width.
The front of each cage was a sheet of glass, about 14
inches in width and 12 inches in height, as it started at 3
inches from the bottom; it was vertically divided down
the middle by the thin deal partition between the two
compartments. The backs were of perforated zinc, corre-
sponding in size with the glass fronts, and the outer sides
were of deal containing the doors. The tops were open,
but covered with woven material as stated below. The
framework of the top, and generally, was about an inch
wide, and there were many angles more or less shady,
vertical, and horizontal, where the parts of the framework
met at right angles.

The compartments were covered internally as follows:—
The whole of the interior, except the glass front and the
perforated zinc backs, was covered with tissue-paper ; black
in the black compartment, white in the white compartment,
green in the green compartment, and about one-halt orange
and one-half yellow (the division being vertical) in the
orange-yellow compartment. The perforated zinc backs
and the open tops were covered with woven material, viz.
the black compartment with double black muslin (covered
with a slate), the white compartment with double white
calico, the green compartment with threefold yellow-green
art muslin (fourfold on the top except for the brassicz experi-
ment hereafter described when it was twofold), the orange-
yellow compartment with orange-yellow leno; this leno,
instead of paper, being also used in some of the angles of
the framework.

The fronts of the cages were placed within a few inches

378 Mr. F. Merrifield and Mr. E. B. Poulton on

of a second-floor window about 3 feet wide and 6 feet high,
they were facing the W.N.W. and looking into an open
country, and were often screened by a muslin window-
blind during hot sunshine. No direct light from the sky
reached any part of the interior except a small part of the
bottom and of the sides.

Much orange light came through the roof of the first
compartment, very little greenish light through the top of
the green compartment, and much white lhght through
the top of the white compartment; in addition to white
direct light, much was reflected from the coloured interior
of these three compartments, but the black compartment
was very dark inside except the part close to the glass
front.

The effect was that the black compartment was for the
most part very dark; the white, green and orange-yellow
compartments much lighter, as the tops transmitted much
light, and the colours were such as to reflect a great deal
of light. In the green compartment, however, while
covered with the fourfold art muslin, very little light
came through the top. In all cases a little ght came in
through the chinks of the doors, and a very little through
the draped perforated zinc at the back. In all four com-
partments there were shady regions in the angles of the
framework, and I found so many of the larve had a dis-
position to select these shadier regions for pupating, where
of course the coloured light would operate less strongly,
that I found it expedient to transfer them, when I could
do so im time, to receptacles where they were exposed to
stronger light. For these purposes I prepared glass
cylinders of the dimensions before mentioned, covered at
the bottom and for about two-thirds of the outside cireum-
ference with paper of the appropriate colour, leaving clear
the one-third next the window, and the tops being usually
covered with paper of the same colour, but sometimes with
clear glass. In this way I fitted up cylinders for the follow-
ing colours: black, white, green, yellow, orange, Dutch
“gold,” all provided with sticks covered with paper of the
corresponding colour. These cylinders were placed on thin
pieces of wood or cork carpet with tintacks driven through
the bottom, forming spikes on which the coloured sticks
were fixed. The bottoms, as well as about two-thirds of
the circumference opposite the light, and the tops, were
covered with single or double paper of the proper colour

Adjustinent of colowr in various pupe, etc. 379

secured by three ties of thread and on the tops by a sheet
of clear glass. In the case however of the dark sticks the
bottom was of dark cork carpet, and these cylinders were
clear all round, with a sheet of clear glass on the top.

In order not to crowd the larve, it was my practice to
transfer them, when spun up on the sticks, to wide-mouthed
Bordeaux plum-bottles of clear glass having a greenish hue,
of about the same size as the cylinders, but fitted for two-
thirds of their circumference with coloured paper inside
instead of outside, and I sometimes transferred larvze direct
from the compartments of the breeding-cage to these bottles,
as pupating against glass in front of coloured paper did
not appear to me to be the same thing in effect as pupat-
ing against the coloured paper. In these bottles the
coloured paper was inside, and therefore nearer the larve
than in the cylinders. The sticks in the bottles could not
be prevented from shifting as heavy larve crawled over
them, and the larve appeared to dislike this, and to be
more restless and slower in pupating than when the sticks
were fixed.

The cylinders and bottles were placed within a few
inches of the window above referred to, or of another
window having the same aspect, but only about 4 feet
high by 3 feet wide. As the objects in both were neces-
sarily near the light, and, except in those given over to
black, light, coloured or uncoloured, was admitted all round
and by the tops, the larve in these were as a rule exposed
to much more light, both direct and reflected, than those
in the breeding-cages.

I also lined a clear glass saucer with green carrot-tops
and placed it in a second saucer with green carrot-tops,
among which some larvze pupated, the surface being covered
with a sheet of clear glass.

Most of the machaon larve in the eylinders or bottles
spun up against the coloured sticks provided for them, but
in a few cases they spun either against the glass where it
was covered outside by coloured paper or against the clear
glass front of the cylinder or bottle. In these cases I
always classified the pup as “orange through glass” or
as ‘“‘orange on glass,” the pupz in the latter case being
often not near the special colour, although the larve in
moving about had probably been at times exposed to the
colour influence. If, however, these larvee follow the same
laws as those of Vanessa wrticx, they are only sensitive when

380 Mr. F. Merrifield and Mr. E. B. Poulton on

at rest on the surface upon which they will afterwards
pupate. (Phil. Trans. 1887, /.c.) Allowance must be made,
on the other hand, for the possible disturbance of larve
after they have entered the susceptible phase.

Colours of the different surroundings.

Besides these black or coloured cylinders or bottles I
had clear glass cylinders or bottles supplied with the other
objects enumerated below for the pupze to attach themselves
to, some also being shut up in absolute darkness. The
result was that I obtained several classes of pupe, viz.
from—

1. Black paper (a) in strong light, (>) in more or less
shade.

2. Dark sticks from old faggots or freshly-cut dark
alder.

3. Dirty white paint, being that of a breeding-cage ten
years old.

4. Darkness.

5. Light-coloured dry stems (dead stems of Hpilobiwm
hirsutum).

6. Dead reeds, light brown in colour.

7. Dull green reeds.

8. Bright golden yellow oatstraw.

9. Dutch “gold,” with embossed pattern.

10. Green paper (a) in strong light, (6) in more or less
shade.

11. Green carrot-tops.

12. White paper (a) in strong light, (2) in more or less
shade.

13. Yellow paper (a) in strong light, (>) in more or less
shade.

14. Yellow orange leno (a) in strong light, (>) in shady
corner.

15. Orange paper (a) in strong light, (6) in shady
corner.

Some of these had the further differences which are
indicated in the exhibit and described in the classification
of the pupe.

T exhibit the results of all these exposures, 72 individuals.
All of these may repay study ; the details will be described
by Prof. Poulton. I could perhaps have made the Exhi-
bition Case more attractive as well as more effective for
purposes of comparison had I detached the pupz from the

Adjustment of colour in various pupa, ete. 381

coloured backgrounds that in most cases adhered to them ;
but I thought it eminently desirable that the exhibition
should supply its own evidence, so as to enable all who see
it to draw, independently, their own conclusions. I supple-
ment it, however, with all the other pup obtained in the
course of my experiments. As general results it will be
enough perhaps here to say that the 16 pupe of P.
machaon on the black paper-covered sticks and on dark
natural sticks are all grey, with the exception of one,
which is yellow-green ; those that were in a strong light
are very dark; that of the 4 on carrot-tops all are
yellow-green ; that of the 9 on white paper 1 is grey
and the other 8 all yellow-green or light grey with a
greenish tinge; that of the 6 on yellow paper 2 are
greenish-grey and the other 4 yellow-green ; and that of
the 4 on orange paper all are yellow-green.

Theseresults seem to me clearly to prove the susceptibility
of P. machaon.

The pups of P. machaon on the other coloured sur-
roundings employed did not give such definite results;
particulars are given in the next section.

4. RESULTS OF THE ABOVE EXPERIMENTS. (E. B. P.)

The pup were compared May 2nd, 1899.

The various conditions to which ihe larvee of P. machaon
were subjected will be considered in the same order as that
adopted by Mr. Merrifield on p. 380.

(1) BLACK PAPER. (a) In strong light.

2 pupe, fixed to black tissue-paper, were very dark
grey (2b).

1 pupa, fixed to black tissue-paper, was intermediate
between dark and light grey (2 a, b).

1 pupa, fixed to black tissue-paper, had emerged, and
was grey, probably similar to the last.

1 pupa, fixed to black net, was dead, and was grey,
probably similar to the last.

(b) In fair light.

The single pupa (dead and discoloured) was fixed to
black net. It was impossible to place it with any certainty,
but the appearance suggested that it had been yellow-
green.

382 Mr. F. Merrifield and Mr. E. B. Poulton on

(c) In a dark corner.

All the pupz were fixed to the black paper.
1 pupa was very dark grey (2b).
3 pup (one dead) were grey, probably light (2 a).
1 pupa was grey, probably intermediate between dark
and light (2a, b).

5

}
}
7
1

(2) DARK STICKS (ONE MUCH LIGHTER THAN THE REST).

All the pups were attached to the sticks.
2 pups were dark grey, (2b) one very dark.
1 pupa, on the lightest bark, was light grey (2 a).
2 pup (dead) were grey, probably light (2 a).
1 pupa (dead) and much altered, was probably the
yellow-green form (1) described by Mr. Merrifield
on p. 381.

ic

(3) DIRTY WHITE PAINT.

1 pupa, fixed to the painted surface, was intermediate
between dark and light grey (2 a, b), but with
some tendency, in the possession of a greenish
tinge, towards the yellow-green form (1).

1 pupa, fixed to the painted surface, was dead and
changed in colour, but it had probably been
yellow-green (1).

ro!

(4) DARKNESS.

2 pupze were intermediate between dark and light
yellow-green (1 a, b).

1 pupa (dead) had altered in colour, but had been a
light form of either (1) or (2).

1 pupa was probably light yellow-green (1 a), but had
darkened, apparently preparatory to emergence.

1 pupa was intermediate between (1) and (2) and also
intermediate between dark and light (a, 0).

Hox!

The absence of strong pigmentation, and the tendency
towards yellow-green rather than grey, were the marked
results of darkness, so far as these 5 pupz are concerned.

Adjustment of colour in various pupe, etc. 383

(5) LIGHT-COLOURED DRY STEMS (Epilobium hirsutwm),.

1 pupa (dead and discoloured) was probably light grey
1 pupa (dead, or perhaps discoloured, preparatory to
emergence) was probably a yellow-green form (1).

re)

(6) DEAD, LIGHT BROWN REEDS.

Both pups had become greatly discoloured, and one at
least was dead. The position of one was uncertain; the
other had apparently been a yellow-green form (1).

(7) DULL GREEN REEDS.

1 pupa, fixed to reed, was intermediate between dark
and light grey (2a, b).

1 pupa, fixed to reed, was intermediate between grey
and yellow-green (1, 2).

2
(8) OATSTRAW (BRIGHT GOLDEN YELLOW).

1 pupa, fixed to straw, was a light grey form (2 a), with
some tendency towards yellow-green.

1 pupa, fixed to straw (dead and discoloured) was of
uncertain position, but clearly it had not been
highly pigmented.

pre)

(9) DUTCH “GOLD” (EMBOSSED).

—

pupa, fixed to “gold,” was intermediate between
dark and light grey (2 a,b), with a slight tendency
towards the yellow-green form (1).

pupa, fixed to “gold” (dead), was intermediate
between dark and light yellow-green (1 a, b).

ft

be |

(10) GREEN PAPER. (a) Ln strong light.

1 pupa was dark yellow-green (1 6}.

1 pupa was light grey (24).

1 pupa was dead and of uncertain position, but pro-
bably it had never been strongly pigmented.

384 Mr. F. Merrifield and Mr. E. B. Poulton on

(b) In good light.

The single pupa was dead and discoloured, and its posi-
tion uncertain. It was obvious however that it had not
been strongly pigmented.

(¢) In fair light.

1 pupa was intermediate between light yellow-green
and light grey (1a, 2 a).

1 pupa (dead or emerging) was probably a yedlow-
green form (1).

Hel

(ad) In shady corner.
1 pupa was light grey (2 a).
1 pupa was intermediate between /ight and dark grey
(2 a, 6).

(e) Fixed to glass near the green paper.

The single pupa was dead, and so discoloured that it
could not be placed.

Hence green paper is far less powerful than yellow or
orange paper (Nos. 13, 14, 15), or the natural green of
chlorophyll (No. 11) in the production of green pupz of
this species, thus agreeing with the results already obtained
in the case of other species (Phil. Trans. 1887, and Trans.
Ent. Soc. 1892, J. ¢.). It is noteworthy that the most
shaded part of the green surface produced the strongest
tendency towards grey forms.

(11) GREEN CARROT-TOPS,
1 pupa was intermediate between dark and light
yellow-green (1 a, b).
3 pup (dead or emerging) were yellow-green, probably
of the same shade as the above (1 a, 0).

| =|

(12) WHITE PAPER. (a) In strong light.

4 pupe were light yellow-green (1 a), one of them
with a tendency towards grey (2), and one very
remarkable in possessing an almost white ground.

1 pupa was intermediate between light and dark
yellow-green (1 a, b).

Adjustment of colour in various pupe, ete. 385

1 pupa was discoloured, but had probably been inter-
mediate between a light and dark grey (2 a, b).

I co |

The form with white ground perhaps indicates some
special influence of the surroundings in the direction of
producing a peculiarly close resemblance; but more
experiments are needed in order to render it certain that
the case was not that of a rare individual peculiarity.

(b) In fair light.
1 pupa was intermediate between light and dark
yellow-green (1 a, b).
1 pupa was light yellow-green (1 a), with a tendency
___ towards bone-colour in the ground (2).
2
7 (c) In dark corner.
The single pupa was discoloured, but was probably
light yellow-green (1 a), with some tendency towards
grey (2).
(13) YELLOW PAPER. (a) Jn strong light.

pupa was dark yellow-green (1 5).

pupz were yellow-green (1), discoloured, but pro-
bably light (1 a), or intermediate between light
and dark (1 a, 0).

pupe were light yellow-green (1 a), with some
tendency towards grey.

bo (IL) |

lo!

(b) Through glass.
The single pupa was dead, and had entirely blackened.
(c) On glass.
The single pupa was discoloured, but was probably
yellow-green, and certainly had not been highly pigmented.
(14) YELLOW-ORANGE LENO.

This label could not be found as distinct from the yellow
(13) and orange (14); or I may have overlooked it.
The results are certainly included in one or more of the
sub-divisions of (13) or (14), inasmuch as the total number
of the pup examined is 72,—the number given by Mr.

Merrifield.
TRANS. ENT. SOC. LOND. 1899.—PART IV. (DEC.) 26

886 Mr. F. Merrifield and Mr. E. B. Poulton on

(15) ORANGE PAPER. (a) Jn strong light.
3 pupe were light yellow-green (1 a).
1 pupa was dark yellow-green (1 b).

\ >!

(b) Through glass.

The single pupa was dead or emerging, but appeared
to have been intermediate between dark and light grey
(2 a, 6).

(c) In shady corner.
2 pupe were light yellow-green (1 a).
1 pupa was light grey (2 a).
3

As Mr. Merrifield has stated on p. 381, the examination
of these pup seems “clearly to prove the susceptibility of
P. machaon.” At the same time there was one unsatis-
factory point in the evidence, viz. the extremely unhealthy
condition of the pupe. A large solitary ichneumon began
to emerge from the pupz in the autumn of 1898, soon
after the date at which Mr. Merrifield showed them before
this Society (October 5th, 1898). At intervals they con-
tinued to emerge until the early summer; and soon after
my examination was made on May 2nd, 1899, they
came out suddenly in large numbers. The ichneumons
almost invariably gnawed a hole in the pupal wing, in
order to escape. The whole batch produced very few
butterflies.

It was unfortunate that press of work prevented me from
comparing the pupze until so late, when the discoloration
of many of them had proceeded so far; but under any
circumstances it was much to be desired that the
susceptibility of undoubtedly healthy pupz should be
tested. Such a test I have fortunately been able to apply,
and the result confirms Mr. Merrifield’s conclusion in the
most complete and convincing manner, as may be seen
from the succeeding section.

5. Mr.C. V. A. PEEL’s WINTER PupP& oF Papilio machaon
DESCRIBED. (KE. B. P.)

Through the kimdness of Mr. Peel I have been able to

compare a fine set of very healthy pup from Wicken

Fen. The pupz were in part attached to green reeds; m

Adjustment of colour in various pupx, etc. 387

part to the wood (somewhat darkened by age), and in
part to the perforated zinc of two ordinary rectangular
breeding-cages. It is clear that the two latter sets of
pupze had been formed from captured larvee, while the set
attached to reeds may have been, in part at least, found
so attached in the open. Mr. Peel is not sure upon the
point; but the fact that reeds, and reeds only, have been
selected is in favour of this interpretation. The majority
of the pupz have now emerged successfully, and nearly all
of those that remain are still healthy. Very few have
died. The entire absence of parasites in this set of larve,
as compared with their excessive abundance in Mr.
Merrifield’s continental individuals is of high interest, and
suggests the same conclusion as that at which I arrived
last year in breeding large numbers of Continental and
English larve of Vanessa urtice, viz. that the greater
abundance of birds in this country may, by destroying
parasites, compensate for their direct attacks on the
species of Lepidoptera (see Report of British Association,
1898, Section D). I will now give the results of the
examination which was made on May 3rd, 1899.

14 pupex attached to reeds,

Thirteen pupz were distinct light yellow-green (1 a).
Of these 2 were fixed at some distance from each other
on the same reed, while 10 were on separate reeds. Just
below the thirteenth pupa was fixed the single exception
a dark grey pupa (2 b).

19 pups upon wood or zine.

In the breeding-cage which contained th2 reeds, one
dark grey (2 b) pupa was lying loose upon the floor, and
another of the same kind (2 }) was attached to the zine.

In another breeding-cage 5 pupze were attached to the
z ne, in a curved line, near together. From above down-
wards their arrangement was—

2 pup intermediate between dark and light grey
(2 a, b).

1 pupa dark grey (2 b).

1 ,, again intermediate (2 a, 0).

1 ,, light grey (2 a).

Thus the central pupa was durkest, as though some
influence had been exerted by the neighbouring larvee or
pupie (as is so markedly the case in Vanessa).

888 Mr. F. Merrifield and Mr. E. B. Poulton on

Above these 5 pupz, on the wooden roof, were 3 pupe,
two close together and one near them in the corner. All
were light grey (2 a).

At the opposite end of the cage 3 pupz were scattered
over the zinc, one being intermediate between dark and
light grey (2 a, b), two, dark grey (2 b).

On the wooden roof over them was a compact group of
5 pupe, all imtermediate between dark and light grey
(2 a, 6); while a single pupa loose on the floor was also
intermediate (2 a, 0).

Thus there was not a single exception among the pup
on the zinc and wood. Furthermore those fixed to the
latter were on the whole lighter than those fixed to the
darker zinc. The singular completeness of the result is
best shown in a tabular form as follows :—

Yellow-green (1) Grey (2)

Light (a) | Dark (6) | Light (a) | Dark (6)

Attached to green reeds 13 _ = 1 = ile

Intermediate.

Attached to wood of cage — — | 8 5 = |= 8

Lying loose on wooden floor
(more in shade than roof
and sides, and with dark:

débris scattered over it) & — — — 1 1 = 2
Attached to zine — == 1 4 4 —9
33

It is unnecessary to examine these data further. It is
obvious on an inspection of the above table that there
is only a single exception to the complete susceptibility
of the pupe.

The much greater susceptibility of this set of pupze as
compared with Mr. Merrifield’s considered as a whole, and
with the few upon which I experimented in 1886 (Phil.
Trans. 1887, /.c. p. 406), is probably due to their more
healthy and vigorous condition, and perhaps in part to
some of the results having been obtained under normal
conditions (if it is admitted that most of the green pupe
were formed in the open). The larve are not gregarious,
so there is no justification for assuming a family tendency
towards susceptibility on the part of the set as a whole.

Adjustment of colour in various pupx, ete. 389

Local differences in susceptibility are of course possible,
and an enquiry directed along this line might lead to
results of high interest.

It is a great pleasure to me to see this species, upon
which so much doubt has been thrown—in the first place
in the discussion which followed Mr. T. W. Wood’s
communication to this Society in 1867 (Proc. pp. xcix—
ci), and nineteen years later as the results of my experi-
ments—now finally proved beyond doubt to be susceptible
to the colours of its environment. This result, which we
owe to Mr. Merrifield, is a further warning against the
errors into which we are liable to be led by relying, as Mr.
Bond did (in the 1867 discussion), upon a general im-
pression gathered from a wide experience not specially
directed towards the solution of the problem, as I did,
upon an insufficient number of individuals subjected to
experiment.

0.—EXPERIMENTS UPON THE PUPA OF
Papilio podalirius. (Cora B. SANDERS and E. B. P.)

Five full-fed larve of this species were found by us
in Switzerland, between Visp and Stalden, on July 22nd,
1898. They were all, except one, of the usual yellow-
striped green form. The single exception was brownish-
green with many red spots somewhat similar to those
which occur upon certain forms of the larvae of Smerinthus
ocellatus and S. popult. It is possible that the darkened
ground colour was due to changes preparatory to pupa-
tion, or perhaps to ill-health, as the larva died without
pupating. Two of the larva were placed in a white
muslin bag and offered green reeds together with the
green twigs and leaves of the food-plant: three were
placed in a black net bag, and provided with dark brown
branches, as well as the food-plant. In a few days four
of them pupated, one being fixed to the white muslin
and three to the black net.

It was immediately seen that the latter were far darker
than the former. A careful comparison was made on
May 3rd, 1899.

The pupa which had been fixed to white muslin was a
pale dull orange tint, especially dull over the wings. Of
the other three which had been fixed to black net, one
was also dull orange but of a distinctly darker shade,

390 Mr. F. Merrifield and Mr. E. B. Poulton on

while the remaining two were much darker still, being of
a purplish-brown deepest in tint over the wings.

These small numbers are not sufficient to prove the
susceptibility of this species; but they render such
susceptibility probable. It is to be hoped that larger and
more varied experiments will be made by those who have
the opportunity of obtaining considerable numbers of the
larvee of P. podalirius.

D.—EXPERIMENTS UPON THE PUPA OF

Pieris nap.

1, EXPERIMENTS UPON THE WINTER Pup oF Preris napi.
(F. M.)

I exhibited to this Society on November 2nd, 1892
(Proceedings 1892, p. xxx), some pup of P. napi show-
ing that the species was susceptible. In this present year
I was experimenting on the species for other purposes, and
determined to avail myself of the apparatus I had to
provide for experiments on the coloration of pupz of P.
machaon. Mr. Harwood supplied me in the early part of
August with a number of females captured in the vicinity
of Colchester, and from them I obtained several hundred
eggs on watercress, on which I fed the larve till about
their last stage when that food was largely supplemented
and finally replaced by cabbage. These when approach-
ing pupation were exposed to the same colour influences
as the machaon larve had been (substituting green
cabbage-leaves for carrot-tops), and to the following in
addition (16) planed deal in shade, (17) planed deal in
light.

They seemed very unwilling to pupate on orange
(except on the glass in front of the orange paper), so I
shut up some in (18) a threefold yellow-orange leno
cylindrical bag with single leno on the top, and orange
paper outside two-thirds of the circumference, and here
they were obliged to pupate.

In this species and in P. brassice my results are too
numerous for me to show the whole in the Exhibition
Case; but I have brought with me in glass-bottomed
boxes, so that they can be seen, all the pupz of both
species which are not thus displayed Gn the Exhibition
Case), duly classified. There are in all about 340 pupz of
P. nap.

Adjustment of colour in various pupe, ete. 391

About 80 of the napi not in the general Exhibition Case
are in a second Exhibition Case under the following circum-
stances. I had an old breeding-cage (somewhat resembling
those previously described, but a little smaller) in two com-
partments with a glass roof over both; one of these was
lined with black and had its glass top covered with double
black tissue-paper, nearly opaque; the other was lined
partly with orange paper, and partly with yellow paper or
yellow-orange leno, and the glass top was covered with three-
fold orange Teno, transmitting much orange light. Finding
that a number pupated on the glass roof, I succeeded in
detaching the plate of glass from the roof and mounting
it for exhibition in a second case. It will be seen that on
the black side there are 34 pupe all bone-coloured and
nearly all much spotted with dark; on the orange yellow
side 46 pupe, all green except 4 which are bone- -coloured,
though with a yellowish tinge, and nearly all the 46
practically unspotted.

I think this second Exhibition Case, in which the pupz
have arranged themselves, atiords a very effective demon-
stration of the sensitiveness of this species.

The following section will contain a detailed statement
of results by Prof. Poulton. Here it will be sufficient in
reference to the first Exhibition Case to point to the contrast
between the 10 on black paper or the 8 on black or
dark sticks, all of which are dusky, with much black
spotting, and the 19 on yellow or orange paper all of
which, with one exception, are green, to the 12 on or
near cabbage-leaves, many being attached to the glass
bottle, most of which are green, and to the varying colour
of those in darkness; the 8 on Dutch gold, the 7 on
planed deal, the 8 on green and the 6 on white paper
being nearly all bone-coloured, and most of them spotless
or nearly so. These results clearly prove the high
susceptibility of the pupa.

All the four species experimented on by me, in pre-
paring to pupate vary in colour according to their
surroundings, and I do not see any room to doubt, when
the 16 pupz of machaon on black paper or dark sticks are
compared with the 4 on green carrot-tops,*or when the
18 pups of napi on black paper or dark sticks are
compared with the 12 which pupated on or near cabbage-
leaves, that the adaptation must be in many cases
protective.

392 Mr. F. Merrifield and Mr. E. B. Poulton on

2. RESULTS OF THE ABOVE EXPERIMENTS. (E. B. P.)

The pup of Pieris napi were compared on January
2nd, 1899, with a view to the construction of a standard
table.

As compared with the allied P. rapa, the dimorphism of
the ground-colour is far more marked. Furthermore, the
ground is almost invariably restricted to bone-colour or
green in P. napi, whereas many different shades are
common in P. rape. Even in pupe with the darkest mark-
ings the bone-coloured ground is far less obscured by
generally distributed pigment in xapi than it is in rape.
There is also in the former a marked dimorphism in the
arrangement of the black markings, which show character-
istic differences even when present in similar amounts in
the two forms—bone-coloured and green. Thus the bone-
coloured forms, however pale, almost always possess a
distinct black patch (made up of two or three spots or
short lines) in the centre of the fore wg. This character,
which I call the “ wing-mark,” is either wanting or far
less developed in the green forms, even when more
richly pigmented in other parts of the surface.

Another difference is the far greater irritability of the
pupe of P. napi. A slight stimulus, such as light breath,
would almost always cause active movements, when the
more stolid pup of P. rape remained quiescent. This
observation, which certainly held at the time the examin-
ation was made, may perhaps have been due to differences
in the degree of development then reached by the two
species.

The forms of the two pup are almost alike, and as each
varies considerably, it is very difficult to distinguish them
with certainty by this means; but the above-mentioned
tests enabled me easily to pick out two pupz of P. rape,
which had found their place among Mr. Merrifield’s numer-
ous napi, having been accidentally introduced as larvee in
the food-plant.

The pup of P. napi are either green or have a bone-
colowred ground with black markings and minute dots. The
latter may be classified, according to the amount of
pigment, as :—

1) Dark, (2) Intermediate, (3) Light. In the light
forms the pigment is so small in amount that the pupz

Adjustment of colour vn various pupe, ete. 393
y)

are practically bone-coloured (3a). The green pup are
equally deficient in pigment and may be regarded as a
dimorphic form of this degree (3). These relationships
may be conveniently expressed as follows :—

1 ADARK.
9, INTERMEDIATE.
3. LIGHT. (a) Bone-coloured.

(b) Green.

(1) The Dark pupe. The ground is bone-coloured.
Black pigment is strongly developed on each side of the
dorsal surface, forming a large sub-rectangular patch on
each side of each abdominal segment, fusing into an
irregular mass anteriorly, on each side of the thoracic
region. The dorsal line is marked posteriorly by a distinct
dot on each segment. Seven dots can be recognized,
including one on the caudal spine. There is also a large
pigment patch on each side of the mesothoracic keel and
on each side of the anterior rostrum. Minute black points
are scattered between the sub-dorsal pigment and that of
the dorsal line.

On the sides, the principal development of pigment is on
the wings, in which part of the venation is thus rendered
conspicuous, while the outline of the hind margin of the
future wing is marked by a distinct row of black dots.
Near the centre of the wing is the ‘“ wing-mark” made
up of two or more, generally three, intensely black patches
—the largest inferior apparently occupying spaces
between the veins. Even in the darkest forms these
patches are nearly always conspicuous from their superior
blackness: they often tend to fuse, forming in many cases
a single large patch.

The eye is strongly pigmented superiorly, but below and
including the crescentic mark upon which alone of the
entire pupal surface the facetted structure is developed
(the pupal eye), it is devoid of pigment.

(2) The Jntermediate pupe. The ground is_bone-
coloured. These pup differ from the last, with which
they are connected by transitional forms, in the lesser
development of black pigment both dorsally and on the
wings. Asa rule the diminution is proportional throughout,
but in certain cases the dorsal pigment may retain its
full development.

The lesser amount of black pigment renders the intense

394 Mr. F. Merrifield and Mr. E. B. Poulton on

black patches in the centre of the wing especially distinct
and sharp.

(3) The Light pupee. (a) Bone-colowred. 'These also are
perfectly transitional into the Intermediate pupe. In
pupee of this degree the pigment is everywhere reduced,
remaining strongest in the black patches in the centre of
the wing, which become excessively conspicuous against
the pale bone-coloured ground. In some of the most
extreme cases these patches become much fainter, but a
trace of them is probably always to be found. In other
parts the pigment spots and patches are much smaller,
and are often represented by minute dots: on the eye
it is often absent altogether.

(3) The Light pups. (b) Green. Pigment is never
highly developed upon the ground-colour of these pup.
With very rare exceptions it does not exceed the amount
present on the light bone-colowred pups (3 a). The green
pupee are far more transparent than the bone-colowred, and
the palest are even more deficient in pigment than the
palest of the bone-colowred forms. 'The most important
difference in marking has already been mentioned, viz.
the absence of the black patches in the centre of the wing
(the “ wing-mark”’) even when pigment is developed else-
where as greatly as in a decidedly dark bone-colowred pup
of the corresponding degree (3 a@)—a pupa which would
always possess distinct and prominent patches. Occasion-

ally, however, faint traces of the marking may be detected,

as minute dots, even in the palest green forms, and very
rarely it is fully developed. All marked exceptions will
be described below. In the centre of the wing the trans-
parency is such that a considerable depth into the pupa
can be seen, and the large tracheze distinctly made out.

Before classifying Mr. Merrifield’s numerous pupe, I am
tempted to suggest what I believe to be the meaning of
this strange dimorphism in marking. The possible failure
of my hypothesis would not, however, alter the validity
of the observations which I have here recorded. Without
attaching too great weight to it, I do not hesitate to
suggest the hypothesis, thinking it possible that observa-
tion and thought may be stimulated by its means.

I have just alluded to the great transparency of the
green pup (3 4), and have stated that this is especially
marked in the centre of the wing, viz. in the exact
position of the dark mark on the bone-coloured pupe (3 a).

Adjustment of colour in various pupe, ete. 395

These latter are also, though less transparent than the
green, chiefly so in the same area,

The green pup resemble green leaves and stems, and
Mr. Merrifield’s experiments show that they are produced
by such surroundings: the bone-coloured pup resemble,
and are produced by, such surfaces as bark, wood, or stone.

Now the transparency is no hindrance to the conceal-
ment of the former: it is rather an advantage. But to
the latter it is a distinct hindrance and, when once seen,
immediately betrays the fact that the pupa is not the
opaque object to which it presents so strong a superficial
resemblance. I believe that this is the reason why this
particular area is so invariably covered up by dark patches
of pigment in the pale bone-coloured pupee of this species.
I believe that the persistence of this particular mark
when the other pigment spots and masses are disappearing
in the palest pupz, is due to the operation of natural
selection.

The experiments and observations Miss Cora B.
Sanders and I were able to conduct, in the summer of last
year, prove that pupze (in this case of Vanessa urticx) are
subject to a tremendous struggle for life: they also
strongly indicate that the enemies are guided by their
sight in hunting for them. Hence it appears to me that
there is nothing improbable about the suggestion that this
dark patch covering the transparent area may have been
retained by natural selection in certain forms, because
transparency would be a danger, may have been dismissed
in certain others, because transparency would be an
advantage,

The pupz were compared and tabulated on Dec. 22nd,
1898, and on the following dates in 1899—Jan. 6th, 7th,
9th,and 28th. Nearly all of them were extremely healthy,
and showed the effects of the conditions in a remarkable
manner. As these are the first complete and detailed ex-
periments which have been conducted upon this sensitive
species, and the material was in such excellent condition,
the comparison was carried out with the utmost care,
and numerous details were recorded which are included
in the tabular statement below. The order is that in
which the pups of P. machaon were considered so far as
the conditions were the same. The conditions which
were different are placed at the end.

It is to be observed that the light-green forms (3 b)

396 Mr. F. Merrifield and Mr. E. B. Poulton on
Degrees of Pupal Colour.
|= | Light (3)
EXPERIMENTS. SUAS A edly 4 R ks.
EIS) ES 23 aS g emarks.
eres aa iees
La! [S)
o
1. Black. aman eee 10 | 5 Three of the (2)s very dark.
2. Black compartment) 15 | 41 | 1 17 | (2) Very dark : (3 a) very
(pupe fixed to black). | light. Also two dead pup
which were probably (1) or
(1) and (2).
3. Miscellaneous black il ree 1
/ compartment.
4. Black through glass it il Very pale pupa.
(in shade).
| 5. Black compartment, | 9 if 3 The (3 a) darkish.
through glass.
|= SS S| —
244 Anterior group} 5 | 6] 6 '17 | Pupe mostly fixed parallel
52 & >|mostly close to | to glass front: the heads of
S° © s\clearglass front. | ‘only five pointed towards
\e¢ ox light, and only two of these
him i=} (ol 5, o
ee Soar pointed directly (the body
obs & line at right angles to glass
Soc | front).
SS |
“ = ae) | |
g Oy fe 3, Group pelea) 1 il | | 2 One pupa pointing to-
= EE B|between bac | wards light; one had be-
22 =aiand front, and | come loose.
03,2 S)towards inner| |
= =%e)side (viz. near-|
eR. = g \est orange).
2B Gam | :
2 oat =| Group in pos-| 1 | 1 | | 9 | Another pupa had died
® § £./terior inner cor- | andisnot classified: another
© S49 | ner of roof. lesion was accidentally mixed
So ae | with next group and is
4255 | there classified. All four
5 2 = | | towards light (direct).
» | Group in poste-| 10 | 2 | | 12 | Including one pupa from
= © & rior outer corner | last-named group. All
&,_ @\and extending | pup towards light and all
z= = along outer side direct except one.
= 5 = of roof. | a fo
7. Black compartment | CANT ae 14 | One (3a) rather dark with
on or near front glass | }an indistinct wing-patch ;
another much lighter with
a faint one. In addition,
an extraordinary intermedi-
ate form not tabulated—a
pale green ground with the
pigment of a (2).

Adjustment. of colour in various pupa, ete.

| Degrees of Pupal Colour.

EXPERIMENTS.

(2)

Intermediate
Bone-
coloured (a)

Light (3)

Green
()

Totals

Remarks.

8. Dark sticks.

Another pupa had died.
The (3 a) was dark.

9. Dark sticks: pupze
fixed to bottle.

The (8 6) very pale and
pigmentless.

10. Dirty white paint. |

9 | 15

bo

11. Glass of old cage.

12. Darkness.

13. Dead reeds.

14. Dead
on glass.

reeds ; pup

15. Dull green reeds.

dl

~I

Two had died and are
unclassified. It is certain
that neither was green. One
pupa was withdrawn as a
P. rape. One (3 6) dark
but no wing-mark to be
detected.

All light for their degrees
except one (3 a).

Of the (3 5)s two are deep
green with much pigment
for this degree, two similar
with little pigment, two
pale and almost pigment-
less.

Wing-mark faint in three
(3a) ; two of them dark for
this degree.

(36) deep green with little
pigment.

Sr
a

(3 6) pigment very dark,
equal to darkest (3) or
even light (2) : wing-mark
small but dark. One (3 a)
with very slight wing-
mark. The (2)s light.

16. Dull green reeds ;
pup on glass.

Another pupa dead.
Wing-mark indistinct in one.

| 17. Oat-straw:

On

(3 @) wing-mark indistinet
in two. (35) one pale, one
deep green, both with very
little pigment.

18. Oat-straw ; pup on
| bottle.

to

Pup with little pigment
and wing-marks indistinct.

397

» | Light (3) |
ene ea]
| ws i SS a; L
EXPERIMENTS. |fo|/8&S|/eg/S-| 8 Remarks.
Ja |e |SE/ES| 2
7 ie ce Mie
| 19. Dutch ‘‘gold” on | 8 8 | Six pupe with very little
_ sticks. : | pigment. Wing-mark very
faint in two, invisible in!
one.

20. Dutch « gold” | 1 1 | Deep green; little pig-

through gla s. ment. |

21. Dutch ‘‘gold” on 1 1 | Almost pigmentless except

glass. for very distinct, dark wing-
mark.
. 22. Green paper spills. ii 7 Another pupa dead.
EADS 2A : pup :
| Five pup with very little
pigment, and wing-mark
| indistinct, oreven invisible.
23. Green paper, through il 1 | Very little pigment.
| glass. |

24, Green paper, on | 1|} 1] Very little pigment |

glass. | deep green. |

25. Green cabbage-leaves ; 4| 6|10) Another pupa dead: (3a)s_

pupe chiefly on bottle. | very light, but with wing-
mark conspicuous. (3 b)s
pale green except one, all
with little pigment.

26. Green cabbage-leaves ; 1 1 | Very pae pupa with

puja on white muslin. faintly transparent yellow-
ish appearance. |

27. White paper. 6 6 | Very little pigment ;
wing-mark distinct and
dark on three, indistinct on
one, hardly visible on two.

28. White paper, through | 1 1 | Almost pigmentless.

glass.

29. Yellow paper. 1} 7| 8 | Nearly all pupe almost |
pigmentless, The (3 a)|
transparent yellowish and

really transitional to (3 3d).
Of the latter, two pale,
two intermediate, and three

Mr. F. Merrifield and Mr. E. B. Poulton on

Degrees of Pupal Colour.

|

deep green.

Adjustment of colour in various pupe, cte. 399

| Dexrees of Pupal Colour.

> | Light @) |
By) 9)
= | ©
ad So
EXPERIMENTS. Tei) micailli te ar a Remarks.
|AS| ES | coal ae ly oes
= ® Baelos iS)
~ os /Hw~ =|
a |mola
ase |
| = | |
30. Yellow paper, through | 1) 1) 2) The (3 a) is transparent |
glass. | | | | _greenish-white, and transt-
tional to (36). The latter
|deep green with much

| pigment for this degree.

31. Yellow paper, on 1 1 Like the (3 a) described
glass. | | -above and, like it, very
| | | _ pale and pigmentless.
32. Orange leno bag. | 11 | 11. ‘Twopale, nine deep green.
One of former fixed to a
leaf. Seven with very
little pigment and the
| | | others not very dark.
38. Orange-yellow com- | | 8 | 8 Another pupa dead. One
partment ; miscellaneous | j | | pale, two intermediate, five
pupe. | | deep green. All very pig-

| mentlessexcept three of lat-
_ter (not very dark), and one
remarkable deep green form
with the pigment of a (2)
and a small distinct wing-
| |mark ; also present on two
of the last-mentioned set of

| | three.
34. Orange compartment, [ot Ny See sei (Suey) very pale and pig-
near leno and dirty white mentless with no wing-
paint. mark. (3 0) one pale, two

deep green, all dark pig-
mented ; faint trace of
wing-mark on one.

|

oes, Anterior ie fel) |S) We vive pupe parallel to
> | :

g ed group, many glass front (four close to
4 close to clear glass | it), one with tail, seven)
& front. with head pointing directly |
Fo

to light : one with head to.
light, but body line not
direct (viz. forming angle
of about 45° with front). |
| (3 a) very pale, trans-
parent, and pigmentless,
much more so than any on
| the black side of same
sheet. One (3 6) pale)

jo yey Moped asurig “cg

ey} uo ednd ayy,

400

Mr. F. Merrifield and Mr. E. B. Poulton on

Degrees of Pupal Colour.

| 2 | Light (@) |

wale | S|. |
EXPERIMENTS. | Pom Eel oq |e | “ Remarks.
9) Bee ie AS) ae
[0S ee Sa oe
| = | iS | |
| |
ta | yellowish-green, and really

a | 'transitional to (3 a). It
22 | and five others almost pig-

nest | ‘mentless, six with very
oe | little pigment, and one

2 8. | with much, like a dark

(e= | (3a) and yet hardly a trace

Se of wing-mark. Twelve of
03, the (3 0) distinct green
eS | | ground.

aos | [sacra Tp yea ariece

= Compact groupin | Viet | 7| 8] Headsof all pupz directly
= | the outer side of | point to light. The (3 a)
5 the middle part of | very pale, but more pig-
‘<4 | roof (viz. away | | mented than any green
= | from the black). | | | | (8 6) in this group and
gs | with traces of wing-
a | | mark. (3 4) all distinct
= | green and very little pig-
= | | ment.
o. | asi
=| Group in the 1 8 9) Another pupa dead. All
S | inner side of the | ie 4 heads pointed directly to
o | middle part of roof) | light. (3 a) with very
F | (towardsthe black). | | little pigment. Seven
~ | | \(3 6) similar, and one
og | darker, but no trace of
ag | wing-mark. All (38) dis-
= tinct green.
ea ee ee
=| Elongated group | 1313 All directly faced light
._ | along outer side of | except one parallel with
= | posterior part of front and two reversed
% | roof. | | (tails directly pointing to
= | | light). All distinct green,
g and, except three, with
_very little pigment.
a | |
= Inner side of 1| 1 Distinct green, very littl
gg | posterior part of pigment. Directly faced
* | roof. | | light.

36. Orange, through | 3| 3 | Another pupa dead. One
ee | pale, two deep green; all
c=) r

very pigmentless.

Adjustment of colour in various pupex, ete.

EXPERIMENTS.

37. Orange compart-
ment ; leno through glass.

Degrees of Pupal Colour.

=>

(2)

Bone-

Intermediate
coloured (a

|
|

bo

]

Light (3)

Totals

Remarks.

(3 a) very pale and pig-

mentless. (3 5) one pale,
two deep green, one of

‘latter with medium pig-

ment, other two very

pigmentless.

38. Orange-yellow com- |
partment, on or near
front glass.

39. Planed deal, in light.

40. Planed
shade.

deal, in

41. Planed
glass lid.

deal,

on

| transitional to (3 b).

Another pupa dead. (2)
very pale. (3a) ten pupe
very pale and pigmentless
with wing-mark minute,
faint, or absent ; of these
two are greenish and
Six
are light with wing-mark
prominent. Three rather
darker, but only average
for this degree. (3 0) one
pupa pale, five deep green.
Two almost pigmentless
(including pale one): four
rather dark for this degree
and minute wing-mark on
two.

(2) very pale. Very little

pigment on (3 a)s and very
faint wing-mark on one:
prominent on others.

Four with very little
pigment, but wing-mark
distinct : one typical.

Very pale and pigment-

less, but wing-mark dis-

| tinet.

which presented any trace of a “ wing-mark ” are specially
so described: when no reference is made to this character
it may be assumed that it was absent in (3 0)s, normal in

(3 a)s.

I will now briefly summarise the results obtained with

this highly interesting and sensitive pupa.

For the sake

of brevity (8 a)s are called light pup; (3b)s green pupe.

Black (1 to 7).

The very powerful effect of black in

TRANS. ENT. SOC, LOND. 1899.—PART IV. (DEC.) 27

401

402 Mr. F. Merrifield and Mr. E. B. Poulton on

producing the darkest pupz is very clearly seen. The
black paper did not act quite as strongly through the
thickness of glass (4, 5, 6) as when the pupz were in
direct contact with it. When the pupz were upon the
class window at a varying distance from the black paper,
the effects were much less marked (7). Black in shade (4)
produced an intermediate pupa, but the parts of the roof of
case(6) which were farthest removed from the light produced
pup which were rather darker than those nearer to it.

Dark sticks (8) produced as great an effect as black, but
little or no influence was exerted at a distance (9).

Dirty white paint (10) gave rise to a great variety of
pupe, all forms being represented among the thirty-one
forms tabulated, the dark and intermediate degrees (1) and
(2) strongly predominating.

Clear glass (11) produced intermediate and light pupe,
while in Darkness (12) they were strongly green—six out of
seven being (3 0). This result is so remarkable and
extreme an effect of darkness that further experiments are
to be desired. It is probably to be explained by the
strong normal tendency of this species to produce green
forms in the absence of any effective stimulus (see 9, 14,
24, in support of this: the pupz on the glass removed
from the stimulus are greener than those subjected to it).

Dead reeds (13) produced intermediate and light pupe ;
at a distance (14) light and green ones were formed.

Dull green reeds (15) produced chiefly light pup (3 a)
with some intermediate (2) and a single green one (3 0);
on the glass they were all /ight (16).

Oat-straw (17) caused the pupz to become light (3 a) and
green (3 b), while at a distance they were light (18).

Duich “ gold” (19, 20, 21) tended strongly towards light
pupz (8 a), the only exception being in the case of a green
pupa (3 6) formed when the gilt ‘surface was the other
side of glass (20). Ata distance (21) the single pupa was
similar to those fixed on the gilt.

Green paper (22, 23, 24) produced almost exclusively
light pupze (3 a), the action through glass (23) being similar
to that of the coloured surface itself. At a distance a
single green pupa (38 6) was formed.

Green cabbage-leaves (25, 26) on the other hand produced
far more green than light pupe, affording a most interesting
comparison with the artificial colour. On this point see
the Appendix in which the constitution of the light

Adjustment of colour in various pupe, etc. 403

reflected from the latter is described. The pupa on white
muslin (26) was light (3 a), and probably affected by this
surface rather than the leaves.

White paper (27, 28) is of great interest, invariably
producing light pups (1 a); the influence through glass
(28) being the same as that exerted directly (27). "We
see in this and in the pupe produced by Dutch “gold”
(19—21) a great advance in the susceptibility of ‘these
pup over those of the Vanesside. Thus the highly
sensitive pupa of V. 7o is influenced in the same direction
by bright green (such as those of nature), yellow and
orange as it is by golden metallic surfaces and by white.
These all alike tend to produce brilliant green pupee
with a golden sheen over much of the surface. And
yet such pups would only be concealed on the bright .
green backgrounds. The pupa of P. napi is similarly
influenced by bright green, yellow, and orange, but is quite
differently affected by white and gilt. The pup in the
latter case are light bone-colowred (3 a), and certainly much
more effectually concealed on a white surface than if they
were green. ‘Traces of the same kind of sensitiveness at a
much lower level of development are rendered probable in
Papilio machaon from the results already recorded, and
will be seen to exist in Pieris brassice and P. rape, in the
formation upon a white background of intermediate, grey,
and pale forms rather than green ones. The similar effects
of bright green, yellow, and orange are certainly to be
explained as they have been in many other species, both
larvee and pup, by the fact that all these colours reflect a
high proportion of the effective rays, viz. the yellow and
orange. This will be proved in the case of P. napi by the
results of the spectroscopic examination of the back-
grounds which were made use of (see Appendix).

Yellow paper (29—81). Produced only the light (3 a)
and green pup (3 0), the latter strongly predominating.
The influence through glass (30) and at a distance (31)
was less strong in the direction of green, although the
number of pupze was too small to carry much weight.

Orange leno and paper, in some cases combined with
yellow (32—88), were even more powerful than the yellow
in producing green pup (30); in fact if we consider
those experiments only in which the pupz were directly
placed on the backgrounds or were only separated by the
thickness of the glass (32, 33, 35, 36, 37), no less than sixty-

404 Mr. F. Merrifield and Mr. E. B. Poulton on

seven green pupe (3 b) were formed as against five light
ones (3 a). The influence through glass (35, 36, 37) was
undiminished; but that at a distance (34, 38) was im-
mensely reduced, a single intermediate pupze (2),a largenum-
ber of light (3 a), and only a few green (3 b) being produced.

Planed deal (39—41) seemed to produce effects com-
parable to those of white, viz. that form of pupa out of
the various possible degrees, which harmonized best with
the background, viz. light (3 a), a single intermediate one
(2) also appearmg. No appreciable difference is to be
noted between the effects in strong light (39), shade (40)
and at a distance (41).

Looking at these experiments as a whole, the much
smaller effect produced by the coloured backgrounds
- upon the pupz “on the glass,” viz. at a more or less
distance from the effective stimulus, harmonizes well with
the results of previous investigation.

The strong tendency of the pupe, or rather the larve, to
face the light directly is well shown in the results of
Experiments (6) and (35) where the positions are recorded.
The great majority of the exceptions were due to the larva
placing itself parallel and in close proximity to the clear
glass front. A similar tendency to seek the angles
between a horizontal and vertical surface probably in part
explains the fact that so high a proportion directly faced the
light, the body line having been drawn into parallelism and
often into contact with the sides. In these experiments
only one pupa was parallel with and close to the back, but I
have noticed very many pupee of Pieris brassicx in shallower
breeding-cases, in this position. The few complete ex-
ceptions in (6) and (35) in which the head pointed directly
away from the light are enough to show that susceptibility
is unaffected by orientation.

Mr. Merrifield is to be congratulated in having proved
the high degree of sensitiveness possessed by this pupa.
While the susceptibility is as great as that of any other
species, even of the highly sensitive Vanessa io, the range
of controllable modification is probably wider than in any
other as yet investigated in sufficient numbers. This is
shown by the production not only of green and dark forms,
but of pale and grey pupe upon such backgrounds as
white paper and planed deal. At the same time the
range is not nearly so great as that of the larva of Amphi-
dasis betularia, which can produce on appropriate dark

Adjustment of colour in various pupex, ete. 405

surfaces many distinct shades ranging through Olack,
brown, and grey ; and also on the provision of the appro-
priate stimuli can become white or green (Trans. Ent. Soc.
1892, 7. ¢:),

The reactions of the pup of Pieris napi to the colours
employed agree well with the observations recorded in the
case of other susceptible pupee if allowance be made for the
wider range of controllable modification.

L—EXPERIMENTS UPON THE PUP OF
Preris brassicex.
1. EXPERIMENTS UPON THE WINTER PUP OF
Pieris brassice. (F. M.)

I took advantage of the pestilential abundance of the
larvee of this species to experiment on them and I exhibit
samples of the results, which will be described so far as
necessary in the following section. It is the less necessary
to refer to them at any length, because the species has
been so fully experimented on by Prof. Poulton as de-
scribed in the papers before referred to, and as will appear
later in this memoir, also during the present year (1898).
But I would call attention to one feature, that has been
carried perhaps a little further than had been done before,
in my experiments on this species and the next referred
to, P. rape. It seemed a fair inference both from Prof.
Poulton’s paper and from my personal observation of the ex-
periments with darkness on the other two species, that the
positive application by reflection or otherwise of some
decided colour (including black and white among colours)
was necessary to affect the colour of the pupe in a marked
degree. Accordingly I tried the experiment of surrounding
some pupze with clear glass away from all near reflecting
objects. For this purpose I placed the full-fed larvee in clear
glass cylinders covered by clear glass and resting on a clear
glass sheet several inches above the table, so that light
reached them all round. They were placed on a table
near the window. It will be observed that in the case
both of this species and the next, the pupe of the larvae
thus exposed to uncoloured light on all sides, rather closely
resemble those in darkness, the former being somewhat
darker than the later. In the case of P. napi darkness pro-
duced much variety of colour: of seven all but one are green
but of somewhat varying tint, and the one bone-coloured
and two of the green ones are much spotted with dark,

406 Mr. F. Merrifield and Mr. E. B. Poulton on

I found this species very troublesome as to the place of
its pupation. In no one case did I succeed in getting it
to pupate on a stick whatever its colour, and it had a way of
pupating on the clear glass in front of the cylinder or bottle.
To circumvent it I procured some white photographic
trays, some of which I covered with orange glass, and
others with deep green glass, while one was lined with
black paper and covered with clear glass, and another, left
white, was covered with clear glass. The space left
between the bottom of the tray and the glass covering varied
from about } inch to 3 inch or a trifle more, and in some
cases it will be seen the pupa bears marks of squeezing.
These instances are indicated by the word “screen,” in the
case of trays thus provided with screens of coloured glass,
“tray” where the covering was clear glass.

The whole of the pupz obtained (about 80) are displayed,
part in the Exhibition Case, and part in the glass-topped
boxes.

2. RESULTS OF THE ABOVE EXPERIMENTS. (KE. B. P.)

The pupe of P. brassice were compared May 38rd and May
6th, 1899, the results being shown in the following table (see
pp. 407, 408). The degrees of pupal colour are the same
as those suggested and fully described in Phil. Trans. 1887
(1.c. pp. 409, 410). The letters g. 0. w.y. indicate the faint
greyish green, orange, yellow, or white tint of the pupz in the
darker degrees. The tint is however usually very faint,
being greatly obscured by the dark pigment (see also
Trans. Ent. Soc. London, 1892, p, 439).

These results afford a very useful confirmation of
those which have been previously obtained. Thus the
black (1, 2,3) produced uniformly dark pupe, the effect
being as strong when the dark surface was behind glass (3)
as when it formed the surface to which the pupa was
attached (1, 2). The black surroundings also produced a
considerable effect upon pupz which were attached to the
clear glass at some distance (4).

Darkness (5) produced far more intermediate pupe,
although still upon the dark side of intermediate. As Mr.
Merrifield has suggested, this result is to be compared in an
interesting manner with that of clear glass (28). It is
probable that in these two cases we witness the results of
pupal tendencies undirected by any effective stimulus.

Adjustment of colour in varius pupe, etc.

DEGREES OF PuPAL COLOURS.

407

()
EXPERIMENTS. ee ——— Whitin Remarks.
‘k- rme- ‘ ritish|
ae diate Light. Yellow... Green.
(a) (8) (y) (2) | (8)
1. Black. lo | 389 The darker pupa
dead.
2. Black tray. lw} 29,y | Also one dead and
| one emerged: both
| probably (1 8).
3. Black, through) 1g | 2g |
glass. |
4, Black, on glass | 4 wy Also one dead,
9 probably a (1 8).
5. Darkness. | 2 9,0 3 g,W,Y Probably one more
pupa accidentally
transferred to those
on clear glass.
6. Green paper. ly ly 2 1
7. Green gauze. | 1 | 1
| | =|
8. Green, through | 2 w,g 3 1
glass. | |
9, Green screen. wo} al Also one dead,
| probably (2) ; and
| one emerged ; not
| darker than (1 y).
The (8) was a pe-
culiar greyish-green.
10. Green,on glass. 3 G,g9,u 3 1 Also one dead ; a
| (1 y), or darker.
11. Cabbage leaves. eae 1 1 | Also one dead, evi-
| dently a (2) or (8).
12. White. teedeee | 1 ;
| |
13. White paper ly |
(cylinder).
14. White paper. 2 avg) |) ly Also one dead, but
certainly a (1 8).
15. White calico. 2y,w | lw
16. White tray. lw | alae
17. White,on glass. | 2y 1 | |

408 Mr. F. Merrifield and Mr. E. B. Poulton on
DEGREES OF PuPAL CoLours.
‘i (1)
XPERIMENTS. : ae Remarks.
Fae ee aaer |e ineutes || Mesnee Green.
(a) (8) (y) (2) | (3)
18. Yellow paper. 2 Also one dead.
19. Yellow, in 3 0,99 Also one dead, but
shade, | probably a (1 8). j
20. Yellow, 2 W,Y
through glass.
21. Yellow, on 1
glass.
22. Yellow com- 1
partment, on glass
with patch of white
paper behind.
| 23. Orange leno. lw lg Also one dead or
emerging, about a
(1 ).
24. Orange. lw il 1
25. Orange, 1
through glass.
26. Orange screen. 4
27. Orange on 1 3 One (8), a curious
glass. sreyish form.
28. Clear glass. 6 Also one dead,
probably a (2) or
(3). The (1 8)s are,
one greyish-orange,

}

four green, one
white.
Probably one pupa

belongs to the group

in darkness.

Green paper, gauze and cabbage-leaves (6, 7, 8, 10, 11)
caused the appearance of far lighter pupa, the majority

being the whitish-yellow degree (2).

Here too the influ-

ence through glass (8) was as strong as in the other

experiments

(6, 7).

The few pupe subjected to light

through green glass (9) were of the two lightest and
greenest degrees, thus confirming the effects described in
Trans. Ent. Soc. Lond. 1892 (pp. 429—432, 446, 466—

Adjustment of colour in various pupe, ete. 409

468), and ascribed to the greater concentration of the
effective rays. The stimulus produced a strong effect at a
distance (10). The powerful effect of a natural green
environment (11) is of great interest.

The experiment with white (12—17) was especially
useful, as this environment had been almost omitted from
the previous investigation of this species. It is at once seen
that the effect is to produce intermediate pups, inclining
towards the dark side. In this, the results differ widely
from those obtained with Vanessa urtice and V. io in
which white surroundings produced strong effects in the
direction of the golden and green pupz respectively. There
is great uniformity in the results obtained by the different
white backgrounds, and the action at some distance (17)
was clear. It will be found that my own experiments (43
to 48) in 1898 (see pages 415, 416) lead to the same
conclusions as those which result from Mr, Merrifield’s.

Yellow surroundings (18 to 22) produced the usual
strong effects in the direction of the palest, greenest pupe,
the influence being much reduced in shade (19), and also
reduced when acting through glass (20). An influence at
some distance was probably exerted im (22).

Orange surroundings (23 to 27), for the most part, pro-
duced the same effect as the yellow, the orange leno (23)
being an exception. Influence was strong through glass
(25), and at some distance (27). The orange screen (26)
produced considerable effects, in accordance with the
principles already explained (Trans. Ent. Soc. Lond.
1892, U. c.); although an even stronger result might have
been expected.

This account of the results should be read in relation
to the Appendix, in which the colours reflected from the
various backgrounds are analyzed.

3. EXPERIMENTS UPON THE WINTER Pura oF Pieris
brassice. (HK. B. P.)

I also took advantage of the immense abundance of this
species to repeat some of the experiments made in previous
years upon insufficient numbers. The larvee in part came
from St. Helens, Isle of Wight (experiments 5, 6, 37, 38,
39, 40), partly from St. Helens and near Reading (13, 41,
42), and partly from St. Helens and near Oxford (remain-
ing experiments, including those with conflicting colours,

410 Mr. F. Merrifield and Mr. E. B. Poulton on

and excepting Nos. 4 and 12). They were collected by
Miss Cora B. Sanders (St. Helens and Reading), by Mr.
W. Holland (Oxford), Mr. A. H. Hamm (Oxford), and by
myself (St. Helens). Experiments 13, 41, and 42 were
conducted by Miss Sanders; 5,6, 37, 38, 39, and 40 by me,
and the remainder by Mr. Holland and Mr. Hamm. I
wish to express my warm thanks for all the large amount
of kind help I have received.

The conditions of experiment are sufficiently shown in
the following tabular statement (see pp. 411—416), the con-
stitution of the reflected light being given in the Appendix.

The pupz were examined on May 6, 7, 8, 9, 10 and 11,
1899, when several of them had emerged or were emerg-
ing. All these, however, are indicated below, and nearly
all could be tabulated with considerable accuracy.

The positions of the pupz in relation to the receptacles
employed and to the light were noted in a large number of
examples, with the following results. Thirty-six pups
were fixed to the roof (or in the angle between it and the
back), with a direction parallel with the front (generally
clear glass) and back ; 25 were fixed to the front, back,
or sides in a vertical position with the head uppermost
(including a few with the head downwards—cases in which
pupation certainly occurred during a temporary reversal of
the position of the receptacle); 41 were fixed, almost
invariably to the roof, with the head pointing directly
towards the light, viz. with the line of the body at right
angles to the front (generally clear glass); 11, otherwise
similarly placed to the last, had their heads pointing
directly away from the light; 15, otherwise similar, had
their heads obliquely directed towards the light; 10 their
heads obliquely directed away from it. It is therefore
clear that there is, upon the whole, a tendency to direct
the head towards the light, although the tendency is not
nearly so strongly marked as in P. napi. There was no
appreciable ditference in the colour according as the head
pointed towards or away from the light.

Reviewing the results of the tabulated experiments,
the effect of black (1 to 9) is, with certain exceptions,
similar to that which has been obtained before, and also to
Mr. Merrifield’s investigations carried on simultaneously.
These exceptions are the very dark pup obtaimed in
almost complete darkness (8), and the very unusual hght-
ness of some of those in dim light (6, 7).

|
|
|

Adjustment of colour in various pupe, ete. 411
he
— | er- il 2 i
EXPERIMENTS. Dee | cr Tight. e2| 6 Remarks.
(a) (8) (y) | (2) | (8)

1. Black lined cylinder | | On roof near and parallel
(wide and low, placed on | | with glass front. Emerging.
side with clear glass front). | | |

2. Black lined cylinder , 1a Similar position to above ;
(very similar to last). together with three other

dead pup (unclassified).
Two of the (1 a) greenish, |
| the other emerged.

3. Smallish black eylin- | lg Pupa on black paper roof
der (intermediate in size | | of upper compartment.
between 7 and 24, same |
arrangement) with two |
compartments, moderate
illumination. |

4. Tarred fence near Ox- | | Two pupe found April 2,
ford. | 1899, by A. H. Hamm. They

| were very dark, probably
| | | (1 B) or even (1 a).
sae as | | ; | One (1 8) emerging. Also
= 2 BES ox : moe | | 4g.g,w?\4g, 0,0, Y anes ed ee prob-
i= "so po oO roo}. | | |
Bis 7 tg | | | ably (1 y), two (1 8) or
=o) oa |(1 y), and one uncertain.
=o 252 | | [Als d, and
Ho © 5 | so one emerged, and was
= Bees | a (ly). Pupe chiefly in
a8 2 E = | two crowds of seven to R.,
Bee seo = | and five to L. of roof. The
ze oe oS" | heads were nearly as often
pores | away from light as towards
"S55 Paes it ; often oblique, and often
=e re On parallel with back or front.
a=] 5S SAS Loose on | Also one dead ; it had
© 25 | oor. | lg been a (1 7).
I Attached to roof. 40,w,,; 1y 2 1 Also six dead ; 1=(1 8),
ee 0,0 | 14 Gig) “or Gl +), and
go | 1=(1y). Also two emerg-
Se ing, a (1 y) and a (1 y) or
Es | (2). A group of four at
Se | back and of ten on left side,
Or | heads of those (six) not
By parallel with back nearly
SS always turned to light.
net Attached to black ly Also three dead on the
= | net window. sides of box ; all (1 y) or

A rather lighter or darker than

Be | this. Also one dead, loose

el on floor—a (1 8) or (1 ¥).

412 Mr. F. Merrifield and Mr. E. B. Poulton on
@) aE] 2
EXPERIMENTS. Dark. Eee Light. Ee | & Remarks.
(a) | (8) (y) | (2)| @)

7. Small black com- | 1 Pupa from a single larva
partmented cylinder, in | | | put in lower compartment.
deep shade. The cylinder | | Rather more pigment than
about 10 ¢.m. high by 6 | usual, but deep green
c.m. in diameter. A black | ground. Pupa fixed verti-
paper partition divided cally, head upwards, at top
the cylinder into two of side opposite window, but
chambers illuminated by a | | latter closed above and the
narrow window 2° to 3°5 roof very convex below, so
e.m. wide. | that larva had been in deep

| shade.

8. Black cylinder in| lo 39 | All pupe isolated. All
almost complete darkness : (1 8) very dark and two of
16 ¢.m. diameter by 10 them nearly (1 a).

c.m. deep.

9. Black-lined (3 sides) | TONGA aSrO gn? | Four pup isolated on
rectangular glass case (18 | | |roof; the (1 a) g, on clear
cm. square by 28 c.m. | glass window, just below
high) with perforated zine | black binding and zine at
roof, | top, and near black paper of

| | side.

10. Perforated zine roof | 4 0, 0, g, | lo Also one emerged ; was a
of yellow-lined (three sides) | | Gg | |(18)or(1y). All tabulated
rectangular glass case (28 | | | pup on roof in groups of
e.m. square by 38°5 c.m. | | | two and three.
high). | | | |

11. Similar zine roof of} lw | 20 | Also one dead, probably
similar white-lined (3 sides) a (1 y) or (2). All pupe
case (28 c.m. square, by | isolated on roof.

33 c.m. high). | | |
12. Brown paper. hey al | | Found in O. U. Museum
| _by A. Robinson. Probably
/an escape from my cases.
|Imago had emerged, but
/pupa so dark it may have
| been even a (1 a).

13. Salmon-pink box in 1 All sides and roof sloped
strong light (24 ¢e.m. by | | inwards to a back only 10
13:5 ¢.m. in section in | | ems by) 18 cams eins
front). | | | caused a very strong illu-

mination.

i4. Orange paper-lined | 1 | 2 Also one emerged, a (2)
| cylinder (22 c.m. diameter, | | or (3). The (1 y) bright
| 10.2c.m. deep). Placed on | yellowish green, All pupa |

| | on roof, |
!

side with clear glass front. |

|
}

Adjustment of colour in various pupx, ete.

EXPERIMENTS.

15. Orange-lined cylin-
der (27 c.m. diam. 8 c.m.
deep) arranged as the
last.

16. Orange-lined cylin-
der (30°5 c.m. diam. 9°8
e.m. deep). Arranged as
the last.

17. Compart-
ment I,

quory
‘daap
Wee ral

-ANO,T

18. Compart-
ment IT.

19. Compart-
ment III.

pis uo poovld xog
8 'UL'D OT JUeUyIedUIOD

(1)

ee
Inter-

mediate.

(8)

— Whitish-
LY Yellow.

lg

20. Compart-
ment IY.

*xoq peul[-esurio pojuotmyredur0o

SSBIO TBITO TM 0
‘ul'd F.9 ‘arenb

21. Compart-
ment H, orange-
lined.

o00 Gt)

22. Compart-
ment L, orange-
lined.

‘deop ‘url's ¢.g

‘mo1}0eS UT

G1°¢ Aq ‘u's G.F

PF

23. Lower compartment
of small orange-lined cylin-
der, similar to 7.

24. Lower compartment
of large orange-lined cylin-
der (8 cm. diam. by 18 |
em. high, with window
3°4 c.m. wide) arranged
similarly to 7.

s}uouryred wL0—H

eo Green.

—
wa

415

Remarks,

bo

One (3) on glass front,
three-quarters up. Others
on roof.

Also one emerged, evi-
dently a (3). One of the
(3)s had the pigment of a
(1 y) but bright green
ground of a (3). All pupe
on roof.

Two on roof, one on floor.
The box had clearly been
turned over for a time, so
that floor became roof.

All on roof.

Also one emerging—a (2)
or (3). Two on roof, one on
angle of roof and side, one
head downwards on side ;
clearly due to box having
been turned over.

One pupa emerging, but
probably a (1 7); one dead,
but was a (2) or (3). Both
on roof.

On back, close under
roof.

In angle between roof and
back.

In angle between roof and
side.

Position of pupa clear
although emergence had
oceurred. On roof, close to
side.

414

Mr. F. Merrifield and Mr. E. B. Poulton on

(1) if ES 2
aa | 3
om Inter- So | &
HAP SRIMENS Dark. | mediate. | Light. |FR| S Rens
(a) (8) (vy) | (2) | (3)

25. Orange-lined rectan- lg ly oi ws Also two dead, a (2) and
gular glass case (30°5 c.m. a (2) or (8). One of the
square, 41 c.m. deep) (2)s tabulated was loose
on side with clear glass on floor: all the rest of
window. the pupe on roof. The

darkest pup in dim iliu-
mination.
[eae |
2-2 6 26. Compart- 2 On roof.
© 3 © ment E, orange-
3.8 & %. | lined.
Bava
eee s 27. Compart- 3 Two on roof, one on side.
S 2 “5 | ment F, orange- Also one dead on side and
a3 it = | lined. one on roof ; both probably
Same (8) or one perhaps (2).
a ae g _ Sa
225s) 28. Compart- Zeal All on roof ; one emerged
BC as ment G, orange- and one emerging, but
: wos lined. position seemed certain.
— > |
iJ : ae
ae @ |_29. Compart- lo 1 Also one emerged, almost
F > | ment H, orange- certainly a (2). All on
857 5 lined. roof.
Ss 5 5 =| 30. Compart- 1 Pupa loose on floor.
=~ |mentA, yellow-
& IS & | lined.
ee
es ae 31. Compart- One pupa dead on roof,
a = ment E, yellow- a (2) or (3).
ts @ Wy | lined.
: ae

32. Upper compartment 1 Pupa on side just below
of large yellow-lined cylin- roof.
der Similar to 24.

3. § =| 33. Compart- 1 In back corner of roof.
egy |ment A, yellow-
& 5 Es lined.
ses ert hace Goal
mo 5 34. Compart. 2 One on roof, one on back :
°='3 |ment B, yellow- latter with the green ground
a | lined. of a (3).
EE —'—--- so
pe 35. Compart- ij) al Pup isolated on roof.
moet
=~ % yment C, yellow-
fo} .
5 & | lined.
+O
aris ean ee eth ee A
5 ,| 36. Compart- 40,9,9,Y 1 Four crowded, one iso-
St |ment D, yellow- lated (1 y) y on roof.
5 & | lined.
: |

Adjustment of colour in various pupx, ete.

415

(Q) Se] 3
lm eS co)

EXPERIMENTS. Dae. Aes | Lizht, ae & Remarks.
(a) (8) | (y) (2) | (8)

37. Yellow-lined box lo 1 9 Also one dead, but clearly
placed on side, 20 e.m. | a (3). Four (3) and the
square, 6 c.m. deep. | (1 8) on yellow leno window.
Window 14°5 c.m. square, Six (3)s on roof.
yellow leno covered.

38. Yellow-lined box wy Also two dead, probably
placed on long side: 14 a (2) and a (1 8) or (1 4).
c.m. by 18°5 e.m. in see- All pup on roof; mostly
tion, 8°5 cm. deep. in small groups.

Window 10°5 c.m. by 15
c.m. as above,

39. Yellow-lined box lg ee Also five dead, two emerg-
placed on short side: 21 ed and one emerging, but
e.m. by 12 c.m. in section, all certainly (2)s or (3)s.
9 cm. deep. Window 8 Ten on roof, six on left side
em) bys 15:5 e.m: as (three (2)s and three dead).
above.

40. White box (31°5 cm. 29,Y 2 | Also three emerging; they
by 16 c.m. by 6 c.m. deep) were either (2) or (3),
with yellow leno window or in one case possibly a
biesme by. 23:5) chm), (ly). Allattached to edge
probably placed upwards. of leno, where pasted to

box.

41. Green cabbage-leaves, |) & All seven on separate
leaves, except that there
was another dead one, cer-
tainly a (3), on the same
leaf as one of them. Also
one pupa dead, but certainly
a (3), and two emerged, both
(2) or (3), Also a pair on
one leaf, one dead and one

| emerged, both certainly
(8s).

42. Dark purple-lined 6g yy al Also six dead probably
box with white roof, in three (2)s, one (2) or (1 y),
very dim light: (12 ¢c.m. one (2) or (3), one (1 8) or
by 19°5 eci:m. by 7 c.m. (1 y). Also one emerged ;
deep). probably a (2). All tabu-

lated pupz on white roof,
except two (2)s, loose on
floor. All pupe very
much undersized.

43. White opal gas Se Pup isolated in upper

globe (usual size).

|

part of globe.

416 Mr. F. Merrifield and Mr. E. B. Poulton on
| () aiaier
Se ees
rating | Inter- la3| &
EXPERIMENTS. lenarie | Tiediake lh etheke, le a | oS Remarks.
@) | () | @ |@]®
BSE | 44. Compart- 29 Pupe on roof, isolated.
Es 3 = ment A.
2 2 =a ee
aes 45. Compart- | 1g | 1 | Also two dead or emerg-
Sa~ 5 | ment B. | | ing, probably (1 8), or per-
SEE | | haps (1 y) in one case. All
wees | pupz on roof, three crowded.
+ 26 —— |
e 5 46. Compart- | lg ly | One on roof, one on floor
oO, S| ment C. | (case having doubtless been
Bs | turned over).
ee
2-45) 47. Compart- | 29 ie Two on roof isolated, one
5&6 | ment D. | on back.
5 Lye i i ae ee
48. Rectangular glass 29 lg Two pup isolated on
case with three sides lined | ee ae (1 .) a black
with white paper, and) | binding of angle between
white opal glass roof | | window and side, but close
(20°4 cm. square, 30°6 | to white paper.
e.m. high).

The effects of a dull surface of perforated zine (9—11)
and of brown paper (12) were, as might be expected,
practically the same as those of black.

Salmon paper (18) acted like orange (14—29), and
produced the lightest and greenest form of pupa. The
results of so many experiments with orange are very
striking, especially as a very deep reddish-orange surface
paper was employed.

Yellow (80—40) also produced striking results in the
same direction, but not equal in the proportion of the
greenest pupze, to those of orange.

Green cabbage-leaves (41) acted like orange.

Experiment 42 was very mixed, the pup being, almost
all of them, fixed to a white surface in a dark purple box
in a very dim light. They were strongly on the light side
of intermediate.

White (43—48) has been already described as producing
pupze on the dark side of intermediate (see p. 409). There
are, however, some few marked exceptions in both Mr.
Merrifield’s and my experiments, in which the lightest and
greenest pup were obtained.

Thus Mr. Merrifield’s and my experiments in 1898

a

a, ee

Adjustment of colour in various pupe, etc. 417

afford most useful confirmation in the case of a species
which has not been hitherto sufficiently tested, besides
bringing evidence of its behaviour under conditions as yet
hardly tried at all. In the next section is recorded an
experiment upon the same species, which, more than all
others, needed repetition because of the important con-
clusions which follow from it.

4. EXPERIMENTS WITH CONFLICTING COLOURS UPON THE
WINTER Pup oF Pieris brassice. (E. B. P.)

I had long been anxious to repeat some of these experi-
ments upon the species of Piering because of their extreme
suitability for such an investigation and because of the
important conclusions which follow from the results.

It had been originally supposed by Mrs. M. KE. Barber
(Trans. Ent. Soc. 1874, p. 519) that particoloured pups
are produced by a particoloured surface—a conclusion
which naturally followed from the views held by many at
that time as toa direct “sun-picture or photograph ” on the
fresh, moist skin of the pupa. A single pupa of Papilio
nireus had seemed to support this conclusion.

In 1886 I made a large number of conflicting colour
experiments on Vanessa wrtice (Phil. Trans. 1887, J. c. pp.
368—392). The contrasted colours were, however, only
applied during Stage III when the larve are suspended
preparatory to pupation and are less sensitive than at an
earlier period. Nevertheless the results were sufficient to
make it highly improbable that any parti-coloration of the
pupal surface could occur as the result of such a mixed
stimulus, and led to the conclusion that the effects were
due to the intermediation of the nervous system in the
central parts of which the opposing influences from different
regions of the body met and produced more or less of an
equilibrium, resulting in the dispatch to all parts of the
body surface of stimuli producing intermediate effects.
These conclusions were so far-reaching and important that
it was necessary if possible to repeat the experiments with
other species in which the conditions were more favourable.
Although such experiments were not made in 1886 upon
the Piering, it was clearly seen, when the paper came to be
written, that they would be peculiarly suitable for the
purpose, because of the great length of the whole sensitive
period and the fact that its two stages are both passed
under conditions which are eminently favourable for such

TRANS. ENT. SOC, LOND. 1899,—PART IV. (DEC.) 28

418 Mr. F. Merrifield and Mr. E. B. Poulton on

an investigation. A few such experiments upon P. rape
were attempted by G. C. Griffiths in 1887 (Trans. Ent.
Soc. 1888, pp. 265, 266), and the results upon the whole
supported those obtained in the case of V. urtice. But
the experiments were not very convincing because the
colours employed were not those which produce the most
marked and opposite effects.

In 1888 I made some experiments of the kind upon
P. rape and P. brassicw (Trans. Ent. Soc. 1892, pp. 445,
446 and 484), using a box lined with black and orange
squares, but owing to the excessive mortality from the
attacks of ichneumons only 6 of the latter and 2 of the
former could be tabulated. The results however entirely
confirmed the experiments made in 1886 upon JV. urtice,
intermediate and not parti-coloured pupz being always
obtained. In 1892 I made a large number of experiments
upon Vanessa io (1. ¢. pp. 420—426), and again upon
V. urtice (1. c. pp.891—397). Furthermore a method had
by then been arrived at which enabled the larve of the
Vanessid# to be subjected to conflicting colours durimg
the whole of the sensitive period; and another method
whereby the dorsal and ventral surfaces could be subjected
to opposing stimuli (in the case of V. 20,7. ¢.). In all
previous experiments the anterior and posterior parts of the
body had been thus treated. All such modifications and
additions yielded confirmatory results.

I was nevertheless very anxious to repeat the experi-
ments upon the Pierine and therefore took advantage of
the abundance of P. brassice last year (1898). Here,
again, however, owing to the Jchneumonide, my results
were not at all what I had anticipated; but taking them
in combination with those which have been already
published they leave little or no room for doubt.

Two conflicting colour-boxes were made for me by Mr.
W. Holland and Mr. A. H. Hamm. The first box had an
internal section of 54°5 cm. by 14:2 c.m., and a depth of
74 c.m. (from back to front in the position made use of).
It was used resting on one long side with a clear glass
front. The roof (the side uppermost in the position in
which a receptacle is used is here always called the “roof ”)
was divided into 9 bays of about equal size by means
of 8 hanging partitions (each 74 cm. deep and thus
extending from the back to the clear glass front, and
hanging down for a distance of 3°5.c.m.); while the back

—

a i

Adjustment of colowr in various pupe, ete. 419

was similarly divided into 10 bays by 9 partitions which
alternated with those of the roof. These were 9:0 c.m.
high and their lower borders 2°0 e.m. from the floor, while
they projected 3°5 cm. from the back towards the glass
front. The object of this division of the internal surface
was to separate the larvee as much as possible, and thus
minimise their influence upon each other during the
sensitive period. The whole internal surface, except the
floor, of the box, and both surfaces of all the partitions
were lined with a chess-board pattern of orange and
black. each 14 c.m. square, and thus as nearly as possible
half the length of an average mature larve of P. brassice
when resting in Stage II preparatory to pupation. The
pattern was made by ruling the outlines of the squares in
pencil upon a sheet of deep orange surface paper and then
carefully pasting black tissue-paper squares over alternate
orange squares.

It is much to be regretted that an experiment conducted
with so much care should have produced such limited
results as regards the numbers of pupe.

The first box only contained 3 pupz which could be tabu-
lated with certainty on May 6th when the examination was
made. All were fixed in the left-hand bay of the roof,
near to the glass window.

One pupa was fixed diagonally across a black square
with the end of its tail lying on another one, and the head,
directed towards the light, overhanging an orange square.
It was dead but had clearly been a (2) or a (8).

The second was fixed parallel with the glass, the posterior
3 of its body on the anterior part of a black square, but a
little overhanging an orange square in front, and the
anterior 2 similarly on orange and overhanging black as
well as orange in front (viz. towards light). It was a
greenish (1 £).

The third pupa was also parallel with the glass, although
the tail curved towards it. The posterior 2 of its body
was on the posterior (viz. away from glass) border of (so
that the right side overhung) a black square, the anterior
4 similarly on and similarly overhung orange. The left
side (away from glass) overhung the opposite colours in
each case except the posterior 4 which curved towards
light, viz. towards the middle of the black square, so as to
overhang black. The pupa was a greenish (1 y).

There were also two other dead pupze which could be

420 Mr. F. Merrifield and Mr. E. B. Poulton on

placed with tolerable accuracy, by an examination of the
persistent cuticular pigment.

One of these pup was at the back of the roof in the
same bay as those described above. It was chiefly upon
the black, extending obliquely across parts of two black
squares which were in contact at the junction of its
anterior and middle thirds; at this point therefore and
on each side it, the sides of the body overhung orange.
There was very little pigment and it had been certainly
not darker than a (1 y), and probably either a (2) or (3).

The second pupa was on the roof of the bay at the
opposite end of the box, close to the hanging partition.
Its anterior half was on black, its posterior on orange,
crossing almost at the middle of the adjacent sides of the
squares; its head towards the light. It had probably
been a (1 8).

Although these five pup had been subjected to the
most strongly contrasted influences in various regions of
their bodies, there was not, in a single instance, the
faintest trace of parti-coloration. The opposing influences
gave rise to a general effect which was almost exactly
intermediate between the effects which they would have
respectively produced if they had acted alone. It is to be
noted that if there is any deviation from the intermediate
position it is in the direction of the effects produced by
orange. The larvae seem upon the whole to have rested
in contact with black in preference to orange, and thus
overhung the latter colour rather more than the former.
But many more experiments would be required in order
to estimate exactly the relative strengths of these two
opposing influences; and it is noteworthy that the results
of my experiments upon this species in 1888 led to different
conclusions upon this point.

The second box was 43°6 c.m. x 154 c.m. in internal
section and had a depth of 10° cm. It was similarly
arranged with 6 hanging partitions, and 7 projecting
from the back each 85 c.m. long but in other respects
similar.

The second box was examined on May 7th. In this
case six had pupated on the glass. Of two isolated pupze,
fixed in a vertical position with head uppermost, one had
emerged and one was dead or emerging; both were
probably (1 y). Two more, similarly placed, were dead
and could not be classified, Of two near together, but

Adjustment of colour in various pupe, ete. 421

otherwise similarly fixed, one was a greenish (1 y) and
one, emerging, was probably a (1 y) or (2).

Two pup were fixed to the parti-coloured surface of
the roof. Of these one diagonally crossed two black
squares In a manner very similar to that of the first
described of the two dead pupz in the first box, which
could not be classified with certainty. The direction of
the body was oblique with the head away from the light.
It had emerged, but had clearly been a (2) or (8). The
second was fixed in another bay, near to and parallel with
the glass. The posterior 2? of the body crossed a black
square, the anterior 4 was on the next orange one. It had
emerged but had evidently been a (1 8) with quite dark
pigmentation. There was no trace of Jess pigmentation in
the anterior third of its body.

These results entirely confirm those obtained in the
first box.

Although further experiments of this kind are to be
desired, especially upon so sensitive a species as P. napt,
it may be regarded as certain that the conclusions derived
from the earlier experiments with conflicting colours are
sound, and that not parti-coloured pupz but uniform
intermediate ones are obtained in this way.

F.—NOTES ON THE STRUGGLE FOR EXISTENCE
IN THE LARVA OF Preris brassice. (HE. B. P.)

I have previously noted the numbers of this species
which perish from the attacks of parasites in a year in
which the larve are specially abundant. It seemed of
interest to obtain further records, and I accordingly asked
Mr. Holland and Mr. Hamm to keep notes of the number
of larvze attacked by ichneumons and the numbers dying
apparently from other causes, which they removed from
the cases containing the mixed larvee from St. Helens and
Oxford (see p. 409). The results are recorded in the table
on the following page.

In the breeding-cases from which these larvae were
removed only 121 pup were taken, including the dead
ones; so that the extinction is on a vast scale. Even if
it be conceded that the larve dying without the appear-
ance of parasites, and the dead pupz, were entirely due to
the conditions of experiment (such as the possible intro-
duction and spread of some form of bacterial disease), the

422 Mr. F. Merrifield and Mr. E. B. Poulton on

extinction due to the attacks of a single species of parasite
is still immense, only about # of the whole surviving it.

Number of Number of deaths
Larve attacked apparently due to
by ichneumonidee. other causes.
1898. Sept. 22 23 ste
Qe 25
9 =
9 ©6024 70 —

~)
[es

~~
Sb bw
ooNI OD
lor)
rs
Wm
CO

NG

29 47

oo
i)
Or
rs

m Co oS or Co bo
ee
bo

-
.
et

1899 at least 31 at least
Total 1016 Total 273

G.—EXPERIMENTS UPON THE PUPA OF
Preris rape.
1, EXPERIMENTS UPON THE WINTER PuP# OF
Pieris rape. (F.M.)

The next experiments tried were on this species and
are detailed in the following section. I was late for this
species and did not experiment with more than 50 or 60
larvee, from which I obtained the single row of about
40 pupze which are now shown in the Exhibition Case.
Here I call attention to the contrast between those in
the black surroundings on the one hand and those in
the green paper, yellow, or orange surroundings on the
other, the green in this species seeming more effective than
it proved with the other species and the yellow less so.

2. RESULTS OF THE ABOVE EXPERIMENTS. (E. B. P.)

The colour variations of the pupze of P. rape have been
already described and figured (Phil. Trans. 1887, 7. ¢. pp.
410, 411, Plate 26, figs. 31—41), and it is here only
necessary to state that the standard classification begins
with the darkest pupze (1), ranging through the less dark
(2) and still lighter (3) to the pale (4) and the green (5).

Mr. Merrifield’s pupze were compared on March 20,
1899: the results are given below without much detail,

Adjustment of colowr in various pupe, ete. 423

inasmuch as the species is already known to be susceptible.
In addition to those tabulated below, Mr. A. H. Hamm
found two very dark pups, evidently (1), on a tarred fence
near Oxford, on April 2, 1899.

All the (5)s were bright green and very pronounced

Degrees of Pupal Colour. |

EXPERIMENTS. z B ad =3 3 B Remarks.
(1) | (2) | (8) | (4) | 6)
1. Black. 2 Also 2 dead.
2. Black, on glass. 2 Pinkish ground-colour.
3. Dirty white paint. 1 Very dull, dark pupa.
Removed from P. napt.
4. Darkness. 4 1 (3) Pinkish :
(4) greenish.
5. Green. 1 | 1 | Also 1 dead:
(4) greenish.
6. Green, through glass. 1 Also 1 dead or emerging.
(4) light pinkish.
7. Green screen. 1 Also 1 dead.
Pupa greenish.
8. White, on glass. | 2 1 greenish, 1 pinkish.
9. Yellow. | 1 | 1, (4) greenish.
10. Yellow in shade. il 1 | (4) greenish.
11. Yellow on glass. 2
12. Orange. | 1 1 | 2 | Also 1 dead:
| (4) pinkish.
13. Orange screen. 1 | Also 1 dead (removed
from P. napi).
14, Clear glass. | or ae Also 1 dead.

representations of this degree. A few other conditions
were tried, but the single pupz subjected to them had
died and the colour had changed so greatly that they
could not be tabulated.

The numbers of the pupz are not large and the results
quite confirmatory of previous experiments. (Phil. Trans.

424 Mr. F. Merrifield and Mr. E. B. Poulton on

1887, J. c., and Trans. Ent. Soc. Lond. 1892, J. ¢.: see also
G. C. Griffith’s experiments on this species in Trans.
Ent. Soc. Lond. 1888, p. 247.)

It is interesting to observe the relation between the
effects of darkness (4) and those of clear glass (14). The
latter produced even darker pupz relatively to the former
than in the case of P. brassicx.

The table does not support Mr. Merrifield’s conclusion
that the yellow (9, 10, 11) was less effective than in the
case of P. brassicx. The six pupz were of the two lightest
degrees, four of them green and the other two greenish.

The effect of the green and orange screens (7, 13) is
confirmatory of previous results with other species (P.
brassice and V, 20).

It is not necessary to comment further upon the other
results, all of which will be clear upon an inspection of the
table on p. 423.

H.—EXPERIMENTS UPON THE PUPA OF
Vanesside.
1. EXPERIMENTS UPON THE PuP& OF Vanessa urtice AND
Pyrameis cardwi. (C. B.S. and E. B. P.)

In the course of our investigations in 1898 into the
struggle for existence during the pupal period of Vanessa
urtice it was necessary to produce a very large number
of pup with colours as widely contrasted as possible. In
order to achieve this we made use of black surroundings
on the one hand and gilt (Dutch “gold”), yellow, orange,
and white on the other. There was abundant evidence in
the 700 pups which we obtained of the previously recorded
influence of these surroundings, and also, to our frequent
annoyance, of the effect of the dark surfaces of the larvee
upon one another. In fact so powerful was this influence
and so gregarious were the larve under the conditions of
our experiments that intermediate pups were generally
produced when the lightest forms were desired. The con-
ditions of the investigation rendered it impossible to isolate
so many larve in separate cases.

Many of the larve pupated on the leaves and stems of
the food-plant (nettle), and when isolated brilliant golden
pupe were almost invariably produced.

A few larve of P. cardwi were also found and subjected
to black and white surroundings: the pupz being dark in
the one case, and light, and often brilliantly metallic, in
the other.

ee oe

Adjustment of colour in various pupe, ete. 425

2. EXPERIMENTS UPON THE Pupa& oF Vanessa v0. (MABEL
EK. NotLey, FLorENcE A. Wricut, and E. B. P.)

The experiments of last year upon the struggle for
existence during the pupal period are now being repeated
in the case of Vanessa io. In this case all the pups were
obtained at the outset of the investigation, and the results
as regards their colour susceptibility can now be given.

Weare greatly indebted to Mr. W. Farren of Cambridge,
and to Mr. H. W. Head of Scarborough, who sent us
numerous companies of larvz in excellent condition. Kind
help was also received from Mr. W. H. Harwood of
Colchester, and from Mr. A. E. Holdway of Newton Abbot.

The larve thus obtained were placed in a large number
of “light” and “dark” receptacles, the former being lined
with orange, yellow, or white paper (white opal glass was
used in the case of a few larve), the latter with black
paper (a few were attached to the dull surface of perforated
zinc). All were placed in a strong light and only shielded
from the direct rays of the sun. A few were subjected to
conflicting colours in one of the boxes described on pp. 418,
419.

The results obtained are tabulated below (see p. 426), the
degrees of pupal colour being those described in Trans. Ent.
Soe. Lond. 1892, p. 398; the (1)s and (2)s being the darkest
forms with the underlying green completely or very nearly
concealed by pigment which is blacker in (1), lighter in
(2); the (4)s and (5)s being distinct green forms very
bright and glittering in (5), duller and with more dark
pigment in (4); the (3)s intermediate.

The pup which were attached to the nettle-leaves,
leaf-stalks or stems are indicated by the letter m, and
those found loose on the floor by the letter 7 These facts
were not however recorded at the beginning of the experi-
ments so that more pupe were in reality found in these
positions in the companies received at first. The facts are
important inasmuch as the pupz on the floor were ad-
versely influenced in the experiments with light surround-
ings, the pupe on the nettles in the experiments with dark.

The companies are tabulated separately below, but it
was not thought necessary to describe each separate re-
ceptacle, as this work is confirmatory. The numerous
receptacles in which the larve of each company were
placed are grouped together as “light surroundings,” and
“ dark surroundings.”

426 Mr. F. Merrifield and Mr. E. B. Poulton on

Companies of Conditions of Degrees of Pupal Colour.

Larve. Experiments. (1) (2) (3) (4) (5) Totals.

Company of 64 Light | | 2 11 22 35
larvee (3 died), re- jsurroundings. | | | (3 n.) (3 7.)
ceived July 11, | | |
from W. Farren of Dark 4 | 9 il 1 | i 26
Cambridge. ; : 1 (G72) (3 n.) (4 2.)

surroundings. | (lf) (1 f)

Company of 172 Licht 1 2 7 26 90 126
larve (26 died), aa at ae (Ca) Gin) (8 .) | (21 n.) | (29'2.)
received July 11, ere (3 f.) (8ys) | oy
from W. Farren of SS |
Cambridge. Mack | 9 ' 4 3 | 1 3 20

eases tno) 17) May Aller 1 7.) 17.) | Gas
surroundings. (2 f) (27)

Company of 163 Licht 4 2 2 72 80
larvee (16 died), anid (1/.) (383 n.) | (33 2.)
received July 13, era aay)

from W. Farren of | |] —— — | —_—_
Cambridge. Dark 10 44 y 1 3 67
surroundings.

Company of 87 Light 2 4 3 hes Bil 43
larve (16 died), |surronndings. (ales) | (13 m.) | (14 2.)
received July 14, “a S|
from W. H. Har- Dark 8 10 4 2 4 28
wood of Colches-|surroundings.| (2 7.) | (5 2.) (4m.) | (22.) | @ 2)? | (tare)
ter. |

Company of 57 Lie} 1 2 4 29 36
larve (16 died), on | (9 n.) (9 n.)
received July 14, SUTOURGIngs. | | (i). | 26L 79) aa
from W. H. Har- [Nee SS sy ee
wood of Colches- Dark 3 1 1 5
ter. surroundings. |

Company of 148 Terehe 4 1 She 8) 83 110
larvae (8 died), re-| 718 i (2n:) | (m.) | (16 ns) (19
ceived July 14, surroundings. | CR) (2 Ni)
TOMPE Wie ead || —————— BS
of Scarborough. Dene 2 20 4 1 3 30

_ Dink af) (2n.) | (2m)
surroundings. | (1f,) (2)

Company of 18 | 7
larve (11 died), Licht | (8 2.) | (3 2.)
received July 15,|. ea
fromTorquayfrom S¥*roundings.

A. E. Holdway. |

Company of112) Light | | eee arse 123 29 36
larve (21 died), surroundings. | (1”.) | (11 m.) | (12 2.)
received July 15, —— SSS SSS ee ras =
from Wo Waren Of) Dark Meneame yo.
Cambridge. surroundings | ; f)

gs.
| pekia) | | 2/)

Adjustment of colowr in various pupe, ete. 427
Companies of Conditions of Degrees of Pupal Colour.
Larvie. Experiments. (ld) (2) (3) (4) (6) Totals.
C 5 : ade nce,
me aes pe ee Light 13 11 IY ||) Reka 123
July 18, from H, |SUtFoundings. oe ay eee
W. Head of Sear- |—— Coee ee ee) (27)
borough. Dutk 10 50 1 1 62
surroundings. (2 2.) (1 m.) | (3 1).
(_yf.) (1f)
Company of 46 | coat
larve (2 died), ee
received July 19, | wes i i 9 2 +
Fram ti. Wo kload surroundings.| (1 7.) (3 n.) (2 n.) (2 n.) (8 n.)
of Scarborough. |
Company of 139 . 1 7 8 5 17 38
: Fe Light » ‘
ee faa surroundings. (7f) 5 ie (2 2.) C nu.) | (5 n.)
from H. W. Head — Z aS Rees ee
of Scarborough. Wark Gi ' 32 19 4 1 64
ie : n. 5 n (len: (1 2.) | (8 7.)
gurnanes) (if) | Ger (3B aN Ger) (2 f)
Company of 25 Licht 1 1 ee Sly
yay 21 Received’ surroundings. a a fi bs
’ - . C
W. Head of Scar- $$ | —_—_|—_-——_ ee Sey ees
borough. Conflicting 1 2 5 8
surroundings.
Company of 70
Pave ee deh Dark a 6 3 2 62
from H fa eae surroundings. | (4n”.) | (4m) | (0m) >} (2) | (1)
of Scarborough. |
; Degrees of Pupal Colour.
Totals. () 2) (3) (4) (5) Totals.
Companies ex-| Light | : - ie ) oe ) oe ) on )
aa on in ee | Of) (6 f.) (47) | (Of) | (29,f)
Total number of | - ‘ ‘ ‘
larve, 1302 (180| Dark 1 Zs un oe 22 ate
ea bea omadines: (5n.) | (Ql 2.) | (15%) | (7 2.) | (12 n.) | (60 x.)
a) (47) | (bs) | Co s-))| G75 | Guy oz)
Conflicting 1 2 5 8
surroundings.
3

These results prove the extreme sznsitiveness of V. %o, and they afford valuable and
very extensive confirmation of some of the results described in Trans. Ent. Soc, Lond.
1892, pp. 397—432.

428 Mr. F. Merrifield and Mr. E. B. Poulton on

I—EXPERIMENTS AND OBSERVATIONS UPON
THE SUSCEPTIBILITY OF CERTAIN LEPI-
DOPTEROUS LARVA AND PUPA TO THE
COLOURS OF THEIR SURROUNDINGS. (A.
H. Hamm and E. B. P.)

1. Stauropus fagi. Mr. W. Holland and Mr. A. H.
Hamm of the Hope Department, who are extremely
skilled and accurate observers, drew my attention last year
(1898) to the fact that the larva of this species differ in
tint according as they are reared upon beech or birch, and
that the colours are in each case such as to conceal them.

During the present year Mr. Hamm reared two
batches (from different parents) upon the same food-plant,
beech, but in other respects under very different conditions
as regards environment. One of the batches was reared
in a white tissue-paper lined rectangular glass case with a
perforated zine lid, the other in a similar case lined with
black tissue-paper. We compared the two, placing both
batches on white paper, on July 13, 1899.

There were 24 Jarvee in the batch reared in the white
case, and of these all but 2 were in the last stage and
mostly advanced init. All but 1 were much lighter than
larvee of this species usually met with in nature, and
some most markedly lighter. The other batch consisted of
14 larve, of which 10 were in the last stage (mostly
advanced), 1 in the last but one, and 3 in the last but
two. <All were very dark, and most of them far darker
than those generally met with in nature. It is clear that
this species is highly sensitive, and it would be of great
interest to repeat the experiments under conditions which
have been found in other larve to produce the strongest
effects. It is remarkable that such considerable results
followed from surroundings which were not apparently in
contact with the larvee (for these at any rate when they
were examined rested upon the twigs and not upon the
walls of the case). I do not think, in any of the previous
experiments with larve, that equal effects have been
produced in this manner ; and one is tempted to enquire
whether it is possible that the larve in earlier and
specially sensitive stages, did actually rest upon the black
and white walls of the cases.

2. Notodonta ziczac. My friend Mr. Arthur Sidgwick
showed me, in the summer of last year (1898), a larva of

Adjustment of colour in various pupe, ete. 429

this species which he had obtained from Populus alba, and
I was greatly struck by the remarkable lightness of its
tint. It was quite unlike any larva of the species I had
ever seen. Mr. A. H. Hamm informs me that he has
also observed great differences in the depth of colour of
this Jarva according to the food-plant upon which it has
been found. It is highly probable therefore that this
species is also sensitive, and searching experiments upon
it are greatly needed.

This case and the last are of high interest, inasmuch as
no larva at all closely allied to these two species has yet
been shown to be sensitive to the colours of its environ-
ment.

3. Amphidasis betularia. Mr. A. H. Hamm tells
me that he has again and again observed in nature the
wide differences between the colours of this highly sensi-
tive larva upon various food-plants, and that the differ-
ences are invariably in the direction of concealment. Mr.
Hamm's experience in the field is so wide and his
powers of observation so keen that the strongest confirma-
tion is afforded to the observations recorded in my previous
paper (Trans. Ent. Soc. Lond. 1892, pp. 359, 360).

4. Hupithecia pimpinellata. Mr. Merrifield informs me
that Mr. Nicholson (of Lewes) has noticed that this larva
appears in two forms—reddish-brown and green—which
correspond to the two forms of the seed-heads of Pimpinella
saxyfraga on which it feeds. Mr. Nicholson states that
the green larve are nearly always found on the green heads
and the others on the brown.

The power of colour adjustment is probably present in
a very high degree in the larve of the genus, and numer-
ous carefully conducted experiments are much to be desired.

5. Vanessa polychloros, I may also mention that a
pupa of this species which I found (July 1899) upon the
dark painted iron railing at the North Entrance Gate
of the Oxford University Museum, harmonised very per-
fectly with the surface from which it was suspended.

6. Vanessa antiopa. In August 1897 I found several
living pupe of this species attached to fences, buildings,
etc., at the Hunt Club, Scarborough Heights, Toronto. I
specially noticed that there was a marked resemblance
to the environment. This was all the more noticeable
inasmuch as the colours differed very widely, some surfaces
being very dark and others very light.

430 Mr. F. Merrifield and Mr. E. B. Poulton on

K.—OBSERVATIONS ON THE COLOUR-RELA-
TION BETWEEN A COLEOPTEROUS SPECIES
(Cleonus sulcirostris) AND ITS SURROUNDINGS.
(W. HoLuanp and E. B. P.)

The Rhynchophorous species Cleonws sulcirostris 1s
described as possessing very variable markings (Fowler,
British Coleoptera, London, 1891). Mr. W. Holland has
recently found it upon the red sands of Boar’s Hill near
Oxford, and it is most interesting to observe that all the
specimens are reddish-brown in colour, entirely different
from the grey forms found by him on the sand-hills at
Deal, and from the darker grey ones which he finds on
“hotover Hill also near to Oxford and only a few miles
from Boar’s Hill. There are yellow and red sands on the
top of Shotover, but Mr. Holland has as yet only searched
for this species in localities where they are not exposed
on the surface. There have been no exceptions to the
colours of the very large numbers found on both hills. It
is reasonable to suppose that these colonrs, which certainly
harmonise with the ground of each locality, are protective ;
inasmuch as the species possesses in a very high degree
the instinets which lead to concealment.

So far as Iam aware this is the first time that such
loeal adaptation of colour has been shown to occur in a
Coleopterous insect, and the interesting question arises as
to whether the species possesses the power of varying its
colour during growth according to the stimulus provided
by the colours of its surroundings, or whether the results
are due to the varying operation of natural selection in

different localities leading in each case to the survival of

the individuals which are best concealed.

It will be of the highest interest to look for further
examples in Coleoptera, as well as to attempt to ascertain
the manner in which the colour adaptation is brought
about.

eS

Adjustment of colour in various pupe, ete. 431

L.—APPENDIX.

THE QUALITY OF LIGHT FROM THE COL-
OURED AND OTHER BACKGROUNDS
EMPLOYED IN THE EXPERIMENTS RE-
CORDED IN THE PRESENT MEMOIR. (Sir
JOHN Conroy and E, B. P.)

My kind friend Sir John Conroy, F.R.S., again helped
me to make a correct analysis of the light reflected from
the backgrounds employed in our experiments. The
papers, etc., were examined in the Laboratory of Balliol
College, Oxford, on July 24th, 1899. The beam from an
electric arc was passed through a bisulphide of carbon
prism, and the spectrum thrown on a white paper screen
in a darkened room. ‘The coloured papers, etc., were held
so as partly to cover the spectrum, and sometimes two
were held in the spectrum side by side for purposes of
comparison. ‘T'he method was thus the same as that
made use of on the previous occasion (Trans. Eut Soc.
1892, /. c. pp. 459 et seqq.), except that the lime-light
illumination was then employed, and our spectrum was
therefore weaker at the blue end than with the electric
are. The results obtained are recorded below.

1, Coloured backgrounds, etc., employed by F. Merrifield :—

Black net and black tisswe-paper (used as a plane surface
and also rolled round sticks). The reflected light from
both gave a feeble continuous spectrum. The trans-
mitted light was the same but still feebler.

Dark sticks also gave a faint continuous spectrum:
cork carpet was similar except that the reflected red was
prominent.

Orange tissue-paper (used as a plane surface and also
rolled round sticks). Some absorption of green and more
of blue and violet; absorption more marked in the sticks
where there was more than one thickness of the paper.
No appreciable difference between the reflected and trans-
mitted spectrum of the paper. The spectrum was far
more like that of a yellow paper than of the deep
orange used in my experiments, which removed everything
except the red, orange, and yellow.

Orange leno gave a very similar spectrum, the blue and
violet being almost cut off, and the green a little darkened.

432 Mr. F. Merrifield and Mr. E. B. Poulton on

The absorption was more marked in the transmitted
light.

Fale tissue-paper (also rolled round sticks). Violet
cut off and blue much diminished ; hardly any blue in the
transmitted light, and in the thicknesses of paper rolled
round sticks.

Golden yellow oat-straw gave a very similar spectrum,
the blue and violet being absorbed and the rest
unaffected.

Unfaded bright yellowtsh-green tissue-paper (also rolled
round sticks). The red shortened and dimmed; the blue
and violet much absorbed. Transmitted light similar, as
also the reflected light from the paper round sticks.

Bright yellowrsh-green art muslin. A similar spectrum.
Not much difference between transmitted and reflected
light, except that two thicknesses produced far greater
effects in the former.

Dull green reeds. Whole spectrum somewhat weakened,
the blue most and then the red.

Yellow metallic surface of brass (Dutch gold), also rolled
round sticks. Appears to give a strong continuous spec-
trum, but the yellow colour is due to absorption of the
blue end.

Dead reeds. The lightest of them gave a typical yellow
spectrum with absorption of the blue end only; in the
others there was diminution of all other regions, although
the blue end was still most reduced.

Turned cylindrical wooden sticks (probably deal). A
very similar spectrum; the blue end was still more
absorbed than any other part, although less so than in the
dead reeds.

Dirty white paint gave a very similar spectrum with
some absorption throughout, most in the blue, least in the
green.

White paper rolled round sticks, white calico, shaved white
sticks all gave a strong continuous spectrum with no se-
lective absorption.

Yellow glass (ased as screen). Blue and violet cut off,
the rest unabsorbed.

Deep green glass (used as screen). The green almost
unabsorbed. The whole of the blue end, and nearly all
the red and orange absorbed.

II. Colowred backgrounds employed by E. B. Poulton and
those who worked with him.

Adjustment of colour in various pups, ete. 433

The Black papers, both “surface papers” and_ tissue-
papers, gave the same results as Mr. Merrifield’s, and
those previously recorded (E. B. Poulton in Trans. Ent.
Soc. Lond. 1892, pp. 461—464), and the same was true
of the yellow papers (surface and tissue) and yellow leno,
all of which were typical; the white paper, and the white
opal glass. The deep orange paper was similar to that I
have previously used, and very different to Mr. Merrifield’s.
The following backgrounds had not been examined
before :—

A dull surface of metallic zinc (perforated) gave a dim
continuous spectrum (general, but no selective absorp-
tion).

Salmon pink surface paper. A very faint absorption of
the blue was all that could be seen. The blue also looked
redder, an effect which may have been due to the stray
white light reddened by selective absorption in the
paper.

Violet paper. Much of the blue end, and considerable
red were reflected. There was an absorption band be-
tween the green and the blue, and the yellow, orange,
and green were much absorbed.

28 a

*

Extracted from the PROCEEDINGS OF THE ENTOMOLOGICAL
Soctety oF LONDON of December 7th, 1898.

Dr..Drxey exhibited a series of Pierid butterflies from the
Neotropical region to show the existence among them of
seasonal forms. The species shown were Parura rurina,
Feld., ¢; P. neocypris, Hiibn., 6; Phabis argante, Fabr.,
36, 9; P. agarithe, Boisd., 3 ; Callidryas senne, Linn.,
6, 23; C. philea, Linn., g. In each case three specimens
were shown, exemplifying the “wet season,” ‘dry season,”
and ‘intermediate ” forms of the species. These had been
selected from the Hope Collection by permission of Professor
Poulton,

With reference to the exhibit, Dr. Dixrty remarked that
although direct evidence as to the seasonal changes in these
Neotropical species was at present scanty, the indirect evidence
was strong ; inasmuch as the range of variation here shown
was analogous with that existing in certain African and
Oriental species, whose seasonal relations had been put on a
firm basis by the work of Watson, Barker, Marshall, and
others. In every one of the present instances the ‘‘ wet
season” form tended to be more deeply coloured, and to have
its markings on the underside more fully developed ; while
the “dry season” form was usually smaller, had its forewings
more sharply pointed, and was nearly or entirely devoid of
markings on the underside. He stated further that where
evidence of the date of capture existed, it accorded very fairly
with the supposition that these variations bore relation to
seasonal conditions, though of course more data of this kind
were much needed. He had selected the specimens of each
species so far as possible from the same locality, endeavouring
thus to meet the possible objection that these varieties, several
of which had been described as distinct species, might have a
local rather than a seasonal significance. He added that by
an “intermediate” form he meant simply one that came
somewhere between the other two—not necessarily half-way.
He was accustomed to use the term in a general sense as
equivalent to the German “ Zwischenform,” not as conveying
the idea of an arithmetical mean (Mittelform).

The Prestpent observed that the exhibit was of special
interest, as affording the first recorded evidence of the exist-
ence of seasonal dimorphism in Neotropical butterflies.

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RECEND EXPERIMENTS IN HYBRIDISATION,

T has long been recognised that the facts of hybridisation
have an important bearing on many subjects of biolo-
gical interest. The question of the infertility of crosses was
very carefully investigated by Darwin, and since his time
has been more than once discussed afresh ; while the struc-
tural and other characters presented by hybrid offspring
are of undoubted weight in reference to the problem of
heredity.

For some years past Dr. M. Standfuss, of Ziirich, has
been experimenting ona large scale in the production of
insect hybrids. The results of these experiments, some of
which confirm previous views, while others appear to sug-
gest conclusions not hitherto reached, have now been collec-
ted by their author in his Handbuch der paliarktischen
Gross-Schmetterlinge, published at Jena in 1896. Many of
the data thus made available derive especial value from the
fact that they are expressed numerically, and the whole
series forms a noteworthy contribution to our knowledge of
the subject. It is proposed to give here a short account of
these experiments, with their results, for the benefit of those
readers to whom the original records may be difficult of
access."

The author prefaces his account with a list of fertile and
infertile pairings between different species of macrolepi-
doptera that have been observed by himself and others,
both under natural conditions and in captivity. Details are
first given of twenty-four cases of pairing between Lomébyx
neustria L. 6 and B franconica Esp. %. The results
showed every transition between complete absence of issue
and the deposition of eggs normal in numbers and fertility.
Failure was in some instances plainly due to inadaptability
of the genital apparatus. On this series of pairings Stand-

1Several of Dr. Standfuss’s specimens have lately been exhibited in
London, at the rooms of the Royal Society, Burlington House and at the
Natural History Museum, South Kensington.

2 SCIENCE PROGRESS.

fuss remarks that the absence of uniformity in the results
shows that several trials should be made before any given
cross is pronounced infertile.

Five crosses are mentioned from which only male off-
spring are known to have resulted; these are Dezlephila
porcellus L. 3 and D. elpenor L. 2 ; Smerinthus austautd
Steger. d and S. atlanticus L. 2? ; Fumea nitidella Hof. S
and /. affints Rutt. 2, with the reciprocal cross /. affints
3d and F. nitidella 2 ; Bombyx neustria L. $ and B. fran-
conica Esp. ?.

In the following five cases only female offspring have
been observed, and these contained no eggs capable of de-
velopment : Lombyx neustria L. S and B. castrensis var.
veneta Sidfs. $ ; LB. franconica Esp. & and B. castrensis
var. veneta % ; B. quercus L. 3 and B. trzfoliz Esp. ¢ ;
Saturnia pyrt Schiff ¢ and S. pavonia L. 2 ; Drepana
curvatula Bkh. 3 and D. falcataria L. ?.

In the following seven the males predominate and the
females are again sterile: Dezlephila euphorbie L. 3 and
D. vespertiho Esp. 2; D. hippophaes Esp. & and D.
vespertilio Esp. 2 ; Smerinthus ocellatus L. 3 and S. popula
ie 2 ; Saturnia spent Schitf 3 and S. py77 Schitt 25s:
spint Schiff 3 and S. pavonia L. ? ; Harpyia vinula L. S
and /7. erminca Esp. ? ; Notodonta dromedaria L. & and
LV. torva Hb. ¢.

The author points out that no fertile female offspring
having been observed in any of the above seventeen crosses,
none of these hybrids, so far as our present knowledge goes,
could continue their race. It is unfortunate that the actual
numbers of the broods in question are not given; in some
instances, as in that of B. neustria S and B. franconica &
they are presumably large.

In three more crossings the offspring consisted of both
sexes in normal proportions; the females, however, con-
tained at best but few eggs, and these abnormal in structure.
The crossings were of Smerinthus populi L. & and S.
ocellatus L. ¢ ; Saturnia pavonia L. 3 and S. pyr Schiff ¢ ;
S. pavonia L. S and S. spzeuz Schiff ?.

Finally, in two more the sex-distribution appears again

RECENT EXPERIMENTS IN HYBRIDISATION. 3

to be as usual, and the females contain ova normal in
appearance. Their fertility, however, has not been proved,
and Standfuss thinks he has reason to doubt it. These are
Zygena trifolit Esp. 3 and 2. filipendule L. ¢ ; Biston
hirtartius Cl. & and B. pomonarius Hb. ¢.

In no single instance, according to Standfuss, has the
female of any true hybrid among lepidoptera been shown
experimentally to be fertile. Haeckel’s counterstatement
with regard to the genera Saturnia and Zygena is probably
erroneous. '

The most complete series of Standfuss’s experiments in
hybridisation is one carried on for over ten years with three
species of Saturnia, viz., S. pavonia L., S. spent Schiff, and
Sarzyee och... Vhe results) of these. experiments ate
recorded and compared by their author with great minute-
ness, and excellent figures are given by him of several of
the resulting forms in their immature and final stages. A
brief outline of these records, omitting details, will here be
attempted. |

1. Saturnia pavonia S and S. spint ¢. In this hybrid,
called by Standfuss S. dornemannz, the eggs first laid were
regularly deposited, and were fertile to the extent of from
60 to 80 percent. The larve in all five stages bore a much
closer resemblance to those of S. sfzzz than to those of SS.
pavonia, though this became less pronounced after the
second stage was passed. The cocoon and pupa were both
intermediate in structure, but the perfect insect was nearer
to S. sfzuz than to S. pavonza. This applies to both sexes,
but is more easily seen in the male, the males of the two
parent species differing (as is usual) more than the females.

2. The reciprocal cross (S. sfenmz ¢ and S. pavonia ¢ ;
= S. hybrida Ochs.) has not been produced in captivity,

1 Standfuss does not notice the case cited from Quatrefages by Wallace
(Darwinism, 1889, p. 163), of fertility zz¢fer se in the hybrids between
Bombyx cynthia and £B. arrindia. Fletcher also has obtained fertile
hybrids of both sexes between Zygena lonicere Esp. and Z. trifolit (Proc.
Ent. Soc. Lond., 1893, p. ix.) On the other hand, the hybrids of
Z. lonwere and Z. filipendule proved infertile (Zézd., 1891, p. ix.), and the

hybrid progeny of various species of Platysamia and Actias were found by
Miss Morton to be sterile in both sexes (Zé:d., 1895, p. Xxxiv.).

4 SCIENCE PROGRESS.

but is repeatedly found in the wild state, generally as a
larva. The question naturally suggests itself, how the
parentage of this hybrid is known. Standfuss replies that
the larva, though intermediate in character, shows constant
differences from that of S. d0vnemannz, and that the condi-
tions in nature for the pairing of S. sfzmz 3 and S. pavonza
? are favourable, whereas those for the converse cross are
difficult or impossible. The males of both species emerge
before the females, and in regions where both occur SS.
pavonia is out first. Hence, though the females of 5S.
pavonia are out with the males of S. sfzzz, the converse does
not take place. It must be allowed that it would be well to
test this conclusion as to the parentage of S. Lybrida by
experiment. The larva of S. Zyérzda resembles S. pavonza
somewhat more than does that of S. dornemannz; it pos-
sesses, however, the greasy polish of S. sézuz to a greater
extent than the latter. The cocoon and pupa both show a
nearer approach to S. spzuz than do those of S. dornemannz,
and the same applies to the perfect insect; in fact the
resemblance of both sexes to the male parent is remarkably
close.

3. S. pavonia S and S. pyvt %. This crossing pro-
duced hybrids which fell into two classes, a dark form
called by Standfuss S. dauézz, and a paler form to which he
gives the name SS. ewzlze. The latter is by far the com-
moner. As in former cases, if the laying is much deferred
after pairing, the eggs are apt to prove infertile. The larva
in its early stages is very like that of S. Ayvz; it becomes
more and more like that of S. Aavonza, and finally bears a
close resemblance to the latter species. The cocoon is
intermediate ; the pupa is nearer to S. Aavonza than to S.
pyrt. The perfect insect, except for a reduction in the
sexual disparity of size, is more like an enlarged S. Aavonza
than a diminished S. pyvz. In a majority of specimens some
of the veins were forked terminally at a greater or less dis-
tance from the margin of the wings.

The males of this hybrid paired readily with the females
of S. Zavonia ; they were also attracted by the female hy-
brids, though in a less degree. The latter were, as already

RECENT EXPERIMENTS IN HYBRIDISATION. 5

stated, infertile, the oviducts containing no ripened eggs.
Pairing also took place, though with much less readiness
than in either of the preceding cases, between the hybrid
males and some females of S. Ayrz. The number of fertile
egos produced in the latter case was far smaller than in the
cross with S. pavonza.

4. S. dornemanni 3 and S. pavonia ? (S.bornemannt is the
cross-product of S. favonza $ and S. spenz ¢). The larvee
at first closely resembled those of S. favonza, but in subse-
quent stages the influence of S. sfzuz began to assert itself.
They presented a very variable appearance on reaching the
fourth stage, which period unfortunately they did not sur-
vive. Fresh batches, however, have since been raised to
maturity, though no record of their history has yet been
published.’

5. S. emtle & and S. pavonia 2 ( S. emili@ is a cross-
product of S. pavonia $ and S. pyvz 2). The larve of this
hybrid (named S. stanxd/usst by Wiskott) are again very
variable. Their general appearance is that of a large S.
pavonia. The cocoon and pupa are also near the same
parent species. The perfect insect, like the larva, is vari-
able ; it always, however, shows much resemblance to 5S.
pavonia, and the sexes are dissimilar as in that species.
The margins of the wings are apt to be scalloped. The
oviducts of the only female that emerged contained mature
eggs, but only about twenty, or one-tenth of the normal
number in S. pavonza or S. pyre. It is possible that these
hybrids may be fertile zzzer se.

6. S. emilie & and S. pyri 2 (S. rvesez Stdfs.). This
pairing was only obtained with great difficulty, and in four
cases out of nine the eggs, though laid in normal numbers,
did not hatch. In the remaining five broods only one per
cent. produced caterpillars. These at first closely resembled
S. pyri, and in the second stage still showed more likeness
to S. Ayvz than to the male parent form. In the third stage
the S. pyvz characters began to be lost, and in the fourth

! Specimens of this hybrid have been reared by the present writer from
eggs kindly sent him by Dr. Standfuss in 1895. Other individuals raised
from the same batch of eggs are in the Hope Collection at Oxford.

6 SCIENCE PROGRESS.

those of S. Ayrvz and S. pavonza were fairly balanced. After
changing its last skin the larva resembles a large S. avonza,
though its parentage is three parts S. fyrvz. The perfect
insects, of which only six were reared, were in many respects
remarkable. Unlike S. Ay7z they were sexually dimorphic,
the females differing little from those of .S. evzzlze, while the
males showed a nearer approach to S. fyvz. The females
were not dissected, but were probably sterile. Of the six
specimens, one was hermaphrodite and three others showed
a tendency in that direction. These four were the produce
of three separate females out of the five that laid living ova.
Standfuss draws attention to the fact that the normal oc-
currence of hermaphrodites in lepidoptera is given by Speyer
as about 1 in 30,000. The latter number he considers
rather too low than too high. There was no hermaphro-
ditism in any known member of the families of the progeni-
tors of these hybrids ; and its appearance in this proportion
must be, he thinks, a consequence of their exceptional origin.
On the other hand, he has bred more than 1000 genuine
hybrids without one such case occurring,’ nor was any such
tendency shown by a single one of the sixteen specimens of
S. standfussi—a form whose origin is analogous to that of
the present cross-product.

In the well-marked hermaphrodite mentioned above, the
distribution of sexual characters is remarkable. The shape
of the fore wings is rather female than male, the colouring on
the upper side of both is male; on the under side, the right
is mostly male and the left female. In both hind wings the
upper surface has the costal portion male, the remainder
female ; the right, which is about one-fifth larger than the
left, has the female area more extensive. The under
surface of the right or larger hind wing is female, of the
left mostly male. The right antenna is male in form; the
left partly male and partly female. The external genital
organs on the right side are of a malformed male type; on
the left side absent.

1 Barrett (Lepidoptera of the British Islands, vol., il., 1895, pp. 10,
11) gives some details of the cross between Smerinthus ocellatus and SS.
popult. These are said to be “ often gynandrous”’.

RECENT EXPERIMENTS IN HYBRIDISATION. 7

The want of vigour in this cross is shown by the fact
that out of nine pairings, each resulting in an average of
200 eggs, only ten larvae were produced ; and of these only
Six, as we have seen, attained the perfect condition.

7. S. pavonia 3 and Actias luna L. ¢. Nine apparently
normal pairings took place, and over 1000 eggs were laid ;
but none of these hatched.

8. S. pavonia & and A. zsabelle Graélls ?. As a re-
sult of this crossing, ninety-eight eggs were laid, seven of
which hatched. The caterpillars however did not long
survive their first change of skin.

9. S. dornemannt S and S. pyrt 2. As S. bornemanut
is the cross-product of S. Aavonza S and S. spinz ?, this
hybrid is descended from all three species.‘ Only one
pairing was obtained, and 92 per cent. of the eggs hatched.
Some of the resulting perfect insects were lately shown in
London with the other exhibits of Dr. Standfuss, who
promises to publish further details of their history.

Relative phylogenetic age of the three European species of
Saturnia. Before stating the general conclusions at which
he has arrived as a result of the above series of experiments,
the author proceeds to discuss the question of the relative
phylogenetic age of the three species Saturnia spinz, S.
pavonia and S. pyrz. Of these he considers S. sfzuz to be
the oldest form and S. Ayzz the most recent. His main
reasons are briefly as follows :—

(1) The larva of S. sfzzz maintains its original black
colour throughout its life. .S. Aavonza loses this sometimes
in the third stage, always in the fourth and fifth. S. pyrz
abandons it almost completely in the third stage and on-
wards. The succeeding green colour, which is no doubt
adaptive, is acquired by S. pyrz at an earlier stage and more
completely than by S. Aavonza.

(2) In the larva of S. spzzz the tubercles are not very
prominent even in the adult, and the knobs at their summit
are not distinctively coloured until the last stage. In S.

.1 Mr. A. G. Butler informs me that an analogous triple cross has been

obtained between a goldfinch and the hybrid issue of the English and
Japanese greenfinch.

8 SCIENCE “PROGRESS.

pavonia the tubercles are conspicuous and the knobs acquire
an appearance distinct from the general surface of the body
in the third or fourth stage. In .S. Ayvz on emergence from
the egg the knobs are already coloured and indications exist
of the extreme prominence of the tubercles.

(3) The tactile bristles are least developed in S. spznz,
most in S. pyvz.

(4) The cocoon of S. sfzzz is simpler than that of 5S.
pavonia. That of S. pyrz is the best defended of the three.

(5) S. sfznz is almost sexually monomorphic, though
the male is somewhat the smaller. The female is very
sluggish, and the antennz of the male are pectinate to a
high degree. In .S. favonza there is well-marked sexual
dimorphism, as regards both size and colouring. The
female resembles that of S. sfzzz in aspect; it is less
sluggish, though not very active. The antennz of the
male are less strongly pectinate. .S. Ayrvz again is sexually
monomorphic ; the female is a tolerably good flier, and
the antennez in the male are less strongly pectinate than
in many other species of Saturnza.

Judging from the perfect insects alone, we must con-
clude, according to Standfuss, that the less specialised .S.
spinz is older than S. Aavonza. We should, however, find
it difficult to determine whether S. pyvz arose before or
after the divergence of these two. The larval and pupal
stages enable us to answer the question. The three species
form a progressive series in protective adaptation, S. spzzz
always taking the lowest and S. pyvz the highest step in
the scale. It must therefore be concluded, on the whole
evidence, that S. sfzzz is phylogenetically the oldest, and
S. gyri the youngest of the three forms.

General conclusions with reference to hybridisation be-
tween distinct species. As the author elsewhere expresses
it, the crossing of two distinct species gives rise to a
‘“ Zwischenform ” but not to a “ Mittelform”. The Mittel-
form may, however, exist as a temporary stage in larval
growth. This depends on the following principles, which
Standfuss considers to be warranted by the above experi-
ments with species of Saturnza :—

REGEN exer RIMEINTS IN HYBRIDISATION: 9

1. The freshly-hatched larva closely resembles the female

parent.
2. With the process of growth a resemblance to the male

parent gradually increases.

3. The final extent of approximation towards the male
parent depends on the relative phylogenetic age of the two
species ; the older being able to transmit its properties, whether
of structure or habit, better than the younger.

Thus the crossing of S. pavonza 3 with the phylo-
genetically younger S. pyrz ? gives rise to a larva in which
at first the maternal and afterwards the paternal characters
predominate.” The resultine’ perfect inseet is by more
than two-thirds of its external appearance S. favonza, and
by less than one-third S. Ayvz. Its habits and functions
correspond with its external aspect. It prefers to fly by
day, like S. pavouza 3, and pairs easily with the female of
that species, from 43 to 62 per cent. of the eggs being fer-
tile. On the other hand, it does not pair readily with S.
pyri, and the resulting eggs on an average of nine cases gave
only one larva from 180.

Similarly S. Aavonza 3, when paired with the phylo-
genetically older S. sfzmz 2, gives a form of which in the
perfect state about two-thirds of the external aspect belong
to the type of S. sfzzz. The male flies by night. After
crossing with S. pavonza ¢ the resulting eggs were only
fertile to the extent of 16 to 22 per cent.; while the crossing
with .S. sfenz ?, though not easily brought about in con-
sequence of their diverse times of appearance, yielded eggs
of which from 94 to 98 per cent. were fertile.

Thus the male S. Aavonza is able to influence the issue
of the relatively gigantic S. Ayvz ? much more than that of
S. Spint 2.

Again, the issue of S. spzuz o and S. favonza ? is much
nearer .S. sfzuz than is that of S. pavonza 3 and S. spzni &.
Hence,

4. In reciprocal pairing the male is able to transmit the
characters of the species in a higher degree than the female.

1 Barrett (doc. cit., p. 11) says, quoting Porritt, that the larvee of the
cross between Smerinthus ocellatus and S. populi, ‘though like those of
S. popult when young, appear to become intermediate, or even to re-
semble those of .S. oced/atus when fuli grown ”.

10 SCIENCE PROGRESS:

The above cases show that the former influence, v2z.,
that of the older-established species, is the more effective of
the two. The sexual prepotency of the male S. pavonza
counts for less than the specific prepotency of the female S.
spint. The highest effect of course is produced when the
two influences concur, as in the hybrid of S. spzuzz 3 and
S. pavonia ¢.

A further result of the experiments is that while no
female hybrid was proved to be fertile, there are undoubted
cases of fertility in male hybrids. This has been shown by
crossing with the females of both parent species, and in one
case with a female of a third species (S. dornemannz 3 and
S. pyri ?).

Hence hybridisation is not necessarily a merely tran-
sitory phenomenon. There seems to be no reason why a
male hybrid should not propagate by ‘‘ back-crossing ” under
natural conditions; and since the product of this kind of
crossing is not found to show a complete reversion to the
type of the female parent, it is possible that the existence of
various intermediate forms in such genera as Melilea,
Zygena and Agrotzs may be accounted for in this manner.
Cases of simple pairing between distinct species of the two
former genera have been observed by the author in nature.

The above conclusions with the facts on which they
rest, though much condensed from the original, have been
stated as far as possible in Dr. Standfuss’s own words. A
few brief comments seem called for.

(1) The rule laid down by Standfuss as to the pre-
potency of the phylogenetically older species is probably
another expression of the fact so clearly established by Dar-
win! that hybridisation frequently leads to reversion. It is
significant that Standfuss considers the hybrid form S.
emilue as partly reproducing an ancestral stage in the
history of S. Aavonza rather than the form of that species
at present existing.

(2) It is well known that when plant hybrids are crossed
with one of the parent species, considerable variability may

1Animals and Plants under Domestication, 1868, vol. ii., p. 254, etc.

RECENT EXPERIMENTS IN HYBRIDISATION. “x

occur in the offspring (Focke, quoted by Weismann). The
facts above recorded as to the results of back-crossing (see
accounts of S. standfussi, S. visit, etc.), show that the same
rule holds good with hybrid Saturuzas.

(3) Some of the above generalisations, at present rest-
ing on only a few instances, can hardly be regarded as more
than provisional. It will be interesting to see whether the
further results of these experiments, which are being con-
tinued by their author, tend to confirm or to modify the con-
clusions previously reached.

Cross-breeding between local races. Akin to the above
experiments are others in which local races of the same
species were paired. These gave results in many respects
analogous with the foregoing.

1. Callimorpha dominula L. & and do. var. persona
Hb. ?. The crossing of C. dominula 3 with var. persona
¢, the latter being a local race found only in Tuscany and
Calabria, generally produced issue that, though very variable
in the perfect state, bore on the whole a closer resemblance
to C. dominula than to var. fersona. Inone case, however,
a majority of the brood came nearer to the latter, and one
individual even went beyond the persona type. From 3
to 5 per cent. of the eggs were sterile.

2. C. dominula var. persona 3 and C. dominula 2. In
this, the reciprocal cross, the resulting perfect insects again
varied between the parental types, but, on the whole, came
nearer to C. domznula than to the variety, though less so
than those of the former cross. Of the eggs, from 10 to
15 per cent. were sterile, from which Standfuss concludes
that the male of var. Persona has already diverged from the
physiological standard of the species.

The products of each crossing are fertile in both sexes,
but Standfuss is unable to say whether they have undergone
any diminution in fertility as compared with the parent
forms.

3. Ocnogyna hemigena Grasl. 3 and O. zoratda Grasl. °.
These are regarded by the author as local races of the same
species, the former inhabiting the Pyrenees and the latter
the mountains of Andalusia. The cross-product (called by

12 SCIENGE PROGRESS:

Staudinger O. zoragena) resembles a large O. hemigena.
According to Kréning the mongrel issue are fertile zxZer se,
but quickly degenerate in size and vitality. This of course
may be due to other causes than their mixed origin.

4. Spilosoma mendica Cl. $ and do. var. rustica Hb. ¢.
The larve were nearly always formed within the egg ; but in
some broods not one, and in others only from 8 to 12 per
cent. hatched. In one case, however, as many as 93 per
cent. of the eggs gave living larva. All the broods suffered
severely from disease. The perfect insects, which did not
show much variation, diverged only slightly in appearance
from var. rustica. The males were mostly light-coloured,
as in that variety, and the darkest of them were far lighter
than any male of S. mendvca.

5. 8. mendica var. rustica 3 and S. mendica 2. The
reciprocal cross, which failed with Standfuss in consequence
of an epidemic among the larve, was obtained by Caradja.
Pupz sent by him to Standfuss gave thirty-one perfect in-
sects, seventeen being males and fourteen females. These
were more variable than the former cross-product, but on
the whole inclined towards var. rustica.’

It is observable that in this case of reciprocal crossing,
the influence of the females appears to be equal to if not
greater than that of the males. This shows that the
generalisation as to male prepotency founded on the case of
the Saturnzas is not of universal application. It will be
remembered also that the issue of one of the pairings
between C. domznula S and var. persona 2 showed female
prepotency, though the general result in the case of this and
the reciprocal cross tended in the opposite direction.”

Standfuss considers that the present experiments with
species and their local races favour his view as to the
superior influence of the phylogenetically older form.

Aberrations. Some very remarkable facts are recorded

1This corresponds, as has been pointed out by Mr. South, with the
result obtained by Mr. Adkin, who effected the same cross in 1889. Vid.
Entomologist, for August, 1897, p. 206; Proc. Ent. Soc. Lond., 1890,
p. xl.

2 See also p. 199, zz/ra, note.

RECENT EXPERIMENTS IN HYBRIDISATION. 13

as to the effect of crossing a sport or aberration with its
parent form. The result, which is entirely different from
that which follows the crossing of distinct species, or even
of local races, may be broadly stated as follows: When an
aberration is crossed with its parent form the issue is
sharply divided, in both sexes, into specimens of the aberra-
tion and of the normal form of the species.

Thus in the dark aberration zatima Cr. of Spilosoma
lubyicipeda Esp., there are many degrees from the least
dark form of the aberration (ab. zv¢termedia Bang-Haas) up
to the darkest (ab. deschangei Depuis); but no transitional
forms occur to bridge over the wide gap between zntermedia
and /ubricepeda, nor can they be produced by crossing these
two. “It seems,” so Standfuss expresses it, ‘‘as if there
were antagonistic characters which cannot coexist in the
same individual ”.' Instances follow which will be briefly
noticed here. For full details, which are of great interest,
the reader is referred to the Handbuch, pp. 305-321.

1. SAzlosoma lubricipeda Esp. 3 and do. var. zatima
Cr. 9. These, crossed by Burckhardt in 1889, gave /udrict-
peda, intermedia and zatima (intermedia being, as just stated,
merely a less dark form of zatzma). ‘Two of these zxtermedia
were paired, giving again Zubricipeda, intermedia and zatima.
In this third generation several pairings were effected, as
follows: zatzma 3S and lubricipeda ? ; lubricipeda S and
zatima 2; intermedia 3 and do. 3; tntermedia S and
zatima ¢. <All these gave “4ubrictpeda, intermedia and
zalima in varying proportions, except the cross Zubricipeda
d and zatzma ¢, from which only za¢zma resulted. A pair
of Zubricipeda from this fourth generation gave a brood of
34 lubricipeda and 1 extreme zatima.

In all these successive broods, carried on into the fourth
year from the date of the original pairing, there were no
transitional forms between ludrviczpeda and intermedia.”

‘The bearing of this may be noted on the theory of ‘‘ determinants ”.
See Weismann, Germ-flasm, 1893, pp. 293, etc. See also Bateson, JJa-
terials for the Siudy of Variation, 1894, esp. Intro., Sect. iii.; and this
Review, 1897, pp. 554-569.

* The above case of cross-breeding between colour-varieties of the same
race, in which there are no intermediates and the colours of the ancestors

14 SCIENCE PROGRESS.

2. Psilura monacha L. 3 and do. ab. evemita O. 2.
In 1893 Standfuss raised a brood from a pair of normal
P. monacha received from Silesia. This brood contained
one female specimen of the dark aberration evemzta, which
was paired with a normal P. monacha 3 from Zirich. The
issue consisted of 22 (2 d and 20 ¢) typical monacha, 23
(18 3 and 5 ¢) typical evemita, and 6 forms (5 ¢ and
1 ?) in which the characters of the two are unsymmetrically
mixed, not harmoniously blended. There is no apparent
tendency to hermaphroditism. This unsymmetrical mixture
resembles that occasionally seen in the male of Ocneria
dispar 1.., where also, according to Standfuss, the patches
of the female pattern that sometimes appear do not betoken
organic hermaphroditism. |

3. P. eremita S and P. monacha ¢. This was a
natural pairing found by Standfuss in Silesia. The result
was entirely different from that of the former cross, inas-
much as the issue contained every kind of transition be-
tween the two parent forms, while a few even transcended
the male parent in darkness.

Thus the two specimens of so-called evemzta in this and
the former observation, though externally so much alike,
possessed entirely different properties in regard to the
power of transmission to descendants. Standfuss explains
the apparent contradiction thus: The first evemzfa was a
true sport or aberration, and in its case the rule held good
as usual. The second, also called “eremzta” (which did
not show the evemz¢a characters so well as some of its own
offspring), was a link in the chain leading by slight varia-
tions to a darker and presumably better protected form of
P. monacha, which, under the influence of natural selection,
is gradually developing itself in certain parts of the range

of 195 individuals are known for three generations, seems, like that of
Agha tau and its aberration /ugens (infra, p. 199), to promise suitable data
for the verification of Mr. Francis Galton’s law of heredity. According
to this law the sum of the ancestral contributions is expressed by the series
(0°5) + (0°5)7 + (0°5)2? +... (0°5)”, each term standing for a generation.
See Galton, Vatural Inheritance, 1889, p. 134, and ‘“‘ Average Contribution
of each Several Ancestor to the Total Heritage of the Offspring,’ Proc.
Roy. So¢., 1897, pp. 401-413.

RECENT EXPERIMENTS IN HYBRIDISATION. 15

of the species. It took rank therefore not as an aberration,
but rather as a member of a local race, and with this its
behaviour accorded.’

4. Agha tau L. & and do. ab. degens Stdfs. Like the
dark aberration zatzma of S. lubricipeda, the dark Agha tau
(called by Standfuss ab. /wgens) exists in several degrees of
development, from the ‘ab. /eve nigra” of Thierry-Mieg to
the “ab. xzgerrima” of Bang-Haas. All these different
stages of darkening have been obtained by Standfuss from
pairings between A. éaw and ab. Zugens, but no transitional
form between the normal A. éau and ab. fere nigra has ever
been so produced.

In 1888 Standfuss crossed males of ab. Zagexs, which had
been interbred for two generations, with females of the nor-
mal A. aw. In 1889 Zegens derived from these were em-
ployed in the following pairings: Zzgens S and tau ¢ ; tau
& and lugens 2 ; lugens S and do. %. The specimens of
A. tau were in each case of different ancestry from the
lugens stock. In 1890 two separate pairings were procured
of dugens 3 and ¢ from the third of the above broods, so
that the perfect insects emerging in 1891 had both parents
and all grandparents of the /ugens type. The results of
these experiments were curious, and should be studied in the
original account (//andbuch, p. 312). Here it may be
briefly stated that of the 1889 pairings, the first two (Zugens
Sd and ¢fau ? ; tau 3 and lugens ) produced about 50 per
cent. of each form.”

The third (4ugenxs 5 and $) gave 36 per cent. of tax to

1 Compare the result of crossing C. dominula with the local race
persona, and see especially Standfuss’s figures, Op. ce¢., Taf. V. Compare
also the case of A. éetudaria and its aberration doubledayaria (inf., p. 201),
which is strikingly analogous with that of P. monacha.

* In both of these reciprocal crosses there is a slight preponderance of
forms resembling the ma/e parent—in the first instance /ugens, in the
second ¢aw. ‘This case is fairly analogous with that of the breed of Basset
hounds lately investigated by Mr. Francis Galton (“ The Average Contribu-
tion of each Several Ancestor,” etc., Proc. Roy. Soc., 1897, pp. 401-413),
where the reciprocal crosses of two colour-varieties show a male prepo-
tency in the proportion of about six to five (doc. ci#., Table L., p. 409). As
in the instance of Zaz and /ugens, there are no intermediate forms between

16 SCIENCE PROGRESS.

64 per cent. of Zugens. In the 1890 broods, al] of whose
ancestors were /ugens for two generations, the proportion
of au had fallen to a little over 11 per cent. in one anda
little under the same figure in the other. Jn each of the
five cases about twice as many females as males were of the
tau form. Hence, as Standfuss puts it, it is more difficult
to transform the female of Agta ¢au than the male.’

5. Grammesia trigrammica Hufn. 3 and do. ab. dzlinea
Hb. ¢. The female éz/znea, taken by Gross at Garsten in
Austria, laid eggs of which the male parent was pre-
sumably a normal G. ¢vzgrammica. Of the sixty-seven per-
fect insects that resulted, thirty-eight were ¢vigrammuca
and twenty-nine dz/zzea. ‘There were no intermediates.

6. Angerona prunaria L. 3. and do. ab. sordiata
Fuessl. 2. This cross, procured by Zeller, gave seventeen
prunaria and fourteen sordzata.

7. A. prunaria ab. sordiata S and A. prunaria ¢.
This cross, also obtained by Zeller, gave eighty-four of
prunarva to sixty-eight of sordzaza, 2.e., as in the reciprocal
cross, about 55 per cent. of the type and 45 per cent. of the
aberration. In neither of these cases were there any inter-
mediates.

8. A. prunaria ab. sordiata 3 and ¢. Among a large
brood reared from the eggs of a pair of normal A. pru-
naria, there appeared three males and two females of the
aberration sovdiata. From a pair of these Standfuss ob-
tained thirteen prunaria (three 3 and ten ?) and forty-two
sordiata (twenty-four ¢ and eighteen ?). Again there were
no transitional forms.

9. Amphidasis betularia L. & and do. ab. doudbledayaria
Mill. ¢. A female doudbledayaria found by Steinert near

the “tricolour” and ‘‘ non-tricolour”’ hounds, though there are different
degrees of development of black in the ‘tricolours ”
parable with the range of variation between “‘ feve migra” and “ nigerrima”
in lugens. It is noticeable that in the case of S. dudricipeda and do. ab.
zatima (supr. p. 197), the reciprocal crosses do not afford evidence of male
prepotency. ‘The numbers, however, are small.

which may be com-

1 Compare the result of the cross between /. monacha § and ab.
eremita 9. Here the typical monacha form was retained by twenty females
and only two males.

a ~~

RECENT EXPERIMENTS IN HYBRIDISATION. 17

Dresden produced seventy-five Jdetularia (thirty 3 and
forty-five 2) and ninety doubledayaria (thirty-four ¢ and
fifty-six °). The male parent was doubtless an ordinary
betularia. Two of the examples classed as Jdetularia
were darker than the normal, but otherwise no trans-
itional forms occurred. Standfuss is of opinion that even
these two need not be regarded as owing their darker
coloration to the cross, for it is well known that A. detu-
laria, like P. monacha, is undergoing a gradually increasing
melanism, which is probably protective, in many parts of
its area of distribution. The extreme aberration doudble-
dayaria, which thirty years ago was known only from
Great Britain, has now appeared in Westphalia, the Rhine
Provinces, Hanover, Gotha, and lastly in Dresden and
Silesia. In several of these places it is becoming more and
more common, and in at least some of them it is found side
by side with the darkening forms of A. detularia, which,
though of different nature and origin from the sport doudle-
dayaria, are no doubt being preserved and brought up to
its level (in aspect) under the influence of natural selection.?

10. Loarmia repandata L. & and do. ab. conversaria
Hb. ¢. A large brood raised from the eggs of a pair of
normal 4. xefandata contained three males and one female
of the aberration conversarza. This female, which was paired
with a wild male 4. repandata, produced twenty-eight ve-
pandata (of which ten were males and eighteen females) and
six conversarta (four being males and two females). The
majority of the larve died during the winter. Here again
intermediate forms were entirely absent.”

From the above experiments in the pairing of normal
forms with aberrations and local races, performed or recorded
by Standfuss, he arrives at the following conclusions :—

‘ Compare the facts with regard to Papilio Sarpedon given by Jordan,
(“On Mechanical Selection,” ov. Zool., 1896, p. 431) in illustration of the
position that ‘‘ abnormal varieties show distinctly the directions in which a
species is able to develop”. Compare also the result of an experiment
cited by Tutt, “ Melanism,” 1891, p. 15.

2 Compare the result of an observation by Mr. South, who raised

hybrids from the same two forms in 1883 (Proc. Ent. Soc. Lond., 1887,
p. xliv.).

18 SCIENCE PROGRESS.

1. When the normal form of a species (Grundart) is crossed
with a gradually formed local race of the same species, the re-
sult is a series of intermediate forms.

' 2. When the normal form is crossed with a sporadic aber-
ration, the result in many cases is that the issue divides itself
sharply between the normal form and the sport, intermediate
forms being absent.

Hence, according to Standfuss, the process of species-
formation must be gradual; for when two distinct species
are crossed, the issue does not split up into the two parental
forms as in the case when one parent is a suddenly formed
aberration. On the contrary, the behaviour of the issue of
two distinct species is very similar in kind to that of a
species crossed with a local race or variety which is being
gradually established by the accumulation of slight changes.
It would seem therefore that although an aberration or sport
may be perpetuated by inheritance, it can never acquire
distinct specific rank. No doubt however it may, if se-
lected, eventually replace the original form of the species.

To the foregoing considerations it may here be added
that these sporadic colour-aberrations seem to have many
points of resemblance with the colour-varieties in domestic
animals, such as the ‘‘lemon-and-white” and ‘‘tricolour” of
the Basset hounds referred to above (p. 199, note), or the
well-known tortoiseshell, tabby and black of cats. The
fact that these domestic varieties exist side by side in the
same race and even in the same litter, and that true inter-
mediates are rare or absent, seems to suggest that they
originally appeared as sports, and that their perpetuation
has been ensured or favoured by artificial selection, just as,
if Standfuss is right, the dark aberrations of P. szonacha and
A. betularia are being perpetuated and multiplied by natural
selection. Finally, the well-known case of the ‘‘otter sheep”
(Darwin, Varzation of Animals and Plants under Domestt-
cation, 1868, vol. i., p. 100), where also intermediates are
said to be absent, and other instances on record, show that
the phenomenon of sharp division between types in the off-
spring is not confined to crosses between colour-varieties of
the same race, but occurs in other kinds of aberration as

well.

[From the PROCEEDINGS OF THE AMERICAN ASSOCIATION FOR THE
ADVANCEMENT OF SCIENCE, VOL. XLVI, 1897.]

MODIFICATION AND VARIATION AND THE LIMITS OF ORGANIC SELECTION.
By Prof. Henry F. Osporn, Columbia University, New York and
Prof. Epwarp B. PouLron, Oxford University, England.

es

(Abstract of Professor OSBORN’s paper.)

ORGANIC selection is the term proposed by Prof. Mark Baldwin and
adopted by Prof. C. Lloyd Morgan and the author, for a process in nature
which is believed to be atrue cause of definite or determinate evolution in
certain stuctures. The hypothesis 1s briefly as follows: that ontogenetic
adaptation is a very profound character. It enables animals and plants to
survive very critical changes in their environment. Thus all the individ-
uals of a race are similarly modifiid over such long periods of time that
very gradually congenital or phylogenetic variations which happen to
coincide with the ontogenetic adaptive variations, are selected. ‘Thus
there would result an apparent, but not real transmission of acquired
characters.

This hypothesis, if it has no limitations, brings about a very unexpected
harmony between the Lamarckian and Darwinian aspects of evolution,
by mutual concessions upon the part of the adherents of both theories.
While it abandons the transmission of acquired characters, it pliuces indi-
vidual adaptation first, and fortuitous variation second, as Lamarckians
have always contended, instead of placing survival conditioned by fortu-
itous variations first and foremost, as selectionists have contended.

This hypothesis has been endorsed by Alfred Wallace It appears to
the author, however, that it is subject to limitations and exceptions which
go far to nullify its universal application. This is especially seen in the
fact that the law of Determinate Variation is observed to operate with
equal force in certain structures, such as the teeth, which are not im-
proved by individual use or exercise, as strongly as in structures which are
so improved. <A very large class of determinate variations in other
stationary characters, such as the inner parts of the skull, also remain
unexplained. ‘The author’s studies of teeth in a great many phyla of
Mammalia in past time have convinced him that there are fundamental
predispositions to vary in certain directions; that the evolution of the
teeth is marked out beforehand by hereditary influences which extend
back hundreds of thousands of years. These predispositions are aroused
under certain exciting causes and the progress of teeth development takes
a certain form converting iuto actuality what has hitherto been potential-
ity.

This paper was discussed by Dr. Theodore Gill and Prof. Edward D.
Campbell.

(Abstract of Professor POULTON’s paper. )

Ir must be admitted that the adaptation of the individual to its en-
vironment during its own lifetime possesses all the significance attributed
to it by Professor Osborn, Professor Baldwin, and Professor Lloyd Mor-

(239)

240 SECTION F.

gan. These authorities justly claim that the power of the individual to
play a certain part in the struggle for life may constantly give a definite
trend and direction to evolution, and that although the results of a purely
individual response to external forces are not hereditary, yet indirectly
they may result in the permanent addition of corresponding powers to
the species;— inasmuch as they may render possible the operation of
natural selection in perpetuating and increasing those inherent hereditary
variations which go further in the same direction than the powers which
are confined to the individual.

° . . . . . . . . . . .

Professor Osborn’s metaphor in opening this discussion puts the matter
quite clearly and will be at once accepted by all Darwinians. If the hu-
man species were led by fear of enemies or want of food to adopt an
arboreal life. all the powers of purely individual adaptation would be at
once employed in this direction and would produce considerable individ-
ual effects. In fact the adoption of such a mode of life would at first
depend on the existence of such powers. In this way natural selection
would be compelled to act along a certain path, and would be given time
in which to produce hereditary changes in the direction of fitness for
arboreal life. These changes would probably at first be chiefly functional,
as Mr. Cunningham has argued (in the Preface to his Translation of
Eimer). On these principles we can understand the arboreal kangaroo
(Dendrolagus) found in certain islands of the Malay Archipelago, which
is apparently but slightly altered from the terrestrial forms found in Aus-
tralia. Professor Osborn has alluded to the arboreal habits said to have
been recently acquired by Australian rabbits; these and the similar modi-
fication in habits of West Indian rats, are further examples of individual
adaptive modification which may well become the starting point (in the
sense applied above) of specific variation led by natural selection in the
definite direction of more and more complete adjustment to the necessi-
ties of arboreal life. Although this conclusion seems to me to be clear
and sound, and the principles involved a substantial gain in the attempt
to understand the motive forces by which the great process of organic
evolution has been brought about, I cannot admit that the importance
of natural selection is in any way diminished. I do not believe that these
important principles form any real compromise between the Lamarckian
and Darwinian positions, in the sense of an equal surrender on either side,
and the adoption of an intermediate position. The surrender of the
Lamarckian position seems to me complete, while the considerations now
advanced only confer added significance and strength to the Darwinian
standpoint.

I propose to devote the remainder of the time at my disposal in support
of the conclusion that the power of individual adaptation possessed by
the organism forms one of the highest achievements of natural selection
and cannot in any true sense be considered as its substitute. Professor
Baldwin and Prof. Lloyd Morgan thoroughly agree with this conclusion

ZOOLOQ@Y. 241

and have enforced it in their writings on the subject of organic selection.
The contention here urged is that natural selection works upon the highest
organisms in such a way that they have become modifiable, and that this
power of purely individual adaptability in fact acts as the nurse by whose
help the species, as the above named authorities maintain, can live through
times in which the needed inherent variations are not forthcoming, but
in part acts also as a substitute, not indeed for natural selection, but for
the ordinary operation by which the latter produces change. In this latter
case natural selection acts so as to produce a plastic adaptable individual
which can meet any of the various forces to which it is likely to be ex-
posed by producing the appropriate modification and this, it is claimed,
is in many instances more valuable than the more perfect, but more rigid,
adjustment of inherent variations to a fixed set of conditions.

A good example of the eminent advantages of adaptability in many
directions, over accurate adjustment in fewer directions, is to be found
in a comparison between the higher parts of the nervous system in insects
and birds. The insect performs its various actions instinctively and per-
fectly from the first; it is almost incapable of education and of modify-
ing its actions as the result of the observation of the effects of some new
danger. It would appear that the introduction of the electric light can
only affect the insects which are most attracted to it, by the gradual oper-
ation of natural selection. In the clothes-moths, which infest our houses,
we may see an example of this; for these insects seem to be compara-
tively indifferent to light. Birds, on the other hand, have the power of
learning from experience, of reasoning from the results of observation.
At first terrified by railway trains they learn that they are not dangerous,
and cease to be alarmed; while the effect of firearms results in their in-
creased wariness.

If this view of individual adaptability as due to natural selection be
not accepted it may be supposed that the individual modifications are due
either to the direct action of the external forces, or to the tendencies of
the organism. But it is impossible to understand how the mechanical oper-
ation of such forces as pressure, friction, stress, etc., continued through
a lifetime, could evoke useful responses, or why the response should just
attain and then be arrested at a level of maximum efficiency. The other
supposition, that organisms are so constituted that they must react under
external stimuli by the production of new, useful characters, or the use-
ful modification of old ones, seems to me to be essentially the same as the
old ‘‘innate tendency toward perfection” as the motive cause of evolu-
tion —a conception which is not much more satisfactory than special crea-
tion itself. The inadequacy of the view is clearly shown when we consider
that the external forces which awake response in an organism generally
belong to its inorganic (physical or chemical) environment, while the use-
fulness of the response has relation to its organic environment (enemies,
prey, etc.). Thus one set of forces supply the stimuli which evoke a re-
sponse to another and very different set of forces. We can therefore

242 SECTION F.

accept neither of the suggestions which have been offered. Useful indi-
vidual modifications are not directly due to the external forces and are
not due to the inherent constitution of the organism.

The only remaining hypothesis is that which I have already mentioned
—theview that whenever organisms react adaptively under external forces
they do so because of special powers conferred on them by natural selec-
tion. This hypothesis will, it seems to me, meet and satisfactorily ex-
plain all the facts of the case— whether employed as a preparation or as
a substitute for hereditary variations accumulated by Natural Selection.

In closing the discussion, in reply to Professor Poulton, Professor
Osborn stated that natural selectionists were wont to embrace every new
process discovered in nature in the old comprehensive power of selection ;
but that he believed that plastic adaptation had not been acquired by
Natural Selection but was a fundamental property of protoplasm.

Cr BaD

IV. Notes on American and other Tingitide, with
Descriptions of two new Genera and fowr Species.
By GEORGE CHARLES CHAMPION, F.Z.S.

[Read December Ist, 1897.]

Prares hii:

THE following notes were made while studying the
numerous Central-American species of this family of
Heteropterous insects (¢/. Biol. Centr.-Am., Rhynchota, 11,
pp. 1-48). Most of the corrections in the synonymy of
various South-American species have already been noted
by me, but as they may be overlooked in a work dealing
with a limited fauna, they are here repeated. Professor
Poulton having kindly lent me the small but valuable
collection of American Tingitide belonging to the Oxford
Museum—mostly from the Antillean island of St. Vincent,
or from the Amazons (Bates),—some remarks on these are
added, with descriptions of one new genus and three
species. Herr A. Handlirsch, also, has sent me for exa-
mination a large number of American Tingitide belonging
to the Vienna Museum: amongst these, there are speci-
mens of an interesting new genus, from Lower California,
allied to Acalypta, Westw., a description of this insect
being included in the present paper. The opportunity is
taken, too, of figuring a number of the types of Stal’s
South-American species, very many of which, including
those of all his numerous American genera, have been
communicated by Dr. Aurivillius of the Stockholm Mu-
seum. The two South-American species of Monanthia
described by Mr. Distant (Ann. Soc. Ent. Belg. 1888,
p. Ixxxii) are also figured.

PHYLLOTINGIS, Walker.

This genus (= Alyattes, Stal, 1865, and Luloba,
Uhler, 1884), based upon a single species from the
Amazons, P. arida, Walk. (= eximia, Hagl., and pallida,

TRANS, ENT. SOC. LOND. 1898.—PART I. (APRIL.)

56 Mr. G. C. Champion’s notes on

Uhler), is incorrectly referred to the Tingitide: it belongs
to the Aradide. The name <Alyattes being preoccupied,
Dr. Bergroth [Verh. zool.-bot. Ges. Wien, xxxvi, p. 59
(1886)] has substituted Phyllocraspedum for it; Walker's
name, published in 1873, must however be retained.* His
species will stand as Phyllotingis eximia, Haglund’s name
having five years’ priority. Two others are now known.

SOLENOSTOMA, Signoret.

Coleopterodes, Philippi, Stett. ent. Zeit. xxv, p. 306
(1864).

This genus includes a single species, S. laliputianum,
Sign. (= Coleopterodes fuscescens, Philippi), from Chili.
There are two specimens of it in the Oxford Museum.

DOLICHOCYSTA, gen. n.

Rostrum extending to the meso-metasternal suture. Rostral groove
uninterrupted, narrow and parallel on the pro- and mesosternum,
closed in front, the buccal laminze long and very prominent. Intercoxal
portion of the metasternum broad and transverse. Antenne slender,
widely separated at the base. Head without spines, the eyes small.
Pronotum tricarinate, with broad, rounded, closely reticulated mem-
branous margins, and a long decurved, anteriorly prominent hood,
the triangular posterior portion largely developed, acute at the tip.
Elytra short, broad oval, a little longer than the abdomen; dis-
coidal area large, occupying more than two-thirds of the length of
the elytra, raised externally, and surrounded by prominent nervures,
the outer (median) nervure subfoliaceous ; subcostal area about as
wide as the costal ; costal area of uniform width from the base to
near the apex, biseriate almost to the tip. Orifice not visible. Legs
very slender.

This genus bears a general resemblance to Acalypta,
Westw. (= Orthostira, Fieb.); but differs from it in the
very prominent buccal laminze, which close the rostral
groove in front, the long, anteriorly prominent, decurved
pronotal hood, &c. It is perhaps best placed near Corythaica,
Stal. The specimens described may perhaps belong to
a brachypterous form; they are from the northern part of
Lower California.

——

* The antenne are incorrectly described by Walker, the type
having a very elongate basal joint

American and other Tingitide. 57

I. Dolichocysta venusta, sp. n. (Pl. IL, figs. 1; 1a, profile.)

Broad oval, short, dull ; fuscous, the pronotum with the areolz at
the sides in front and behind, the areolz of the hood in part, the tip
of the posterior process, and the three carinz, a broad blackish patch
on the median one excepted, more or less whitish and subhyaline;
the elytra greyish, with a spot on the discoidal area before the apex,
a patch on the costal area before the middle, and some spots or
some of the nervures before and beyond it, black, the other portions
of the costal area whitish and subhyaline with pale nervures, the
nervures on the rest of the elytra blackish or fuscous ; the antennz
testaceous, with the outer half of the apical joint black ; the legs
testaceous, the femora black or piceous in the middle. Antenne
with joints 1 and 2 stouter than the others, 2 much shorter than 1,
3 very slender and elongate, 4 lanceolate and pilose, about as long
as land 2 united. Pronotum closely punctured on the disc ; the
membranous margins with 3—4 rows of small areola ; the median
carina strongly, the two outer carine feebly, foliaceous, the
latter sinuous and connected anteriorly with a layer of mem-
brane which fills the space outside the base of the hood ; hood
extending beyond the head, narrowed and strongly curved down-
wards in front, rather widely reticulated. Elytra a little wider than
the pronotum ; discoidal, subcostal, and sutural areas (the margin
of the latter excepted) closely reticulated ; costal area slightly re-
curved, with larger areole, the two rows diminishing to one at the
apex.

Length 24—23, breadth 13—1 millim.

Hab.—Lower CALIFORNIA, Guadalupe (Bilimek, in
mus. Vind. Cexs.).

Three examples, two of which are from Guadalupe, the
other without definite locality. The insect is not included
in Prof. Uhler’s list of Hemiptera Heteroptera of Lower
California [Proc. Calif. Acad. Sci. (2) iv, pp. 223-295
(1894) ].

CoRYTHUCHA, Stal.

Numerous closely allied American species belong to
this genus. -Monanthia lucida, Walk. (= C. fuscigera, Stal),
and Tingis spinosa, A. Dugés, both from Mexico, also
appertain here. Stal’s type of C. fuscomaculata, from
Brazil, is figured on Plate II, fig. 2. This last-mentioned
insect is extremely like C. fuscigera, Stal, but it appears to
have the apical margin of the elytra less rounded, the
inner portion being oblique.

58 Mr. G. C. Champion’s notes on

TINGIS, Fabr.

Stal (Enum. Hemipt. i, p. 123) uses the name
Stephanitis for the European Tingis pyri, Fabr., and
T. oberti, Kol. and a very dissimilar South- American
form, S. mitrata, Stal, but this course has not been
followed by Lethierry and Severin. The American
species ought to be separated, and the name Stephanitis
could be retained for it. Stal’s type is figured on Plate II,
figs, 3, 3a.

It may be here noted that the insect figured in Cuvier’s
Regne Animal, Ins., Atlas, 1, pl. 91, figs. 5, 5a, 6, under
the name Tingis cristata, Panz., is evidently referable to
T. pyri, Faby. It bas nothing to do with Panzer’s species.

LEPTOBYRSA, Stal.

The type of this genus is Z. stew, Stal, from Rio
Janeiro. It is figured on Plate II, fig. 4. Two others
have been described by Berg from Buenos Ayres and ae
by myself from Central America.

GARGAPHIA, Stal.

This genus seems to be well characterised by the pro-
minent transverse sinuous ridge between the meso- and
metasternum, the cordate intercoxal portion of the meta-
sternum being surrounded at the sides and in the front by
a continuous membranous ridge. The species are all
American.* Stil’s type of G. trichoptera, from Bogota,
Colombia, is figured on Plate II, fig. 5

CoryTHAICA, Stal.
Typonotus, Uhler, Proc. Zool. Soc. Lond. 18938, p. 716.

The characters of this genus were taken by Stal and
Uhler from the same species, C. monacha, Stal (= Tingis
cyathicollts, Costa, and Typonotus planaris, Uhler), the
type of C. monacha being trom Rio Janeiro and that ot
T. planaris from the Island of St. Vincent. Numerous
specimens of the species are contained in the Oxford
Museum. Stal’s type has been communicated by Dr.

* Tingis formosa, Géldi, from Para, probably belongs here or to
Leptostyla, Stal.

American and other Tingitide. 59

Aurivillius. The insect has been beautifully figured by
A. Costa. The second known species of the genus, C.
carinata, Uhler, is from the Island of St. Vincent, and it
has also been found in Guatemala by myself.

PACHYCYSTA, gen. n.

Rostrum reaching the first ventral suture. Rostral groove sub-
parallel, uninterrupted, closed in front, the buccal lamin not promi-
nent, the meso- and metasternal lamin slightly converging behind.
Antennz pilose, distant at the base, with moderately long, slender
joints, 1 and 2 stouter than 3, 1 a little longer than 2, (4 broken off)
Head with five short decumbent spines, very short, obtuse antenni-
ferous tubercles, and an oblong, smooth, convex prominence in front.
Pronotum with a very large, oval, anteriorly truncate hood, three
foliaceous carinee, and very broad, shell-like, incurved membranous
margins. Elytra oval, rounded at the tip; discoidal and subcostal
areas closely, the costal and sutural areas more widely, reticulated,
the costal area very broad, with three or fours rows of hyaline
areole, the discoidal area extending to the middle and surrounded
by prominent nervures. Orifice visible. Nervures of the pronotal
processes and elytra stout, thickly pilose.

The single species from which the above characters are
taken is perhaps nearest allied to the monotypic genus
Megalocysta, Champ. ; but differs from it in having the pro-
notum strongly tricarinate, with the membranous margins
greatly extended, vertical, and shell-like, and the reticula-
tion of the elytra and of the membranous portions of the
pronotum very much closer, &c. It cannot be included in
Leptostyla or any of the other genera characterized by Stal.

Pachycysta diaphana, sp. n. (Pl. II, figs. 6, 6a.)

Body ferruginous, the integument testaceons, the areole of the
elytra and pronotal processes more or less hyaline, the nervures and
the costal margin thickly clothed with very short pallid hairs ;
the legs and antenne ferruginous. Pronotum with the three carinz
strongly foliaceous, each with a row of transverse hyaline areol,
the median carina angularly raised anteriorly, the outer carinz
curved inwards and partly covered by the hood; the hood extending -
to about the middle, rather closely zveticulated ; the membranous
shell-like margins rounded externally, with numerous small areolee ;
the triangular posterior portion large, and, like the disc, closely

60 Mr. G. C. Champion’s notes on

reticulated. Elytra with the subcostal area biseriate ; the broad
costal area with three to four rows of rather small areole; the
sutural area unequally reticulated. Wings extending to a little
beyond the abdomen.

Length 3%, breadth nearly 2 millim. (@.)

Hab. AMAZONS (Bates, in Mus. Oxon.).
One example.

LEPTOSTYLA, Stal.

Four American species were referred to this genus by
Stal; one other has since been added by Berg, and seven-
teen by myself; Monanthia linetfera, Walk., from Brazil,
also belongs to it.* It is one of the most characteristic
genera of the family in Central America. Stdal’s type of
L. furcata, from Rio Janeiro, is figured on Plate I,

figs. 7, Ta.
LEPTOPHARSA, Stal.

This genus will probably prove to be inseparable from
Leptostyla. Two species were known to Stal and a third
has been added by myself, all from Tropical America.
Stal’s type of LZ. elegantula, from Colombia, is figured on
Plate Il, fig. 8.

LEPTODICTYA, Stil.

Five Tropical-American species were included by Stal
under Leptodictya, and two others have been added by
myself. Monantiia tabida, H.-8., from Mexico, also belongs
here. Stal’s type of LZ. fuscocincta, from Rio Janeiro, is
figured on Plate II, fig. 9.

ACANTHOCHILA, Stal.

The type of this genus is A. armigera, Stal (= spinuli-
gera, Stal), from Rio Janeiro, this species being now known
to extend northwards to Southern Mexico. Two others have
since been added by Buchanan White and Ubler respec-
tively. A. armigera has been figured by me elsewhere.

* The type of an allied species, M. lanceolata, Walk., from Brazil,
is apparently lost, as it cannot now be found in the British Museum.

American and other Tingitide. 61

LEPTOCYSTA, Stal.

Stal’s single species, LZ. sewnebulosa, from Rio Janeiro, is
figured on Plate II, figs. 10, 10a.

SPHAROCYSTA, Stal.

Of the two Brazilian species included in this genus, one,
S. globifera, Stal, is figured on Plate II, figs. 11, 1la.

AMBLYSTIRA, Stal.

Stal included in this genus a single species, A. pallipes,
from Rio Janeiro, and four others have been added by
myself. The type of A. pallipes is figured on Plate IT, fig. 12.

LEPTOYPHA, Stal.

The type of this genus is Tingis mutica, Say, from Texas.
Two allied forms from Mexico and Guatemala have been
figured and described by me elsewhere.

TIGAVA, Stal.

This genus was based upon a single species, 7’. pracel-
lens, Stal, from Rio Janeiro; two others have been added
by me from Central America. Stal’s type is figured on
Plate III, fig. 1.

TELEONEMIA, Costa.

Tingis, subgen. Americia, Stal, Enum. Hemipt. iii,
p: 131.

Lasiacantha, Lethierry et Severin, Cat. Hémipt. Hété-
ropt. ili, p. 18 (part.).

This appears to be the most characteristic genus of
Tingitide in Tropical America, fourteen being known to
me from Central America alone.

The types of Stal’s species of Teleonemia have been com-
municated by Dr. Aurivillius, also those of 7ingis (Americia)
albulatera and 7. (Americia) limbata. T. albilatera, as I have
noted elsewhere, = 7ingis triangularis, Blanch., Stal appar-
ently not having seen Blanchard’s figure. The types of the
following species are figured on Plate IIL: 7. (Amauro-

62 Mr. G. C. Champion’s notes on

sterphus) morio, fig. 2; T. validicornis, fig. 4; T. luctwosa,
fig.5; 7. proliva (?=elevata, Fabr.), fig. 6; 7. belfragii,
fig. 8; also that of 7. (Americia) limbata, fig. 10. A figure
of 7’. aterrima is given, fig. 3, from a specimen from the
Amazons in the Oxford Museum, Stil’s type having the
apical joint of the antenne broken off.

Of T. sacchari, H.-S. and Stal, there are numerous
specimens from the Island of St. Vincent in the Oxford
Museum.

The two following species from the Amazons cannot be
identified with any of the known members of the genus,
and they are therefore treated as new.

Teleonemia longicornis, sp. n. (PI. IU, fig. 7.)

Elongate, narrow, dull ; fusco-ferruginous, the spines on the head,
and the margins, carine and hood of the pronotum testaceous, the
triangular posterior portion of the latter yellowish ; the elytra
fuscous, with the base, the outer and inner margins of the sutural
area, the costal margin, and the nervures of the costal area from a
little beyond the middle to near the apex, testaceous, the areole of
the latter for a similar distance, and also some of those along the
apical margin, and two of the inner ones before the tip, hyaline ;
the antenne ferruginous, the two basal joints and the apex of the
third darker, the fourth joint black; the legs ferruginous. Head
with decumbent spines ; antenne very elongate, about as long as
the elytra, moderately stout, joint 2 slightly shorter than 1, 3 three
times as long as 4, 4 pilose. Pronotum sharply tricarinate, and
with a small, compressed, angularly projecting hood in front, the mar-
ginal carinee moderately prominent, the interspaces coarsely, closely
punctate. Elytra long, constricted beyond the middle, and slightly
dilated at the apex, the apices bluntly rounded, the median
nervure prominent ; discoidal area with rather deeply impressed
areola: ; subcostal area biseriate ; costal area with a single row
of transverse areole from about the middle onwards, becoming
vertical and cariniform towards the base. Rostrum reaching the
meso-metasternal suture.

Length 5, breadth 15 millim. (¢.)

Hab. AMAZONS (Bates, in Mus. Oxon.).

One example. This insect resembles Costa’s figure of
T. funerea, but has shorter, less parallel, and differently
coloured elytra, the small pronotal hood more prominent in
front, &c. Amongst the numerous species described by

Amerwan and other Tingitide. 63

Stal (the types of all of which are before me) it is per-
haps nearest allied to 7. serupulosa, from which it may be
separated by the very long and less distinctly pilose
antenng, the longer elytra, with glabrous discoidal area
and the hyaline areolz of the costal area not extending to
the base, &c.

Teleonemia brevipennis, sp.n. (Pl. III, fig. 9.)

Moderately elongate, narrow, dull ; blackish-fuscous, the spines on
the head and the front of the pronotum obscure testaceous, the tri-
angular posterior portion of the pronotum, and the hood in part,
yellowish ; the elytra fuscous, with the costal margin and the
nervures of the costal area for some distance beyond the middle
testaceous, the areole of the latter to near the apex hyaline. Head
with decumbent spines; antenne long and moderately stout, joint
2 considerably shorter than, and not so stout as, 1, 3 rather more
than twice the length of 4, 3 and 4 clothed with very short hairs,
Pronotum sharply tricarinate, and with a small, compressed, sub-
angularly projecting hood in front, the marginal carinee prominent,
the interspaces coarsely, closely punctate. Elytra (when closed)
suboval, comparatively short, rounded at the apex ; discoidal area
with deeply impressed areole, the nervure limiting it externally
rounded and prominent; subcostal area biseriate ; costal area rapidly
widening beyond the middle, very narrow towards the base, with a
single row of areolz, increasing to two in the widest part, the areole
small to about the middle and then becoming larger and transverse.
Rostrum reaching the meso-metasternal suture.

Length 32, breadth 14 millim. (9.)

Hab. AMAZONS (Bates, in Mus. Oxon.).

This species differs from all the Teleonemix known to
me in the relatively short, somewhat oval elytra. The
costal area is abruptly widened beyond the middle, and at
the widest part there are two additional hyaline areole,
there being thus two rows at this place. The wings
extend beyond the abdomen. The insect approaches
the Tingis (Americia) limbata, of Stal, which would be
better placed in Teleonemia.

EURYPHARSA, Stal.

The type of this genus is Tingis nobilis, Guér., from,
Tropical South America, and 7’. circwmdata, Blanch.,

64 Champion: American and other Tingitide.

belongs to the same species. There are two specimens
of it in the Oxford Museum: one labelled “5S. Amer.,
@Orbigny, ex Mus. Jard. Plantes,” this being, no doubt,
one of Blanchard’s types, which were from Chiquitos,
Bolivia; and the other from the Amazons (Bates). A
second species has been described by me from the State of
Panama. Both have been figured.

MonantTHtA, Lep.

The four ‘Tropical-American representatives of this
genus known to me—J/. monotropidia, Stal, M parmata,
Dist., and JL loricata, Dist., and a new species described
by me elsewhere—are closely allied.

Figures of Mr. Distant’s species are here given on
Plate IIT: fig. 11 being taken from JZ loricata, and fig. 12
from J. parmata.

EXPLANATION OF PLATES II anp III.

[See Explanation facing the PuaTEs.]

EXPLANATION OF PLATE II.

[llustrating Mr. G. C. Champion’s “ Notes on American and other
Tingitide.”

Fie. 1, la Dolichocysta venusta, gen. et sp. n.
2 Corythucha fuscomaculata, Stal.
3, 38a Stephanitis mitrata, Stal.
4 Leptobyrsa steini, Stal.
5 Gargaphia trichoptera, Stal.
6, 6a Pachycysta diaphana, gen. et sp. n.
7, Ta Leptostyla furcata, Stal.
8 Leptopharsa elegantula, Stal.
9 Leptodictya fuscocincta, Stal.
10, 10a Leptocysta sexnebulosa, Stal.
11, lla Spherocysta globifera, Stal.

12 Amblystira pallipes, Stal.

Trans Ent Soc. Lond 1898 Ft. Lf.

EXPLANATION OF PLATE III.

Illustrating Mr. G. C. Champion’s “ Notes on American and other
Tingitidee.”

Fie. 1. Tigava pulchella, Stil.

2. Teleonemia (Amaurosterphus) morio, Stal.

3. x aterrima, Stal.

4, * validicornis, Stal.

By, s luctuosa, Stal.

6. 1 proliza, Stal.

fe 5) longicornis, sp. n.

8. + belfragii, Stal.

9. 5 brevipennis, sp. N.

10. - (Americia) limbata, Stal.

11. Monanthia loricata, Dist.

12. 45 parmata, Dist.

Trans Ent Soc. Lond l8IS FLW

ifs

From the ANNALS AND MaGazine oF Natura Hisrory,
Ser. 7, Vol. i., May 1898.

Descriptions of some new Species of Syntomide, chiefly in
the Oxford Museum. By Herserr Druce, F.L.S. &e.

Pseudopompilia mimica, sp. n.

Female.—Primaries and secondaries brownish black: pri-
maries shot with greenish blue beyond the cell and along the
inner margin; a tuft of greenish-white scales at the base,
the fringe black: secondaries with a small semihyaline spot
at the base. The underside of both wings very similar to the
upperside; the primaries with a reddish-brown streak at the
apex and along the inner margin. The head, collar, tegule,
thorax, abdomen, and legs black; the legs with a yellowish-
white streak on the upperside; the antenne black, with the
shaft sordid white from the base to the middle.

Expanse 2 inches.

Hab. Amazons (in the Hope Collection, Mus. Oxford).

This very remarkable species closely resembles Myrmecopsis
vespa, Herr.-Schiiffer, also from the Amazon region.

Calonotos tripunctata, sp. n.

Male.—Primaries and secondaries black ; a small green
spot at the base and a green streak partly along the costal
margin; an elongated white spot below the cell about the
middle of the wing, and a rather large round white spot at
the end of the cell nearer the apex: secondaries with a white
spot on the middle of the costal margin. Underside very
similar to the upperside ; the costal margin of the secondaries
streaked with green at the base. The head, antenna, tegule,
thorax, abdomen, and legs black ; the abdomen striped from
the base to the anus with three wide, greenish-white lines,
one down the middle, and one on each side.

Expanse 1? inch.

Hab. Trinidad (Lowsley, in Mus. Druce).

Calonotos chrysets, sp. n.

Female.— Primaries above dark golden green; underside
much paler, with the veins black: secondaries black, with a
few green scales in the cell ; underside pale green, with black
veins. Antenne and front of the head black, with two small
white spots between the eyes; palpi black. Collar golden
green, edged with black and three small white dots, one in

402 Mr. H. Druce on some

the middle and one on each side; tegule black, golden green
at the base; thorax golden green; abdomen black, with
three white spots close to the base and three rows of golden-
green spots extending from the base to the anal angle; legs
black ; the underside of the abdomen with a row of white
spots.

Expanse 2 inches.

Hab. Bolivia (in the Hope Collection, Mus. Oxford).

This species somewhat resembles Hupyra bacchaus, Schaus.

Homeocera lophocera, sp. n.

Male.—Primaries and secondaries hyaline: primaries, the
base, costal, outer, and inner margins broadly black ; a rather
wide black band crosses the wing at the end of the cell,
extending from the costal margin to the anal angle; two
small blue dots at the base of the wing: secondaries broadly
edged with black from the apex to the anal angle; the veins
of both wings black. The head, antenne, tegule, thorax,
part of the abdomen, and legs black ; a large white spot on
each side of the thorax and one at the base of each leg; the
last three segments of the abdomen and the anal tuft orange-
red.

Expanse 2} inches.

Hab. Brazil (in the Hope Collection, Mus. Oxford).

Homeosoma stictosoma, sp. n.

Female.—Primaries and secondaries yellowish hyaline, the
veins all black ; primaries, the base black, the apex broadly
black, the outer and inner margins black : secondaries edged
with black from the apex to the anal angle, where it is
broadest. The head, antenne, thorax, abdomen, and legs
black ; a white spot on each side of the head, a metallic-blue
spot on both the tegule; a white spot at the base of the
thorax ; two white and two blue dots on the first segment,
and a central row of four white spots down the abdomen, also
a row of white spots on each side of the abdomen.

Expanse 2 inches.

Hab. Colombia (Chesterton, in the Hope Collection, Mus.
Oxford) .

Trichura mathina, sp. n.

Male.—Primaries and secondaries yellowish hyaline ; the
outer and part of the inner margin edged with black; the
veins yellow: secondaries edged with black from the apex to

new Species of Syntomide. - 403

the anal angle ; the end of the cell black. The head, collar,
tegule, and thorax black, the collar and tegule edged with
yellow ; the base of the thorax and the first and third
segments of the abdomen yellow; the abdomen black ;
antenne yellow-black near the base; the legs brownish
yellow.

Eixpanse 1? inch.

Hab. Amazons, Pard (Bates, Mus. Druce).

This species is allied to T. aurifera, Butler.

Sarosa xanthobasis, sp. n.

Female.—Primaries and secondaries hyaline; the veins,
costal margin, outer and inner margins edged with black, the
apex broadly black: secondaries edged with black. The head,
antenne, and palpi black, the front of the head and the collar
metallic blue; the thorax, tegule, abdomen, and hind legs
bright chrome-yellow, the first and second pair of legs
brownish black ; the anal segments of the abdomen bluish
black.

Expanse 2,3, inches.

Hab. Ecuador (in the Hope Collection, Mus. Oxford).

Mallodeta sanguipuncta, sp. n.

Male.—Primaries and secondaries hyaline; the veins
black ; the base, apex, outer and inner margins broadly
black, a wide black band at the end of the cell: the secondaries
edged with black from the apex to the base. Antenne black,
tipped with white on the upperside; the head, collar, tegule,
thorax, abdomen, and legs black ; two small spots at the
back of the head, two at the base of the abdomen, and a row
on each side white; the tegule edged with red; two red spots
at the base of the abdomen, and two on the third segment
somewhat indistinct.

Eixpanse 2 inches.

Hab. Paraguay (in the Hope Collection, Mus. Orford).

Mesothen tgnea, sp. n.

Male.—Primaries and secondaries hyaline, the veins all
black ; the apex of the primaries broadly bordered with black ;
the outer and inner margins edged with black : secondaries
with the apex, outer and inner margins edged with black.
The head, antennex, palpi, and legs black ; the collar, tegule,
thorax, and abdomen dark chrome-yellow ; the tegule spotted
with black and a central black spot on each segment of the
abdomen, the first three being the largest.

Expanse 1,7, inch.

Hab. Merida (in the Hope Collection, Mus. Oxford).

404 Mr. H. Druce on some

Phenicoprocta metachrysea, sp. n.

Male.—Primaries and secondaries hyaline, the veins, costal
margin, apex, outer and inner margins of both wings black.
The head, antenne, palpi, thorax, abdomen, and legs black ;
front of the head metallic blue, the collar and tegule edged
with blue, a central row of greyish-white spots on the
abdomen, the anal segments and anal tuft yellow.

Expanse 13 inch.

Hab. Cayenne (in the Hope Collection, Mus. Oxford).

Urolosia opalocineta, sp. vu.

Male.—Primaries and secondaries hyaline ; the veins all
black ; the costal, outer, and inner margins narrowly edged
with black, widest at the anal angle: secondaries broadly
bordered with black from the apex to the anal angle. The
head, antennz, palpi, collar, tegule, thorax, abdomen, and
legs black; the collar, tegule, and base of the primaries
irrorated with a few metallic-green scales; the second and
third segments of the abdomen opalescent white, more so in
the female than the male; the anal tuft black.

Expanse 1? inch.

Hab. Cayenne (in the Hope Collection, Mus. Oxford).

Hlerea xanthogaster, sp. n.

Male.—Primaries and secondaries brownish hyaline, with
the veins and marginsall black. The head, antenna, thorax,
abdomen, and legs all black; the underside of the abdomen
from the base to the anus bright chrome-yellow.

Iixpanse 1 inch.

flab. Cayenne (in the Hope Collection, Mus. Oxford).

Chrysostola fulvisphex, sp. n.

Male.—Primaries and secondaries hyaline, the veins
brownish black : primaries, the costal and inner margins yel-
lowish brown, the apex and outer margin edged with black :
secondaries edged with black. Antenne black ; the head,
thorax, and abdomen yellowish brown, the anal half of the
abdomen the darkest; the collar and tegule black, edged
with yellow ; the legs yellowish brown.

Expanse 1,'p inch.

Hab, Cayenne (in the Hope Collection, Mus. Oxford).

new Species of Syntomide. 405

Chrysostola sanguiceps, sp. n.

Male.— Primaries and secondaries hyaline : primaries
broadly black at the apex, the outer and inner margins black,
the veins black: secondaries edged with black from the apex
to the anal angle, the veins yellow. The head, thorax, and
basal half of the abdomen chrome-yellow, the anal half of
the abdomen black ; antenne black ; legs chrome-yellow.
Expanse | inch.

Hab. Cayenne (in the Hope Collection, Mus. Oxford).

Paramya picta, sp. n.

Male.—Primaries semihyaline, the base and partly along
the inner margin chrome-yellow, the apex broadly brownish
black; a large black spot at the end of the cell, beyond which
the wing is crossed by a rather indistinct pale yellow band;
a small black streak on the inner margin close to the anal
angle: secondaries chrome-yellow, the apex black. The
head, antennee, last three segments of the abdomen, and legs
black ; the thorax and abdomen chrome-yellow.

Expanse 1 inch.

Hab. Cayenne (tn the Hope Collection, Mus. Oxford).

Cosmosoma orathidia, sp. n.

Male.—Primaries and secondaries hyaline: primaries, the
base, apex, outer and inner margins broadly black; a wide
zigzag black band crosses the wing at the end of the cell
from the costal margin to the anal angle: secondaries, the
costal, outer, and inner margins edged with black. The veins
of both wings black. The head, palpi, antenne, thorax,
abdomen, and legs black; the tegule black, edged with
metallic blue; a large red spot on both sides of the abdomen
at the base and a row of metallic-blue spots extending from
the base to the anus on each side.

Expanse 1,°, inch.

Hab. Nicaragua, Chontales (7. Belt, in the Hope Collection,
Mus. Oxford). .

Histiwa glaucozona, sp. n.

Male.— Primaries brown, slightly shaded with pink in the
cell; two minute blue dots at the end of the cell, beyond which
is a very indistinct brownish-white band, similar in shape to
the yellow band on the primaries of H. amazonica : second-
aries brown, with a large square-shaped red spot beyond the

406 Mr. H. Druce on some

middle. Underside: primaries brown, bright pink in the cell,
two small spots beyond and one large spot at the anal angle
cream-colour ; the spot on the secondaries is quite small and
slightly reddish in colour. The head, antenne, collar, tegule,
thorax, abdomen, and legs brown; a row of metallic-blue
spots on each side of the abdomen.

Expanse 24 inches.

Hab. Amazons (in the Hope Collection, Mus. Oxford).

This species is closely allied to H. amazonica, Butler.

Cyanopepla obscura, sp. n.

Male.—Primaries and secondaries dull brown, a small red
spot and a metallic-green dot close to the base of the primaries
and a small red spot on the costal margin of the secondaries
near the apex; on the underside both wings are shot with
bright metallic blue from the base to beyond the middle. The
head, antenne, thorax, and abdomen black, the collar metallic
blue, each segment of the abdomen edged with metallic-blue
scales ; legs black.

Expanse 14 inch.

Hab. Peru (in the Hope Collection, Mus. Oxford).

Neacerea albiventus, sp. n.

Male.—Primaries and secondaries black : primaries with a
small, transverse, white band about the middle, not reaching
either margin: secondaries, the inner margin broadly greyish
white ; the fringe white at the apex of the primaries. Under-
side very similar to the upperside, but with the marking much
whiter. The head, antenne, collar, tegule, and thorax black;
abdomen above dark glossy blue, on the underside white.
Legs black above, white on the underside.

Expanse 1+ inch.

Hab. South Brazil, Minas Garaes (Rogers, in the Hope
Collection, Mus. Oxford).

Neacerea dizona, sp. n.

Male.—Primaries black, crossed by two semihyaline bands,
neither of which extends to the margins of the wing; the first
band before the end of the cell, the second beyond nearer the
apex: secondaries hyaline, broadly bordered with black from
the apex to the anal angle. The head, antenne, palpi,
tegule, thorax, abdomen, and legs black; collar red; the
underside of the thorax and abdomen white.

Expanse 1%, inch.

Hab. Cayenne (in the Hope Collection, Mus. Oxford).

new Species of Syntomide. 407

Loxophlebia postflavia, sp. n.

Male.—Primaries and secondaries hyaline, the veins all
black : primaries, the base and apex broadly black, the outer
and inner margins edged with black: the secondaries edged
with black from the apex to the anal angle. The head,
antenne, thorax, and abdomen black ; the three anal segments
of the abdomen sordid yellow; the legs black.

Expanse 1,5 inch.

Hab. Cayenne (in the Hope Collection, Mus. Oxford).

Lymire strigivenia, sp. n.

Male.—Primaries smoky black, the veins rather darker:
secondaries hyaline dusky black along the costal margin, and
broadly so along the inner margin. The head, antenne,
palpi, collar, tegule, thorax, abdomen, and legs black.

Expanse 1 inch.

Hab. Cayenne (in the Hope Ovllection, Mus. Oxford).

Psilopleura polia, sp. n.

Male.—Primaries and secondaries whitish hyaline: pri-
maries, the base of the cell, a spot at the end of the cell, the
apex, anal angle, and inner margin shaded with dark and
light brown, darkest along the inner margin: secondaries
edged with brown. The head and thorax pale brown; an-
tenne pale brown, tipped with white; tegule dark brown,
with a pale brown streak down the middle; abdomen black
above, with the base and sides yellow; legs brown.

Expanse 13 inch.

Hab. Espiritu Santo (tn the Hope Collection, Mus. Oxford).

Teucer albapese, sp. n.

Male.—Primaries and secondaries hyaline, the veins all
black ; a large spot at the end of the cell black; the apex,
outer and inner margins broadly black, the fringe at the apex
white: secondaries, the costal, outer margin, and anal angle
broadly black. The head, antenne, thorax, abdomen, and
legs black ; the front of the head and the underside of the
thorax and abdomen white.

Expanse 1 inch. :

Hab. Cayenne (in the Hope Collection, Mus. Oxford).

This species is closely allied to Anaphlebia caudatula,
Felder.

408
Holophea coerulea, sp. n.
Female.—Primaries and secondaries uniformly dark blink
black. The head, antenna, thorax, and legs black; abdomen
dark glossy blue-black ; the collar and the edges of ‘the tegulee
bright red.
Expanse 1 inch.

Hab, Ecuador (in the Hope Collection, Mus. Ozford).

Atyphopsis roseiceps, sp. 0.

Male.—Primaries and secondaries semihyaline greyish
white, the veins all dark brown, the apex dark brown. The
head, antenne, collar, tegule, thorax, abdomen, and legs
pale greyish brown; the top of the head and the fourth and
fifth segments of the abdomen pale red; the anus black.

Expanse 1+ inch.

Hab. §.E. Brazil, Rio Janeiro (in the Hope Collection,
Mus. Oxford).

The specimen is in very poor condition and much faded.

ON A

COLLE@GEEON

OF

INSECTS AND ARACHNIDS

MADE BY

MR. E.N. BENNETT IN SOCOTRA,

WITH

DESCRIPTIONS OF NEW SPECIES.

BY
F. A. DIXEY, M.A., M.D., MALCOLM BURR, F.Z.S

AND THE Rev. O. PICK ARD- CAMBRIDGE, M. Ye
F.R.S., C.M.Z.S

_From the PROCEEDINGS OF THE ZOOLUGICAL Society oF LONDON,
May 3, 1898.]

372 MESSRS, DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [ May 3,

On a Coilection of Insects and Arachnids made by
Mr. E. N. Bennett in Socotra, with Descriptions of
new Species. By F. A. Drxey, M.A., M.D., Matcotm
Burr, F.Z.S., and the Rev. O. PickaArp-CAMBRIDGE,
MEAG, BReS., C.MeZis:

(Plates XXX. & XX XI.)

CONTENTS.

I. Lepidoptera. By F. A. Dixny, p. 372.
II. Orthoptera. By Matcoum Burr, p. 384.
III. Insects of other Orders. By several Contributors, p. 386.
TV. Arachnida, By O. Pickarp-CamBriDGe, p. 387.

I. LEPIDOPTERA, with Remarks on Local and_ Seasonal
Forms in the Genus Byblia Hiibn. By F. A. Drxny, M.A.,
M.D., Fellow of Wadham College, Oxford.

Mr. HK. N. Bennett, a Fellow of Hertford College, Oxford,
reached Socotra on December 17, 1896, in company with the
[2]

1898. ] INSECTS AND ARACHNIDS FROM SOCOTRA. 373

late Mr. Theodore Bent and Mrs. Bent. During their visit they
traversed the island from Ghalansyah in the west to Ras Momi
in the east, and thence, after a long circuit to the south-west,
returned to Tamarida on the north coast. The party left the
island on February 11th, 1897. Interesting personal accounts
of the expedition will be found in the ‘ Nineteenth Century’ for
June 1897, by Mr. Bent; and in the volume of ‘ Longman’s
Magazine’ for 1897, by Mr. Bennett. The whole of their sojourn
in the island came within the period of the N.E. monsoon.
Atmospheric conditions were persistently dry, especially on the
plains; in the mountains there was a heavy dew every morning,
which soon dried in the sun. Very little rain fell at any time,
and the thermometer never sank below 60° F. Exactly 100
specimens of insects and arachnids, which are now in the Hope
Museum, Oxford, were collected by Mr. Bennett.

The Rhopalocera consist of 52 specimens, belonging to 15
species, two of which appear to be new to science. Of these,
15 species, 9 were also taken by Professor Bayley Balfour, eRe oe
during his visit to Socotra between February 11th and March 30th,
1880°. The only one of Professor Balfour’s captures not repre-
sented in the present collection is Charaves balfourt Butl.

DANAIN®.
Limnas cHrysippus Linn. (Nos. 1, 2.)

Two specimens; g and 2. ‘These are paler than the average
of African examples, bearing in this respect a greater resemblance
t specimens from India. The white spots forming the subapical
band are in both, but especially in the male, unusually small and
discrete. Some African specimens show the same character, but
rarely in so pronounced a form*. The Socotran male has most
of the veins in the hind wing, especially the branches of the
median, thickly covered with white scales, which also extend to
narrow adjacent areas of the wing, and form a ring around the
black patch marking the position of the submedian scent-gland.
A trace of the same white colouring of veins and adjacent areas
is also visible in the female. This is a first approximation
to the condition seen in var. alcippoides, Moore, where, however,
the veins themselves often retain their brown colour in the midst
of the whitened area of the hind wing. It is noticeable that both
the Socotran specimens are in fine condition, though the collection
asa whole has suffered much from the attacks of beetle larve.

“Seen only in the hills, flying strongly. Not common.”—£Z. N. B.

' See Proc. Zool. Soc. 1881, pp. 175-180, pl. xviii.

* A pair from Aden in Coll. Brit. Mus. closely resemble the Socotran
examples in this respect, and also in the general ground-colour. A female
specimen from Aden in Coll. Hope, of the same ground-colour, also shows
an approach to the discrete condition of the subapical white spots. Prof.
Balfour’s Socotran specitoen, a female, has like Mr. Bennett’s pair a pale
ground-colour, but the subapical spots are less discrete. It is curious that
specimens of L. chrysippus in Coll. Brit. Mus. from Athens, Turkey, and Syria
are as dark as the ordinary form from Africa.

[3]

374 MESSRS, DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May 3,

ACREIN&,

ACRXA NEOBULE Doub]. (Nos. 10-15.)

Acrea neobule Doubl.; Butler, Proc. Zool. Soc. 1881, p. 177,
pl. xviii. fig. 5.

Six specimens; probably all females, but in one the abdomen is
missing. These resemble A. neobule from the African mainland,
but the powdering of black scales at the apex of the fore wing is
much more distinct, and there is little or no admixture of brown
or reddish scales with the black of the apex, as generally occurs in
A, neobule. In all six examples the ground-colour is deeper in
tone, and the dark border of the hind wing is broader and has a
smoother outline than in average specimens of A. neobule, while
the pale spots in the dark border are on the upperside either not
present or comparatively indistinct. All the black spots on the
hind wing are relatively larger than in normal A. neobule; they
are also more uniform in size; the two spots which occur one on
each side of the discoidal vein, usually very small in A. neobule,
are here less different in bulk from the other dark spots of the
wing. The abdomen in these specimens is black with spots of
the pale ground-colour, as in A. horta Linn. A female A. neobule,
brought from Socotra by Prof. Bayley Balfour, has a perceptible
powdering of reddish scales at the apex of the fore wing, but in
other respects resembles Mr. Bennett’s specimens. It was noted
and figured in 1881 by Mr. Butler (loc. cit.), who, however,
refrained from giving it a specific name in the absence of further
examples. From the present series it seems probable that the
differences from normal A. neobule are fairly constant, but not
sufficiently so to warrant separation. It is worth remarking that
in Reiche’s figure of A. neobule from Abyssinia’ the border of the
hind wing is comparatively narrow, denticulate, and furmished
with large light-coloured spots, while the apex of the fore wing
appears to be powdered with red. The specimen represented
differs therefore considerably from Mr. Bennett's series.

‘‘Mostly seen in the hills, at an elevation of about 2000 feet.
Not hard to get, the flight being slow and bold.”—#. N. B,

SATYRIN &.

CALYSISME ANYNANA Butl. (Nos. 3-7.)

Mycalesis anynana Butl. Ann. Nat. Hist. (5) ui. (1879), p. 187.

Calysisme socotrana Butl. Proc. Zool. Soc. 1881, p. 175, pl. xviii.
figs 7.

Five specimens; 3 ¢,2 9. Two of the males and one of the
females are much worn. The iris of the large ocellus on the
underside of the fore wing, which is whitish in the male described

‘ Ferr. et Gall., ‘Voy. en Abyss.’ ii. p. 466, pl. 53, fig. 5. See Trimen,
‘South Afr, Butt.’ vol. i, 1887, p. 138.
[4]

1898. | INSECTS AND ARACHNIDS FROM SOCOTRA, 370

by Butler (P. Z. S. 1881, p. 175), is in two of the present males,
including the best preserved of the three, distinctly orange as in
the female. The size of all the smaller ocelli on the under surface
seems to vary in both sexes. ‘Te upper surface of the hind wing
carries in the male the glandular patch and tuft of hairs which are
characteristic of the genus (Moore, Lepid. Ceylon, 1830-81, p. 20).
The under surface of the dorsal border of the fore wing, where it
overlaps the hind wing, is similarly clothed in the male with pale
and glistening scales, forming a pearly patch. Five specimens of
CO. anynana in Coll. Brit. Mus. from the Island of Johanna
(Comoro Group) are apparently “ wet-season ” forms ; but another
specimen from the same locality and one from Zanzibar seem
to be “dry-season” forms and are indistinguishable from Socotran
examples.

“The commonest butterfly in the island, inhabiting plains and
mountains alike. A ground-haunting species, apt to take cover.
Never flying high, and always easy to catch.”—E. N. B.

NYMPHALINE.
ByBuia BOYDI, sp.n. (Nos. 16-22.) (Plate XXX. figs.1¢,29.)

Hypanis cora Feisth.; Butl. Proc. Zool. Soc. 1881, p. 177,
pl. xvii. fig. 4.

Types (¢ and 2) in Hope Museum, Oxford.

Seven specimens; 4 ¢, 3 92. Distinguishable from the
“ dry-season ” form of B. gétzius Herbst and B. anvatara Boisd.
by the following particulars :—(1) The area of fulvous ground-
colour lying between the black submarginal band and the oblique
median black patch on the dise of the fore wing is in B. boydi
divisible into two portions, separated by a pair of black denticula-
tions which almost meet one another along the course of the
first median branch. Of these two portions, the posterior is
conspicuously narrower than the anterior, the narrowing being
caused mainly by the encroachment outwards of the oblique
median patch. The outline of this latter patch in the allied forms
tends rather sharply inwards between the first median branch
and the dorsal border, but in B. boydi it is continued to the
dorsal border at such an angle as to preclude the fulvous area from
expanding again posteriorly, as it does in normal B. gotzius.
(2) A chain of small black spots is more or less visible, crossing the
fulvous median area of the hind-wing upperside. These spots,
which correspond to a series constantly present in B. iithyia
Drury, are only rarely indicated in B. gétzius. The above
characters appear to be constant and distinctive. One or more
of the following features may be found in specimens of B. gétzius
from various localities on the mainland, but they do not occur all
together except in B. boydi, where the combination appears to be
constant :—(1) The black costal bar of the fore wing is continued
across the wing to meet the submarginal black band. (2) The
fulvous submarginal spots of the hind-wing upperside are large,

[5]

376 MESSRS. DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [ May 3,

subconical, and only slightly separated by the black-coloured veins.
(3) All the black markings of the upperside are highly developed,
especially the submarginal band of the hind wing, which encroaches
considerably inwards. In the presence of the chain of small
median dark spots and in the large size of the fulvous submarginal
spots of the hind wing, B. boydi approaches B. ilithyia ; in other
respects it is much nearer B. gétzius. The combination of
characters above given renders the Socotran form easily recog-
nizable among its allies, and seems to justify its separation as
distinct. I have given it the name boydi, after the Principal
of Hertford College, to whom Science in Oxford is under great
obligations.

“« Very common everywhere, hills and plains. Not conspicuously
ground-haunting.”—E. N. B.

Remarks on Geographical and Seasonal Forms in the Genus Bybla
Hiibn.— Byblia gotzius Herbst (=Hypanis acheloia Wallengr. ;
=H. ilithyia var. A, Trimen, S. Afr. Butt. vol. 1. 1887, p. 264) is
probably entitled to distinct specific rank beside B. ilithyia Drury’.
Each form, as pointed out by Trimen (loc. cit. p. 266) and by
Barker (Trans. Ent. Soc. Lond. 1896, p. 415), has its owu range otf
seasonal variation. ‘This is also shown by good series of both the
ilithyia and the gotzius (or acheloia) forms in the British Museum
and in the Hope Collection at Oxford. For a large proportion of
these each collection is indebted to Mr. G. A. K. Marshal], whose
specimens all bear such asnple data with regard to locality, altitude,
and exact time of capture, as to throw much light upon questions
of local and seasonal modification.

The geographical distribution of the two forms is interesting.
The ilithyia form, with some local variation in size and in the
relative proportions of dark markings to fulvous ground-colour, is
found in India, Ceylon, Arabia, and the greater part of Wallace’s
“Kast African” subregion, including the West African coast dis-
tricts lying northwards trom the River Gambia and southwards
from the Congo. It also extends for some distance into the South
African subregion, occurring commonly in the Transvaal and Natal
highlands, and coming, though rarely, down to the sea at Durban.
Its distribution is therefore mainly Indian and “ East African ” in
Wallace’s sense*. The gotziws form, on the other hand, is abseut
from India and from a large portion of “ East Africa.” Itis found
at Sierra Leone, Cape Coast Castle, Lagos, Old Calabar, and the
coast districts of the Gaboon and the Congo; but outside the limits
of Wallace’s West African subregion, 7. e. in Senegal to the north
and Angola to the south, it is replaced by typical ilithyia. Begin-
ning again on the south-east coast, about the easternmost districts

2 The evidence on which Mr. Marshall decides against their specifie distinct-
ness appears to me to require confirmation. See Marshall in Ann. Mag. Nat.
Hist. 1896, xviii. p. 338.

2 Wallace, ‘Geographical Distribution of Animals, 1876, vol. 1. pp. 261,
258 and map, .

[6]

1898. | INSECTS AND ARACHNIDS FROM SOCOTRA, 377

of the Cape Colony', it becomes very common at Durban, and
follows the coast-line northwards, occurring at the mouths of the
Zambesi, in Mozambique, and at Wasin. Though it seems to be
rarely if ever met with in the “South African” interior, it passes
inland up the Zambesi and is found in Matabeleland and at Zomba
on the Shiré ; while the British Museum also contains specimens
from Nyasaland, Lake Mweru, Tanganyika, the Victoria and Albert
Nyanza, Wadelai, the Galla country, Abyssinia, Somaliland, and
Aden. Like B. «ithyia, it shows some amount of local variation
which may perhaps justify the specific separation of certain
geographical forms’.

It appears therefore that, so far as is known, the distribution of
the two species (or varietal groups) is fairly distinct, though their
respective ranges coincide for a small portion of the South African
subregion and to a larger extent in ‘* East Africa,” as at Wadelai,
in Somaliland, and at Aden. It is further evident that while the
distribution of the ilithyia form is continuous from India through-
out the “ East African” subregion, that of the gétzius form is
almost if not quite discontinuous, its area being separated into a
western and an eastern division. In the hight of these facts it is
remarkable that the Socotran form is most closely akin, not to the
ilithyia, but to the gétzius type, nearly resembling in fact West
African specimens of B. gétzius, from which it is separated geo-
graphically by the whole width of Wallace’s East African subregion.
It is further of interest to note that the Madagascar and Comoro
Islands form (B. anvatara Boisd.), though no doubt distinct, is also
a modification, not of B. ilithyia, bat of B. gétzius*®. B, ilithyia,
being found at such distant points of the “ Kast African” subregion
as Senegal, Angola, and Somaliland, as well asin Arabia, India, and
Ceylon, might well have been expected to be the form occurring in
Socotra ; and the fact that it is here replaced by a form of B. got-
zius suggests the possibility that this island, like the South African
subregion, and Madagascar with the Mascarene group, contains
relics of a more ancient African fauna that has been expelled or
excluded from the bulk of the mainland by the great irruption of
forms of life which is believed to have taken place from the north-
east *.

‘ It is implied by Mr. Marshall (/oc. cit, p. 3837) that this Southern race of
B. gotzius (to which he restricts Wallengren’s name ache/oia) occurs also in the
Western districts of South Africa, where, he states, the Cunene River (north
of Damaraland) appears to be its northern boundary.

* Mr. Marshall (oc. cit. pp. 837, 338) recognizes three local races—the
Southern (acheloia Wallgr., of which vulgaris Butl. is the wet-season form),
the Western and Central African (gotzius Herbst), and the North-eastern
(castanea Butl.). Prof. Bayley Balfour’s Socotran examples of B. boydi, noted
by Butler as Hypanis cora Feisth., are rather curiously ranked by Mr. Marshall
under var. acheloia Wallgr.

3 Trimen, South Afr. Butt. vol. i. 1887, p. 267.

4 Wallace, ‘Geographical Distribution,’ 1876, vol. i. p. 218, &e. Prof.
Bayley Balfour (Proc. Roy. Instit. vol. x. 1884, p. 296) discusses the affinities
of Socotran with West and South African plants, and observes that “ Sokotra
indecd is, with Madagascar, to be regarded as the remains of a greatly advanced

[7]

378 MESSRS, DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May 3,

The variations of the underside of the hind wing in both forms,
B, ilithyia and B. gétzius, have been well described by Trimen
(loc. cit. pp. 265, 266). They are undoubtedly seasonal, as
pointed out by Barker (Trans. Ent. Soc. Lond. 1895, p. 415), and
by Marshall in the MS. notes and labels accompanying the series
in the Hope Collection above referred to, as well as in the
Annals & Mag. Nat. Hist. 1896, xviii. p. 333, &e. The deeply
ferruginous hind wing, on which the three creamy bands stand out
conspicuously, belongs in each case to the dry-season form, and
there are several intermediate grades leading up to the dull
ochreous yellow of the wet-season form. In addition to the
points noted by Trimen, it may be remarked that in the wet-
season form of B. gétzius the black submarginal band of the hind
wing is relatively broader, and the proximally adjacent strip of
ochreous ground-colour narrower, than in the wet-season form of
B, ilithyia. In the former, indeed, the band of ground-colour is
often reduced to a mere chain of fulvous dots with dark edging,
forming a proximal border to the dark submarginal band. The
pairs of whitish interpervular spots on the dark band are also
much less regular and conspicuous than in B. ilthyia. In the
dry-season forms the veins crossing the median creamy band are
in B. gotzius often traced out with the deep ferruginous tint of
the ground-colour, which marking has the effect of dividing the
median creamy band into spots; this is not seen in B. alithyra.
In B. gétzius also the dark submarginal band seems never entirely
to disappear, even in extreme dry-season forms, as it may do in
B. ilithyia. It soon, however, loses the whitish internervular
spots, which in the wet-season form are already less distinct than
in B, alithyia.

The Socotran B. boydi resembles most specimens of B. gétzius
from the West African subregion in having the dark costal bar of
the fore wing continued rather heavily across the wing to join the
submarginal band. This is also more or Jess the case with two
females of B. gétzius 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 between the
costal and the submarginal dark bands is often slight or absent. On
the other hand, in the submarginal series of spots of the fulvous
ground-colour on the upperside of the hind wing, the Socotran
form comes nearer to specimens of B. gotzius from Somaliland,
Aden, and the Galla country than to any I have seen from West

African coast-line at a remote period.” Messrs. Sclater and Hartlaub (Proc.
Zool. Soc. 1881, p. 167) point out that Drymaca hesitata, one of Prof. Balfour's
Socotran birds deseribed by them, is most closely allied to a form inhabiting
Madagascar. Col. Godwin-Austen (Proc. Zool. Soc. 1881, p. 252) considers
that the land-molluscan fauna of Socotra affords “strong evidence that the
island was once directly connected with Madagascar to the south”; and adds
that “it is not unreasonable to suppose that in Socotra, the Seychelles,
Madagascar, and Rodriguez we have the remnants of a very ancient more
advanced coast-line on this western side of the Indian Ocean.”

(8)

1898. ] INSECTS AND ARACHNIDS FROM SOCOTRA. 379

Africa. The spots in question are in B. boydi, as in the British
Museum specimens of B. gétzius from the localities last named,
larger, closer together, more conical, and less quadrate than in
individuals from West or South Africa, though in dry-season
forms from Natal and East Central Africa an approach is made to
the Socotran condition, The dark submarginal band of the hind
wing in B. boydi is broader than in most specimens of B. gotzius
from E. and S. Africa, whether ‘‘ wet” or ‘‘ dry” (the narrower
band belonging generally to the “dry” form). It is much
broader than in the specimens of B. gotz’us from Somaliland, Aden,
and the Galla country above referred to, but not, perhaps, much
broader on the average than in the female specimens from
Abyssinia.

The present examples of B. boyddi, like Prof. B. Balfour’s pair,
are all dry-season forms. In at least two of the seven (both
males), as also in both Prof. Balfour’s specimens, the whitish
internervular spots on the dark submarginal band of the hind-wing
underside have disappeared, and in one of these the patches of
ground-colour immediately adjacent to the pale median band are
obsolescent. The wet-season form of B. boydi is still unknown.
To judge by the analogy of B. yétzius, its upper surface must be
still more heavily marked with black than that of the specimens
collected by Prot. Balfour and Mr. Bennett.

PyraMEIs carDUI Linn. (Nos. 8, 2.)

2 9. This species was also observed by Prof. Balfour.

‘Common everywhere. The three most abundant species, in
order of frequency, were (1) Calysisme anynana, (2) Byblia boydi,
(3) Pyrameis cardui.”—E. N. B.

JUNONIA CLELIA Cram. (Nos, 23-28.)

Six specimens; 2 ¢,4 @: two of the latter in a battered
condition. These do not differ in any definite manner from
specimens from the mainland and the Comoro Islands *. They are
“ dry-season”’ forms, the colouring of the hind-wing underside
being fairly uniform and the ocelli obsolescent. This species was
not obtained by Prof. Balfour.

‘Very common in the mountains.’"—Z. N. B.

HyponimMnas Misippus Linn. (Nos. 29-32.)

4 9. These are of the ordinary form, showing no tendency
towards var. alcippoides But]. They have suffered much from the
attacks of larve. Not obtained by Prof. Balfour.

“ Fairly common; commoner than ZL. chrystppus. Chiefly in
the hills. Flight strong.”—£. N. B.

1 In J. epiclelia Boisd., from Madagascar, the size of the creamy-white
markings of the upperside is much reduced, but I doubt whether the other
features mentioned by Trimen (doc. cit. p. 216) are constant points of difference

from J. clelia.
[9]

380 MESSRS. DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May 3,

LYCENIN&.

TARUCUS THEOPHRASTUS Fabr. (No. 33.)

lier) Aan species has a wide range throughout the Indian
and Ethiopian regions and the Mediterranean subregion of the
Palearctic. It does not occur in Prof. Balfour’s collection.

ZizpRA LYSIMON Hiibn. (Nos. 34-87.)

Lycena lysimon, 'Trimen, South Afr. Butt. vol. ii. 1887, p. 45.

Four specimens; apparently 3 ¢ and 1 @. This species also
was not obtained by Prof. Balfour.

“Found cominonly everywhere, both hills and plains, but
chiefly the former. Flight always close to the ground.”—
E. N. B.

A Lycenid collected by Riebeck, who visited Socotra soon after
Prof. Balfour, was not determined’.

PIERIN &.
BELENOIS ANOMALA Butl. (No. 38.)

Synchloe anomala Butl. Proc. Zool. Soc. 1881, p. 178, pl. xviii.
fig. 3.

One @. The specimen is broken, but less worn than the type,
which is also a female. The large black spot at the end of the cell
in the fore wing, subquadrate in the type, is here rather sub-
triangular, with the base directed inwards, and showing on both
surfaces a slight proximal indentation. On the under surface the
outer border of the fore wings is greyish shot with pink, not
semitransparent as in the type.

Mr. Butler (loc. cit.) assigns this form to the genus Synchloe,
but adds that * the possession of a male specimen would satisfac-
torily decide whether or not it is an unusually aberrant Belenois.”
The male is still unknown ; the venation, however, is unmistakably
that of Belenois, as the 1st subcostal branch in the fore wing
is concurrent with the costal (cf. B. mesentina, B. creona, B. gidica,
&e.), while the upper discocellular is straight and forms an open
angle with the lower. There can be little doubt that this interest-
ing species comes nearest to B. abyssinica Lue., the dry-season
form * of B. gidica from the African mainland.

“ Rare ; only met with in the Haghier Range, at an altitude of
about 2300 ft. In the same place another white butterfly of
corresponding size was seen, with circular black spots [perhaps
the male]. Both flew fast.”,—E. NV. B.

TERACOLUS NIVEUS Butl. (Nos. 39-42.)

Teracolus niveus Butl. loc. cit. p. 177, pl. xviii. fig. 1.

Teracolus candidus Butl. loc. cit. p. 179, pl. xviii. fig. 2.

One 3, three 9. The male and one female correspond with
' Taschenherg, Zeitschrift fir Naturwiss. Bd. lvi. 1883, p. 182.

* See Barker, Trans. Ent. Soc. Lond. 1898, p. 419.
[10]

1898. | INSECTS AND ARACHNIDS FROM SOCOTRA. 381

Butler’s 7. niveus; another female is more heavily marked ; the
third female agrees with his 7. candidus. In the male a few
yellow and orange scales form a minute speck proximally, adjacent
to the black dot at the end of the cell on the underside of the hind
wing. A similar yellowish speck occurs in the females, but tends
towards the costal rather than the proximal aspect of the dot.

Dr. Butler now considers his 7’. niveus to be the wet-season
and 1. candidus the dry-season form of the same species. In
reference to the faunistic affinity between Socotra and the
Mascarene group (supra, p. 377), it is of interest to note that
T. aldabrensis Holl., from Aldabra, appears to be the nearest
relative of the Socotran 7’. niveus'.

“The male was taken on Dec. 19th in the sandy coast-plain of
Ghalansyah. It was rescued from the jaws of a lizard.”—Z. N, B.

CaATOPSILIA FLORELLA Fabr. (Nos. 43-46.)
Catopsilia pyrene Swains.; Butl. Proc. Zool. Soc. 1881, p. 178.
Four specimens; 1 ¢,3 2. The male and two of the females
are much worn.
“Only seen in the plain of Tamarida. Flght strong.”’—
Hie NB.
PAPILIONIN A.

PAPILIO BENNETTI, sp.n. (Nos. 47,48.) (Plate XXX. fig. 3.)

Type in Hope Museum, Oxford.

Two specimens, both probably ¢, but the abdomen of one is
imperfect. These resemble P. demoleus Linn., from the African
mainland, but may be distinguished by the following characters :—
(1) On the upper surface all the yellow markings are much reduced
in size, and the second spot from the dorsal border of the yellow
median chain in the fore wing ts more or less Z-shaped, instead
of being irregularly rhombic as in P. demoleus. (2) There is a
broad black area of almost uniform width between the median and
the submarginal chains of yellow spots on the fore wing. The
corresponding area in P. demoleus is comparatively narrow, and
conspicuously denticulated in consequence of the relatively large
size of the median yellow spots. (3) On the under surface the
pale submarginal spots of the hind wing are quadrate, or even
elongated in a direction at right angles to the border of the wing ;
whereas in P. demoleus they tend to be oblong, with the long
diameter parallel to the hind border. The same applies to the
series of rudimentary eye-like marks immediately proximal to the
yellow submarginal row. Another feature which is probably
distinctive is the fact that in the eye-like mark within the cell on
underside of the hind wing the blue edging with its accompanying
buff crescent extends only along the posterior side of the tri-
angular black patch, instead of being continued along two sides,
the posterior and the dorsal, as in P. demoleus. An approach to

' See Butler, Ann. & Mag. Nat. Hist. 1897, xx. p. 464.

Proc. Zoou. Soc.—1898, No. XX VL. 26
[113

382 MESSRS. DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May 3,

this condition may occasionally be seen in the latter species.
Many specimens of P. demoleus from Aden resemble P. bennetti
in the narrowness of the pale median band of the hind wing ;
they differ, however, in the other particulars.

Mr. Bennett’s specimens were taken on the extreme summit of
Jebel Dryet (4900 ft.), settled on a Bedaween’s bright-coloured
cotton wrap or loin-cloth. The species is a strong flyer. It was
not often met with, and never at a less elevation than 3500 ft.
It does not occur in the collection made by Prot. Bayley Balfour.

HESPHRIIDA.
RwHopaLocaMpra sucunpA Butl. (Nos. 49-51.)

Hesperia jucunda, Butl. Proc. Zool. Soc. 1881, p. 179, pl. xviii.
fig. 8.

Three specimens, all ¢. This species, as remarked by Trimen
(South Afr. Butt. vol. iii. p. 373) is near R. keithloa Wallgr. from
the East African mainland. It is also allied to R&. taranis Hew.
(R. anchises Gerst.), which has a wide African distribution and
oceurs at Aden (Butler, loc. cit. p. 179).

“Found in the hills, and also in the coast-plain between
Ghalansyah and Tamarida.”—E£. N. B.

GEGENES NosTRADAMUS Fabr. (No. 52.)

One specimen,a @. This species, which extends throughout
the Mediterranean subregion into the North-western districts of
India, was not obtained by Prof. Balfour. Specimens from Aden
(var. karsana Moore) are more sandy in colour than the Socotran
example.

The Heterocera collected by Mr. Bennett consist of 26 speci-
mens, belonging to 16 species. These have been kindly named by
Sir George F. Hampson. There are no new forms among them ;
Oligostigma incommoda, which was described by Dr. Butler (Proc.
Zool. Soc. 1881, p. 180) from a specimen obtained by Prof.
Balfour, does not occur in Mr. Bennett’s collection. The Socotran
Heterocera, so far as they are known, appear to present a mixture
of African and Oriental species, the former predominating, together
with some widely-distributed types such as D. pulchella, As in
the case of the Rhopalocera, the African element does not seem to
be exclusively East African. It is unfortunate that the Moths
collected by Riebeck’ were never determined.

CossipDZ.
AZYGOPHLEPS INcLUSA Wlk. (No. 53.)

One specimen. Another packed by Mr. Bennett was completely
destroyed by beetle larve.
‘In the hills.,—#. WN. B.

___‘ Taschenberg, Zeitsehr. f. Naturwiss. Bd. lvi. 1883, p. 182.
(12)

1898. ] INSECTS AND ARACHNIDS FROM SOCOTRA, 383

ARCTIIDSE.
DEIOPEIA PULCHELLA Linn. (Nos. 54-57.)

“ Ardahan, &c. Always on the grassy slopes of hills. Lived
in the grass; never flew more than 2 feet from the ground.”—
E. N. B.

Four specimens. This species was also obtained by Prof.
Balfour.
Liryosta vetusta Wlk. (No. 58.)
One specimen.
Noovrvuipdp..

AGRoTIs pivisA Wlk. (Nos. 59, 60.)
Two specimens, one much worn.

Euriexia conpucta Wlk. (No. 61.)
One specimen.

BaNIANA INtTORTA Swinh. (No. 62.)
One specimen.

CEROCALA VERMICULOSA H.-S. (Nos. 63-69.)
Seven specimens.

CALPE EMARGINATA Fabr. (No. 70.)
One specimen.
GEOMETRID &,

HyperyrHra tuctcotor Butl. (No. 71.)
One specimen.

Boarmta acactarta Boisd. (No. 72.)
One specimen.

CRASPEDIA DERASATA Wlk. (No. 73.)
One specimen.

CRASPEDIA LACTARIA Wlk. (No. 74.)
One specimen.

CRASPEDIA PULVEROSARIA Wlk.? (No. 75.)

One specimen.
PYRALIDAS,

NepnHorrerrx sp. (No. 76.)
One specimen, too much worn for recognition.

Merasta sp. (No. 77.)
One specimen.
TORTRICIDSE.
Trras sp. (No. 78.)
One specimen.
26*
[13]

384 MESSRS. DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May 3,

II, ORTHOPTERA.
By Maxucoum Burr, F.Z.S.

The Orthoptera collected in Socotra by Mr. Bennett are few in
number, but not without interest. The new species of Pecilocerus
represents a genus found in Northern Africa and in the western
part of Asia, and there is a new Cricket, of the genus Landreva
Walker, with a similar, but wider, distribution. It is probable
that these two species are peculiar to the island, as they are not
migratory in habits, so far as is known, the Cricket at least being
incapable of flight.

There is the usual percentage of cosmopolitan species, but the
collection is hardly large enough to give a fair idea of the relation
of the island to the neighbouring continents.

A Locustid has been described, Pachysmopoda abbreviata *
(Tasch.), which is not found elsewhere, but was not taken by
Mr. Bennett.

FORFICULARIA.

LABIDURA RIPARIA (Pall.).

One male (No. 79).

A cosmopolitan species; originally apparently an inhabitant of
the Mediterranean subregion, also occurring in Java, Korea, South
America (coll. m.), Burmah, and North America.

BruatTropeBaA.
PHYLLODROMIA sp.

A very fragmentary example (No. 80), which resembles, but is
distinct from, the cosmopolitan Ph. germanica (1..).

ACRIDIODBA.
TRYXALIS NASUTA (L.).

One immature specimen (No. 81).
This species occurs in Southern Europe, throughout Africa,
India, Burmah, and in Australia.

ACROTYLUS LONGIPES (Charp.).

The solitary example (No. 82) is a variety with the wings rosy
at the base, the normal colour being yellow. A blue form has
occurred at Zanzibar (Brunner). An inhabitant of South-eastern
Europe, Asia Minor, Abyssinia, and Zanzibar.

P@CILOCERUS SOKOTRANUS, sp. n. (Plate XXX. fig. 4.)
Statura minore. Caput conicum, pallidum, vertice inter antennas
1 Mecopoda abbreviata, Taschenberg, Zeitschr. f. Naturwiss. Bd. lvi. 1883,

p. 184.
[14]

1898. | INSECTS AND ARACHNIDS FROM SOCOTRA. 385

minus producto, horizontale, supra atro, subtus in frontem per-
currente, profundius sulcato, fronte reclinata, a latere visa vie
sinuata ; antennee breves, nigre, apicem versus palladiores, capite
et pronoto unitis viv longiores ; oculi prommuli. Pronotum
eylindricum, fusco-testaceum, carts nullis instructum, margine
postico rotundato, sulco ty pico poullo pone medium sito ; lobi
defleai laterales marginibus, postico sinuato, tnferiore recto, antico
adscendente, angulis rotundatis. Elytra et ale perfecte explicata,
abdominis apicem via attingentia, illa angusta, fusco-testacea,
umcoloria, densissime reticulata, apice obtuse angulata ; he
elytris breviores, badice, apicem versus pallidiores. Femora
tibiceque antica et intermedia fusca, tarsi pallidiores ; femora
postica extus pallida, media macula magna fusca ornata, mtus
testacea, fusco-reticulata ; genibus supra pallidis, lateribus
atris ; tibie postice sordide testacec, spinis octo albidis, apice
nigris, calcaribus terminalibus parvis binis utroque margine
supra armate. Abdomen fuscum, cylindricum ; valvule
ovipositoris longe, sinuate, pubescentes. @.

Dimensions, 2.

Wrong :"Corp? o< ssc 30 millim.
5 elyirorumras so lnoe 5
oy) PEOUOLIE Sy ee se Mare

The two specimens (Nos. 87, 88) from which the d scription
was drawn are somewhat discoloured by spirit, but that is not a
very important injury. P. sokotranus is considerably smaller than
the other species of the genus, and is probably peculiar to the
island.

Two females (Nos. 83, 84). Upon the second segment of
both specimens there is a curious round pale hard knob, so large
that it has caused a space in the elytra where they cover it when at
rest. I have omitted it from the description as it seems to be
a foreign body, possibly a fungus.

There is also an immature Grasshopper (No. 85), possibly to be
referred to the genus Acridiwn.

GRYLLODBEBA.
LANDREVA, sp. n. ?

A male Cricket (No. 86) of the genus Landreva Walk. seems
to be new, but is not sufficiently good for description. It is small,
testaceous, with truncate elytra and no wings. The tympanum is
only visible on the exterior side of the anterior tibie (subgenus
Ectolandreva, Sauss.); the posterior tibie are armed with five
spines on each margin above, and four terminal spines.

[15]

386 MESSRS. DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May 3,

III. Inysucrs or orHer OrpERS. By several Contributors.

Five species of Odonata, two of Hymenoptera, and one of
Diptera were also captured by Mr. Bennett: the first were kindly
named by Mr. R. McLachlan, F.R.S., the second by Mr. W. F.
Kirby, and the third by Mr. E. E. Austen.

The Odonata were all common species with wide distribution,
and it is of interest that they should in this respect contrast so
sharply with the Lepidoptera Rhopalocera and Araneidea, and to
a less extent with the Orthoptera. The contrast is probably to
be explained by the facilities for distribution which the Odonata
possess in their powers of flight.

Mr. Bennett records that the Odonata are very numerous on the’
plains where streams abound.—E. B. Poutron.

Species of Odonata.

PANTALA FLAVESCENS Fabr. (No. 89.)

Mr. Bennett captured one specimen “ on banks of streams pass-
ing T'amarida.”
Mr. McLachlan describes the species as nearly cosmopolitan.

CROCOTHEMIS ERYTHR#A Brullé. (No. 90.)

One specimen.

The species occurs in South Europe, all over Africa, Asia, &c.
RR. M.).

RHYOTHEMIS SEMIHYALINA Dujardin. (Nos. 91, 92.)

Two specimens. ‘Common near the lagoon at Ghalansyah.”—
E. N. B.

Widely distributed in Africa and occurs in Asia Minor (2. 1/.).

CERIAGRION GLABRUM Burm. (No. 93.)

One specimen.

Nearly all over Africa (#. M.). The British Museum contains
examples from Socotra, Madagascar, Mauritus, as well as from the
mainland of Africa.

Species of Hymenoptera.

BELENOGASTER SAUSSUREI Kirby.

Two specimens (Nos. 94, 95).

This species was described by Mr. W. F. Kirby (P. Z. 8S. 1881,
p. 649) among the insects collected by Prof. Bayley Balfour in
Socotra.

HARPACTOPUS sp. inc.
One specimen (No. 96).
A species allied to H. crudelis, Smith, but larger, and with red-
dish mandibles and tibiz. The specimen is, however, in such bad
[16]

1898. ] INSECTS AND ARACHNIDS FROM SOCOTRA. 387

condition that a description would be useless if not misleading
CHE EO):

Species of Diptera.
SARCOPHAGA, sp. inc.

One specimen (No. 97).
The condition renders it impossible to determine the species
(E. E. A.).

IV. ARACHNIDA.
By the Rev. O. Prckarp-CamBripex, M.A., F.R.S., C.M.Z.S.

ARACHNIDA ARANEIDEA.
Fam. EPELRID&.
Subfam. NEPHILIN”,

Gen. Nrepuina Leach.

NEPHILA BENNETTI, sp.n. (Plate XXXI. fig. 2.)

Female adult ? (No. 98), length 83 lines.

This Spider is of the ordinary characteristic Nephila form, and
at the hinder part of the caput are the two characteristic conical
tuberculiform eminences, conspicuous in a transverse line; the
colour of the caput is dark brown, with two large, somewhat oval,
dull yellowish patches at the hinder extremity, each surrounding
one of the conical eminences. The thorax is dull, pale yellowish,
with black-brown lateral converging stripes, and the whole cephalo-
thorax is thickly covered with light grey pubescence.

Eyes normal.

Legs long, moderately strong; 1, 4, 2, 3, furnished with hairs
and slender spines. Femora yellow, slightly tinged with a smoky
hue at the anterior extremity ; genuz of the first and second pairs
dark brown, those of the third ard fourth pairs yellowish; tibia,
metatarsi, and tarsi deep brown, approaching black. The tibie of
the fourth pair are furnished throughout their length with nume-
rous coarse black hairs forming a brush ; the femora of the first and
second pairs have also a similar but not so strong a brush on their
fore-half, the hairs on these last shorten in length gradually into
the normal hairs of the remaining portion of the joint.

Palpi short, normal; humeral and cubital joints yellowish,
radial and digital black-brown, the latter furnished with long
coarse bristly hairs.

Falces powerful, normal in form, black-brown.

Mawille and labium )black-brown, the latter with a central
longitudinal reddish yellow-brown stripe.

Sternum triangular, slightly longer than broad ; deep brown: with

[17]

388 MESSRS, DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [May3,

marginal eminences opposite the basal joints of the legs; and the
labium is yellowish.

Abdomen cylindrical; the hinder extremity somewhat produced
and extending beyond the spinners. The fore extremity is dark
brown, succeeded by a strong transverse pale-yellowish or cream-
coloured band, the ends of which are produced backwards forming
a lateral broken marginal stripe on each side. The upperside
between these stripes is of a dull pale hue, closely reticulated with
brown in a vermiform pattern, and along the middle is a series of
four pairs of roundish pale spots covered with a shining satiny
pubescence ; the spots in each of these pairs are contiguous or
nearly so, and they decrease in size from the foremost to the last
pair, which are seated a little way from the posterior extremity of
the abdomen. The sides are dark brown, marked with various
subparallel undulating yellowish streaks, two largish dark-brown
patches on each side being left untouched by them. The underside
is deep brown, with a yellowish marginal line in front and on the
sides ; and beneath the produced portion is a longitudinal, central,
silvery-white band stopping short of the spinners. The genital
aperture is of great simplicity, consisting of a narrow transverse
opening covered by a short, scarcely projecting lip.

This handsome Spider is allied to several other African species,
such as NV. femoralis Luc., N. sumptuosa Gerst., and NV. keyserlingit
Blackw., but appears to be abundantly distinct. The only example
was not in first-rate condition.

Subfam. GAasTBRACANTHIN ®.

Gen. GasteRAcANTHA Walck.
GASTERACANTHA SODALIS, sp. n. (Plate XXXI. fig. 3.)

Adult female (No. 99), length (not including the posterior abdo-
minal spines) rather over 33 lines (nearly 8 mm.) ; breadth of abdo-
men (including the longest of the lateral spines) 62 lines (14 mm.).

This Spider is much like G'. madagascariensis Vins., to which it
is nearly allied, but the latter has the transverse abdominal black
bars broken off in the middle, and the abdomen itself is wider in
proportion from front to back than in the present species; the
cephalothorax also and the abdominal markings and spines are of
a deeper hue, in fact generally black, whereas in the present Spider
they are red-brown. The posterior and fore-lateral spines also are
longer and sharper pointed in G. madagascariensis, in which also
the underside of the abdomen is less thickly blotched with yellow,
and the sternum has a strong sub-triangular (or heart-shaped)
central, clearly defined, pale yellow spot; while in the present
Spider the abdominal blotching i is reddish orange-yellow, and the
sternum deep brown, with two small indistinct yellowish spots in
a transverse line near the middle.

The legs are of 2 uniform deep brown colour, while in G. mada-
gascaviensis the coxe and femora are of a brightish yellow-brown

[18]

1898.] INSECTS AND ARACHNIDS FROM SOCOTRA. 389

(in some examples, however, of the latter the legs are of a more
uniform dull yellow-brown).

Subfam. TETRAGNATHIN &.

Gen. TerracnatHa Walck.
TETRAGNATHA BOYDI, sp. n. (Plate XXXI. fig. 4.)

Adult female (No. 101), length 4 lines; length of cephalothorax
2 lines ; length of falces 2} lines nearly.

Cephalothorax oblong-oval, truncated at each extremity, and
widest near the middle ; length double its breadth ; lateral marginal
impressions at caput very slight; caput and margins of thorax
darker than the rest ; yellow-brown, but in the dry specimen the
colour is unreliable.

Hyes of posterior row equally separated; in a very slightly
curved line, the convexity of the curve directed forwards ; anterior
row much more strongly curved, but with the same direction of
the curve ; central quadrangle slightly broader than long, and the
fore side distinctly shorter than the hinder one; the fore-central
pair of eyes longest, and seated on a strongish rounded tubercular
prominence ; each of the lateral eyes also on a tubercle. The eyes
of each lateral pair are much nearer to each other than the fore-
central pair are to the hind-centrals. Clypeus rather less in height
than half the facial space.

Falces very long and projecting forwards, slightly longer than
the cephalothorax ; considerably divergent ; slightly curved, rather
constricted at the fore extremity. Fang more than three-fourths
the length of the falx, strong, abruptly bent at the base where it
is somewhat enlarged, and there is another somewhat shallow
dentiform enlargement towards the middle on the inner side; and
each of the falces is armed with a strong, somewhat curved, pointed
tooth at its extremity, just below the outer side close to the insertion
of the fang ; also on the inner side nearly beneath the base of the
fang is another strong sharp-pointed tooth ; besides these teeth each
falx has a double longitudinal row of others along the underside ;
those on the outside are most numerous (10 ?) and most equally
separated, the inner ones (7 or 8?) strongest and more confined
to the posterior portion of the falx, those of both rows diminishing
in strength as they run backwards.

Legs very slender; 1, 2, 4, 3, very little difference between 2
and 4; furnished with hairs and a few short slender spines.

Maville, labtum, and sternum normal.

The abdomen was so shrivelled and devoid of colour that nothing
can be said as to its colours or markings, which, however, are
most probably distinctive of the species.

This Spider is nearly allied to Yetragnatha tayluri Cambr.
(South Africa), but the relative position of the eyes is different, as
well as the form of the fang and the denticulation of the falces.

[19]

390 MESSRS. DIXEY, BURR, AND PICKARD-CAMBRIDGE ON [ May 3,

Fam. THOMISID2.
Gen. SELENoPS Dufour.

SELENOPS DIVERSUS, sp.n. (Plate XXX1I. fig. 1.)

Adult female (No. 100), length 10 lines.

General forin and structure normal.

Cephalothorax considerably broader (at its hinder extremity)
than long. Indentation at the junction of caput and thorax large
and deep. Colour dark yellowish brown, marked with somewhat
irregular pale markings and darker converging lines, and clothed
with short grey and other hairs, and with a rather dense marginal
row of short, curved, prominent spiniform bristles.

Eyes in normal position. The four centrals of the anterior row
form a slightly curved line, the convexity of the curve directed
forwards; the two central eyes are rather the smallest of the four,
and are further from each other than each is from the lateral eye
on its side.

Legs long, strong, furnished with spines and numerous hairs of
varying length; a scopula beneath the tarsi, and below the two
terminal claws a compact claw-tuft. The colour of the legs is
black, marked and, rather irregularly, annulated with reddish yellow-
brown; the annuli are clothed with coarsish grey hairs. The legs
do not differ much in length, the second pair are the longest, and
the first, apparently, slightly shortest.

Falces moderately strong, roundly prominent in front, straight,
and of deep blackish red-brown colour.

Mawille rather short, oblong, rounded at the extremity, inclined
towards the labium; colour dark yellow, pale yellowish at the ends,
where they are furnished with fine white hairs.

Labium slightly longer than broad, oblong, rounded at the apex,
which is yellowish, the rest deep brown.

Sternum rather longer than broad, short-oval, slightly truncated
before, and notched at its hinder extremity. Colour pale dull
yellowish brown.

Abdomen flattened, rather longer than broad, nearly quadrate,
truncate before, the posterior corners well rounded. Upperside
of a dull brownish hue, pretty thickly clothed with short greyish
hairs, some of those on the sides disposed in minute groups or tufts
giving it a spotty appearance; along the middle is a rather diffuse
but symmetrical black pattern, formed by a central longitudinal
stripe, emitting various spots and markings on each side, and
ending in a transverse angulated, black, irregular bar near the hinder
extremity : the surface of the abdomen from this bar to just above
the spinners is distinctly paler than the rest ; sides deep brownish,
and underside like the upper in colour. Genital aperture small,
but of distinctive form. Spinners short, of about equal length,
compact, and similar in colour to the underside.

This Spider is allied to S. dufourti Vins., and also to S. mada-
gascariensis Vins., but may be easily distinguished from the latter

[20]

1898.] INSECTS AND ARACHNIDS FROM SOCOTRA. 391

by the pattern on the abdomen, of which the form also is different.
In S. dufourti the pale posterior extremity shows on its posterior
border five distinct pale points; the present species shows only
three, and those somewhat irregularly defined. The present Spider
is also of much larger size than either of those mentioned, the
length given of S. dufourtt being 12 millim., and that of S. madagas-
cariensis 11 millim., whereas Selenops diversus measures in length
10 lines, or nearly as possible 21 millim.

EXPLANATION OF THE PLATES.

Puarn XXX.
Fig. i ae hoydi, sp. n., 5, p. 375.
6 » &, underside.
a
2a. ,, nop underside.

3. Papilio bennetti, sp. n., ¢, p. 381.
4. Pecilocerus sokotranus, sp. nov., 5 p. 384.

Puate XXXII.
Fig. 1. Selenops diversus, sp. n., 2, p. 390.

E gener oA Genital apertures.
2. Nephila hennetti, sp. n., 9, p. 387.
2a. a Genital apertures,

3. Gasteracantha sodalis, sp.n., 2, p. 388
4. Tetragnatha boydi, sp. u., p. 389. me and eyes.
4a&4b. ,, 5 is Falx and fang in two positions.

[2]

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FredXO Pickard Cambridge delet ith Mintern Bros.imp
INSECTS FROM SOCOTRA.

PZ 5 1806 eons

Fred*0.Pickard-Cambridge delet lth.

Mintern Bros.imp.

ARANEIDEA FROM SOCOTRA.

ONE COLE CrTON
OF

INSECTS AND ARACHNIDS

MADE IN 1895 & 1897,

BY

Mire CVO AL PER, FE 7.8,

WITH DESCRIPTIONS OF NEW SPECIES.

BY
CG. V. A. PEEL, F.Z.S., | GILBERT J. ARROW,
E. E. AUSTEN, R. McLACHLAN, F.RS.,
F. A. DIXEY, M.A., M.D., | MALCOLM BURR, F.ZS.,
HERBERT DRUCE, F.LS.., AND
F.Z8., R. I. POCOCK,
C. J. GAHAN, M.A.,

[From the PROCEEDINGS OF THE ZooLo@icaL Socrery oF Lonpon,
January 23, 1900. |

4 MR. C. V. A, PERL AND OTHERS ON (Jan. 23,

On a Collection of Insects and Arachnids made in 1895
and 1897, by Mr. C. V. A. Peel, F.Z.S., in Somaliland,
with Descriptions of new Species. By C. V. A. Pret,
F.Z.S., E. E. Austen, F. A. Dixry, M.A., M.D.,
Herbert Druce, F.L.S., F.Z.S., C. J. Ganan, M.A.,
GILBERT J. Arrow, R. McLacutan, F.R.S., Matcoumm
Burr, F.Z.S., and R. I. Pocock.

(Plates I.-IV.)

CONTENTS.
Pago
1. Narrative of the Expeditions. By C. V. A. Pre, F.Z.S....... 4
DAS ID ho Weres, Ned D, WOE UA NSO? “Gpoqdaoadcasacadoseoacosbaagadacegaceesede a
3. Lepidoptera Rhopalocera. By F. A. Dixny, M.A., M.D....... 10
4. Lepidoptera Heterocera. By Hersert Drvcr, F.Z.8., F.L.8. 17
5. Coleoptera. By C. J. Ganan, M.A., and Gitpert J. Arrow... 21
6. Neuroptera. By R. McLacuuan, F.R.S. .................c0cceeeeee 34
7. Orthoptera. By Matcoim Burr, F.Z.S., and Dr. C. Bronnur
VW: WATTENWIY Ii cwss ch sianioecssigecicaeareieceneuretbancsiapmseseenceogse 35
8. Insects of other Orders. By various Contributors............... 46
9. Chilopoda and Arachnida. By R. I. Pocock .................2.+8 48
10. General List of the Scorpions of Somaliland and the Boran
@ountry ans yates OCOCKSa-peseecereaeeeeeeeseet eee seeererce 55
I explanationvoftheselatestascssscacsssceccecsteesteeceeeeceee caer 63

1. NARRATIVE OF THE EXPEDITIONS.
By C.V. A. Punt, F.Z.S., F.R.GS.

(First expedition to Somaliland. April 16 to August 7, 1895.)

Somaliland has always been known as the big-game hunter’s
paradise, and in consequence but few have had the energy to
collect insects as well. Mr. BE. Lort Phillips and Dr. Donaldson
Smith, however, are notable exceptions. I always kept a killing-
bottle in my tent and also collected outside whatever I happened
to meet, but I must honestly confess that I never devoted my time
seriously to collecting imsects. Owing to the great sameness of
thorn-bush and undergrowth, the Lepidoptera of Somaliland are dis-
appointing. The Butterflies are few and somewhat uninteresting,

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND. 5

and the Moths are very local. The latter were nearly all
collected on the banks of river-beds where there were trees, long
grass, and undergrowth. The Beetles, on the other band, simply
swarmed, and there was also no lack of Orthoptera, Chilopoda,
Diplopoda, Arachnida, &c.

On April 20th, 1895, 1 started from Berbera, the coast town of
Somaliland, wending my way south-west along the maritime plain
to Hargaisa. Insect-life was by no means plentiful in this hot
parched-up desert country, but birds were numerous by the sides
of the dried-up river-beds.

Hargaisa is a permanent Somali village on a gentle slope, over-
looking a river-bed. There was luxuriant undergrowth and a
few trees. Insect-life was consequently more plentiful, and I
collected my first butterflies and moths here. Of the latter, the
large species Cyllogramma latona Cr. and Sphingomorpha chlorea
Cr. came to my lantern in great quantities at night. After
leaving Hargaisa, I went south across the great waterless Haud
District, through dense thorn-bush jungle. Hn route we suddenly
emerged upon the Bun Saylah,a large open plain literally covered
with game, notably Oryx, Hartebeest, Seemmerring’s Gazelle, and
Ostriches. It took usa day to cross this, and then we entered dense
jungle again. All this time I did but little insect-collecting, as,
owing to the scarcity of water, | was always on the quick march.
At Sassabanah we encountered water in deep wells and under the
surface of a river-bed, the Webbi Jerrar. Here I collected some
of the ticks described. Thence] marched to the Boorgha Country,
and passed the everlasting red sand, entering a stony, hilly district.
In this latter, Orthoptera were very numerous. I went as far
as Mount Kuldush, marching along the Webbi Shebeyli, the great
river of Somaliland.

The banks of this river were lined with trees and dense jungle,
and butterflies and other insects were very abundant. Being
unable to find a path for the camels down to the river-edge, and as
I was running short of food, and many of my men had fever, I
retraced my steps at the end of June. I followed the Sule River
for a long way and reached Bun Jijjiga, a gigantic plain at the
foot of the Harah Hills. Here I fell in with Abyssinians, who,
however, behaved most cordially. Game was extremely abundant
on this plain; but insects were somewhat scarce except at W hardi
Datal, where there was long grass, in which Orthoptera simply
swarmed. After leaving the plain I traversed thick jungle until
reaching Hargaisa, where | rested to take up water before recrossing
the great maritime plain called Guban (the hot country).

During this latter journey I came across a small herd of the
Somali Wild Ass (Equus somalicus), and passed through a dense
locust-cloud, which darkened the sun for hours and looked like a
great fall of snow, the air being white with them. After a very
exhausting march through this desert, where we encounted terrific
dust-storms every day, we finally returned to Berbera and the coast.

6 MR. C. V. A, PEEL AND OTHERS ON ' {Jan. 23,

(Second expedition. June 5th to October 29th, 1897.)

Being anxious to continue my researches in the Natural History
of Somaliland, and if possibie to cross the Ganana River and
explore Lake Rudolph, I set out from Berbera again in 1897,
accompanied by Mr. J. Benett-Stanford, F.Z.S., and his wife.
We took with us a very large caravan of camels and escort. We
crossed the great Gulis Range of mountains by the Gerato Pass,
and encamping at Lehello, there awaited more camels, which were
being bought for us in Berbera. Lehello lay in stony ground by
the everlasting dried-up river-bed. Here insect-life was fairly
abundant, as will be seen from a perusal of the subsequent parts
of this memoir.

We removed next to the Toyo Plain, a vast open space devoid of
trees, and then crossing the waterless Haud District reached
the wells of Farfanyer in Ogaden. Here we were overtaken by a
messenger from the political officer at the coast, forbidding us to
go to the Ganana Ktiver, as the Somalis were reported to be
fiehting the Abyssinians there. This was a great disappointment
to us, after getting together such a large caravan and escort. At
Farfanyer we fell in with a great number of Somalis armed with
rifles, which they had looted from the Abyssinians, whom they had
defeated in two battles on the Webbi Shebeyli. At Farfanyer
Mr. and Mrs. Stanford stayed, looking for rhinoceros, whilst I,
with my separate caravan, went in search of a reported lion.
Eventually I lost the Stanfords altogether, and although I sent
guides to look for them, it was of no use, owing to the denseness
of the bush. I hada very bad attack of fever, and after waiting for
over a fortnight to try and get news of my friends, or to get a guide
to take me back, I was obliged to give it up and march on alone.

Hearing of elephants in the Marehan and Haweea Countries, [
determined to explore this little-known territory, and so marched
across the great Marehan Desert and reached Habr Heshi, where
we at length found water. Here I shot a fine lion which charged
me, but I managed by great good luck to stop the animal when it
was within a few yards.

After leaving Habr Heshi we encountered stony barren country,
and pitched camp at Sinnadogho in the Haweea country, where:
there were fresh tracks of elephants. Here the natives were
extremely troublesome, and I had a very anxious time. Several of
my rifles, cloth for barter, and a pony were looted from me, and
my followers were frequently attacked. I marched to Joh, the
furthest point south-east [ was destined to reach. We were then
about six days’ distance from the east coast. Finding no elephants,
T went on to Kadea, looting on the way a pony in exchange for
the one taken from me. Upwards of five hundred armed men
made their appearance at this point, and 1 thought we were in for a
big fight, some of the young men dancing ihemaelnce into a perfect
frenzy. J was obliged to tire over their heads, to keep them from
looting my camels. They succeeded, however, in stealing some

[4]

nd

1900.1 INSECTS AND ARACHNIDS FROM SOMALILAND. ‘

rifles, and I was obliged to take out my little army twice against
villagers ; but luc ‘ily, no blood was shed, as the villagers, seeing the
rifles coming, immediately restored my stolen property. I did no
collecting to. speak of here, as I was ill with fever and was having
avery anxious time. I could get no guide to take me across the
great waterless desert of the Merchant and was obliged to load up
the water-vessels at Doosa Moreb and start without one. I believe
I was the first white man to visit the heart of the Marehan and
Haweea Countries, and was right glad to shake off the dust from
my feet on quitting those inhospitable tribes. How I lost my
way crossing the Marehan Desert, ran short of water, and all but
died of thirst, I have already descrihed in the pages of the ‘ Wide
World Magazine.’ We reached Galadi in the Mijertain Country,
and found water in the very nick of time, when I was almost at
the last gasp. Here I became delirious, and knew nothing that
was going on around me for hours. After leaving Galadi I became
so ill and weak with fever that I did no further collecting, but was
practically carried by my pony the whole way across the waterless
Haud again to the Gulis Range, where I remained a few days to
rest, and at length reached Berbera more dead than alive.

A full account of my two expeditions, together with a complete
list of every mammal and bird known to inhabit the country, will
be found in my book *Somaliland,’ published in 1899 by Messrs.
F. E. Robinson & Co., London.

The specimens mentioned and described in the following pages
are in the Hope Collection, University Museum, Oxford, with the
exception of those which are expressly stated to be in the British
Museum.

2. DIPTERA.
By E. E. Austen, Zoological Department, British Museum.

Mr. Peel’s collection of Diptera was not extensive, amounting
only to four specimens belonging to three species, one of which,
however, is apparently new.

Fam. TABANID &.
Subfam. PANGONIN#®.
Panconta Latr.
PANGONIA (sens. strict.) Rond.

PANGONIA TRICOLOR, sp.n. (Plate I. fig. 8.)
2. Length 17 millim. ; length of wing 15°5 millim.; length of
roboscis 4 millim.

Shining black ; first and second segments of abdomen (except a
somewhat trian care area in the middle of the second segment, which,
however, like the first and remainder of the second se gment, is clothed
with appressed silvery-white pile) white above; sixth and seventh
segments and the narrow postervor margin of the fifth ochraceous, and

[5]

8 MR. C. V. A, PEEL AND OTHERS ON [Jan. 23,

clothed with golden ochraceous pile; wings dark brown; alula,
greater portion of the area behind the sixth longitudinal vein, and
sometimes a narrow margin extending from the tip of the second vein
to the apex of the anal cell, hyaline.

Head with an area surrounding the bases of the antenne,
extending from eye to eye, and including the lowest third of the
front and an equal space below the antenne, covered with white
dust ; face on each side below the antenna sparsely clothed with
fine silvery hairs ; cheeks dark brown ; occiput covered with greyish
dust, and base of head below thickly clothed with short white
hairs ; antenne uniformly black, a distinct shoulder at the base ot
the third joint above. Thorax with a few short golden hairs in
front of scutellum ; pectus clothed with silvery-white pile, which
extends on to the pieurz above the front coxe, and also in a stripe
running up to the base of the wing, where the stripe ends in a
fork; a narrow stripe of silvery-white pile extends from the base
of the scutellum to the wing on each side. Abdomen: the white
posterior margin of the second segment is narrowed in the middle
above (thus leaving the black triangular area mentioned in the
diagnosis), and continued on the ventral side as a narrow trans-
verse band. Legs: cove greyish pollinose, and clothed with
silvery-white pile; thie with a slight reddish tinge. Wings
with a fleck of silvery-white pile on the base of the first vein ;
halteres tawny.

Two specimens (both 2). Type in British Museum; co-type
im Hope Museum, Oxford. From Bun Feroli, north of Shebeyli
River, West Somaliland; June 10-20, 1895: “biting men and
animals.”

Jn the present species the eyes are bare and the first posterior
cell of the wing is closed; it is therefore a true Pangonia in
Rondanr’s restricted sense.

Pangonia tricolor is closely allied to P. briechettii Bezzi (Ann.
Mus. Civ. Genov. xxx. (1892), p. 181), also from Somaliland
(Milmil). P. tricolor differs from P. bricchettii (which apparently
is a somewhat smaller species) znter alia in only the first two, and
not the first four’, abdominal segments being marked with white,
thus leaving between the white of the base and the ferruginous tip
a broad shining black space, which is absent in Bezzi’s species.

It may be noted that in the marking of the base of the abdomen
of Pangonia tricolor there is a certain similarity to Tabanus
leucaspis, v. d. Wulp (Notes Leyden Mus. vii. (1885), p. 74, pl. v.
fig. 3), from the Gold Coast.

The collection of the British Museum contains two specimens
of Pungonia tricolor, obtained by Capt. Swayne in Somaliland, from

' There is a discrepancy between Bezzi’s diagnosis and his detailed descrip-
tion ; in the former he writes (op. cit. p. 181) “ abdomine fasciis tribus trans-
versis ex tomento allido ad marginem posticum segmentorum,” while in the latter
he describes (p. 182) the fourth segnient also as ‘a orlatura posteriore bianca,”
with the dorsum “ Fernito di peli bianchi”; he deseritbea the 5th, 6th, and 7th

segments as “a peli ferruginosi.”

[6]

1900.) INSECTS AND ARACHNIDS FROM SOMALILAND. 9

nearly the same region as that in which Mr. Peel’s specimens were
found. Capt. Swayne also captured a single specimen of another
species of Pangonia (too much damaged for determination), and
three examples of a smali Tabanid, somewhat resembling a Hemato-
pota in form, but with clear wings; as the latter specimens are
headless, it is impossible to determine them more precisely.

The following extract from a letter from Capt. Swayne, sent along
with the flies to Dr. P. L. Sclater, is interesting as showing the
apparent effect of the bites of these flies upon domestic animals. It
is possible, however, that the real offender in these cases is either
Glossina longipennis, Corti (the Somaliland Tsetse-fly), or else a
species of Stomovys, which abounds all over E. Africa. The latter
species was found by Dr. J. W. Gregory to kill his camels on the
Tana River, and was discovered by Capt. A. G. Haslam, A.V.D.,
to carry the Trypunosoma of Tsetse-fly disease. Since Stomoarys is
a fly of small size, while Glossina longipennis is in shape not unlike
a Hematopota, the true culprits may escape notice, the effects of
their bites being attributed to the Tabanide. In the extract from
Capt. Swayne’s letter the Pangonia are called “ Doog,’ and the
small Tabanid “ Balaad.” Capt. Swayne writes as follows :—

‘“‘T send you three specimens of ‘ Doog’ (a large fly) and three
specimens of ‘ Balaad’ (a small fly) ...... I was very
much pestered by ‘ Doog’ on my way through Ojaden to the
Webbe Shabeyli in Somaliland. They swarmed on my camels,
constantly drawing blood. The other fly, ‘ Balaad, which looks
not unlike the common house-fly, is far the worst fly on the
Webbe; a valuable camel, on which I caught three or four, two
months ago, is now dying, and the Somalis say that this is due to
the bites of ‘ Balaad.’ If there are many of them they kill horses
and camels, and the Somalis will not have their live-stock grazing
where ‘ Doog’ and ‘ Balaad’ are found.”

Fam. ASILID4,
Subfam. LAPHRIN®.

LamMyra Loew.
LAMYRA VORAX Loew.

Lamyra vorax, Loew, Ofvers. af K. Vet.-Akad. Forhandl. 1857,
355. 47; id. Dipt.-Fauna Siidafrika’s, 114 [1860].

A single 2, West Somaliland, between April 16 and Aug. 7,
1895.

I refer Mr. Peel’s specimen to this species with some
hesitation. Its length is 15 German lines, instead of 11 or 12;
the pollinose spots on the second and third abdominal segments
are practically invisible; and there are differences in the length
and coloration of the hair on the ventral surface of the abdomen.
The specimen, however, is in poor condition, and even should it
eventually prove to belong to a new species, it is too much
damaged to be selected as a type.

[7]

10 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

Fam. Musc1rp &.
GLosstna Wied.
GLOSSINA LONGIPENNIS Corti.

Glossina longipennis, Corti, Ann. Mus. Civ. Genoy, xxxv. (1895)
p- 138.

A single 2, West Somaliland, June 23-25, 1895.

Mr. Peel’s note on this specimen says :—‘ Fly-belt sharply
defined from Biermuddo to Boholo Deno.”

This species, which is the Somaliland T'setse-fly, was described
from a male specimen obtained by Capt. Vittorio Bottego in June,
1893, on the Uelmal River, in the Boran Galla country. The
British Museum possesses four examples from Somaliland (the
exact locality not being known), collected and presented by
Mr. Th. Greenfield.

Corti states (loc. cit. p. 139) that G. longipennis is “related to
G. tachinoides, Westw., but differs in having the antenne yellowish
and not brown.” It is, however, much more closely allied to
G, tabaniformis, Westw., in which the length and size of the wings
are even greater.

3. LEPIDOPTERA RHOPALOCERA.
By F. A. Dixry, M.A., M.D., Fellow of Wadham College, Oxford.

DANAIN#.
LIMNAS CHRYSIPPUS Linn.

Twenty-two specimens: 16 ¢,6 92. It is remarkable that not
one of these is of the type form, 14 ¢ and 6 Q being var. Aluyit, in
which the black and white of the apical portion of the fore wing
are wanting; while the remaining 2 ¢ are var. dorippus, which
resembles var. kluguv in every respect except that both surfaces of
the hind wing are more or less suffused with white as in the
aleippoides form of the type. The dates and places of capture
were as follows: Hargaisa (North-west Somaliland), April 25-28,
1895, klugu, 4 3, 1 2, dorippus, 1 $; Arigumeret, Farfanyer
District (Central Somaliland), June 20, 1897, in thick bush, klugii,
4 3, dorippus, 1 $; Haud, Odewein (North Central Somaliland),
June 23, 1897, in dry river-bed with thickly wooded banks,
klugu, 1 6,192; Haud District, Eyk (North Central Somaliland),
July 2, 1897, klugu, 1 ¢,1 9; Habr Heshi, Marehan Country
(East Central Somaliland), Aug. 26, 1897, in thick bush, klugii,
Joys on oe

The ground-colour of the present specimens varies, the majority
being of the usual light reddish amber seen in Oriental specimens
of the type. Two or three of the klugii are of a pale dull fawn,
and one or two approach the duller and darker ground-tint of the
African chrystppus. These differences are not sexual, and there
are transitional forms. The marginal white spots of the hind wing

[8]

1900. | INSECLS AND ARACHNIDS FROM SOMALILAND. sla

are generally obsolete on the upper surface; they are, however,
conspicuous in one of the two dorippus. Most of the male kiugi
show a slight powdering of white scales in the neighbourhood of
the submedian scent-patch.

In describing the collections made fifteen years ago in Somali-
land by Col. Yerbury and Messrs. Thrupp, Lort Phillips, and
James (Proc. Zool. Soc. 1885, p. 756), Dr. Butler remarked of
L. klugu: “This is clearly the prevalent Zimnas in Somaliland ;
L. chrysippus and L. alcippus having, apparently, entirely
disappeared, and ZL. dorippus being scarce.” The fact that the
only specimens of LZ. chrysippus obtained by Mr. Peel in his two
distinct visits to Somaliland were of the klugu and dorippus
varieties, confirms the above conclusion. It is also worthy of
note that the collection made by Capt. Swayne in the Harar
Highlands (Proc. Zool. Soc. 1898, p. 821) contained L. klugu, but
no specimens of L. chrysippus, L. alcippoides, or L. dorippus.

ACRAEIN&.
ACR#A NEOBULE Doubl.

Four specimens: 2 ¢,2 9. Caught at Gonsali (West Somali-
land), June 24, 1895.

ACREA DOUBLEDAY! Guer.
1 2. Gonsali, June 24, 1895.

ACRZA SERENA Fabr.

3 9. Gonsali, June 24,1895. In one of these specimens the
subapical oblique dark bar on the fore wing is well marked, in the
other two it is almost or altogether absent. One of the latter
has the inner portion of the hind-marginal dark border of the fore
wing almost obsolete, except near the apex.

ACR#A MIRABILIS Butl. (Plate I. fig. 4.)

Acrea mirabilis, Butl. Proc. Zool. Soc. 1885, p. 760, pl. xlvii.
fies i

Seven specimens: apparently 5 ¢,2 92. The British Museum
possesses five specimens of this interesting Acrewa, including the
trpes. The present specimens differ from those in the National
Collection (one of which retains a violet bloom) in the following
particulars :—(1) The upper surface is generally of a warmer
chestnut tinge. (2) The paler marks beneath, including the sub-
apical patch of the fore wings, are rich yellow-ochre, instead of
being nearly white as in the British Museum examples. In one
of the latter, a female, the subapical patch on the upper surface
also is creamy white. (38) In the present specimens there is a
definite yellow band immediately external to the darker median
band of the hind wing beneath. This is not the case in the British
Museum examples, where the drab submarginal area gradually
pales inwards towards the darker median band. In the @ type

(9

12 MR. C. V. A. PERL AND OTHERS ON [Jan, 23,

this submarginal pale area is somewhat more distinct than in
the other British Museum specimens, but it does not reach the
condition seen in Mr. Peel’s examples. The dated specimens in
the National Collection were taken at Bundu Maria, Somaliland,
in April. Mr. Peel’s were all captured at Aoho, near Hodayu,
Ogaden Country, Central Somaliland, on Aug. 20, 1897. The
country consisted of stony hills, with thick bush. From the dates
it seems probable that the present specimens belong to the wet-
season, and the British Museum specimens to the dry-season form
of the species.

NYMPHALINA.
JUNONIA CEBRENE Trim.

Six specimens, all males. Three were captured at Hargaisa,
April 25-28, 1895; the other three in the summer of 1897, two
bearing the date June 20, and the locality Arigumeret, Farfanyer
District. These latter have the underside generally darker and
more speckled than the spring examples; this is less apparent in the
third specimen, from Central or East Somaliland, June 5—Oct. 29,
1897.

JUNONIA CLELTA Cram.

Six specimens: 36,39. Hargaisa, April 25-28, 1895. The
undersides of these specimens vary, but in all the ocelli are more
distinct and the general tint is less uniform than in the ordinary
“‘dry-season ”’ form of the species.

Junonra TAVErA Rogenh.
One male. Hargaisa, April 25-28, 1895.

Bysuta tirayia Drury.

Four specimens: 3 6,19. These are of the ‘“ intermediate ”
seasoval form, the female verging towards ‘‘wet”'. All are dated
Hargaisa, April 25-28, 1895.

HYpoLimnas Misippus Linn.

Twenty-eight specimens: 26 ¢, 2 9. It is very remarkable
that of the only two female specimens obtained by Mr. Peel, one
should be of the ordinary form, resembling the type of L. chry-
sippus, and the other of the var. alcippoides, differing from the
former only in the whitish suffusion on both surfaces of the hind
wing. From the facts given above (see under L. chrysippus, p. 10),
it would appear that the form klugii of L. chrysippus occurs in
Somaliland to the exclusion of the type, and it might have been
expected that the form of H. misippus 2 which so closely resembles
klugii, viz. H. maria Cram., would have been the form similarly

' For a discussion of geographical and seasonal forms in the genus Byblia
Hiibn., with especial reference to the relations between the forms occurring in
Somaliland and Socotra, see Dixey, Proc. Zool. Soc. 1898, pp. 376-379.

[10]

_ 1900. | INSECTS AND ARACHNIDS FROM SOMALILAND. 13

prevalent in that region. So far as the evidence of Mr. Peel’s
collection goes, the reverse is the case. Another remarkable fact
in the distribution of these parallel forms is that while H. klugw is
extremely rare in India, the corresponding variety of H. misippus
2? occurs there not infrequently’. It is also worthy of note
that the white-winged West-African form, Limnas aleippus Cram.,
is accompanied by the ordinary, and not the white-winged, form of
H, misippus 2.

The dates and localities of the present examples are as follows :—
Hargaisa, April 25-28, 1895, 6 ¢, 1 2 (ordinary type); Arigu-
meret, Farfanyer District, June 20, 1897, in thick bush, 2 g,1 2
(var. alcippoides Butl.); Bally Maroli, Haud District (North
Central Somaliland), June 25, 1897, in open plain, 14 g; Hyk,
Haud District, July 2, 1897, 3 ¢. One other male was taken in
Central or East Somaliland between June 5 and October 29, 1897,
the exact locality being uncertain.

HAMANUMIDA D&DALUS Fabr.

One male, Hargaisa, April 25-28, 1895. The underside is of
the ‘‘dry-season” form, though not extreme.

LYCANIN2.

PoLyomMatus Ba&Ticus Linn.

Two specimens, both males. On the thickly wooded banks of a
dry river-bed, Haud, Odewein, June 21 & 28, 1897.

PLEBEIUS TROCHILUS Freyer.

Two females. Gerato Pass, Goolis Range (North-west Somali-
land), June 9, 1897.

AZANUS Jusous Guer.

Five males. Of these, four were captured on the dry sandy
plateau of Edegan in the Haud District (North Central Somali-
land), July 9, 1897; the remaining one was taken at Joh in the
Haweea Country (East Central Somaliland), Sept. 20, 1897.

AZANUS THEBANA Stder.

Lycena macalenga, Trim, 8.-Afr. Butterfl. vol. ii. p. 74 (1887).

Three specimens: 1 ¢,2 2. One pair from Odewein, Haud,
June 21-23, 1897, dry river-bed with thickly wooded banks;
the other female from the sandy plateau of Edegan, in the same
district, July 9, 1897.

LYCZNESTHES PRINCEPS Butl.

Two females apparently belonging to this form, though some-
what smaller than the type, which came from Abyssinia. Edegan,
Haud District, July 9, 1897.

* See Swinhoe, Journ. Linn. Soc., Zool. vol. xxv. pp. 340, 341. For a
summary of the facts at present known with regard to the distribution of the
forms in question, see Poulton, ‘Nature’ July 6, 1899, p. 223.

[11]

14 MR.C. V. A. PEEL AND OTHERS ON [ Jan. 23,

SPINDASIS SOMALINA Butl.

Spindasis somalina, Butl. Proc. Zool. Soc, 1835, p. 764, pl. xivil.
fig. 5.

Two specimens, both males. Webbi Shebeyli, near Mount
Kuldush (West Somaliland), June 28, 1805.

ToLaus NURSEI Butl.

One male. Webbi Shebeyli, near Mount Kuldush, June 28,
1895.
PIERIN#.

CATOPSILIA FLORELLA Fabr.

Forty-three specimens; 40 ¢, 3 2. Of these, 34 d on July
2.1897, and 1 2, July 4, 1897, were caught at a pool in the open
plain of Hyk in the Haud District. The other captures were as
follows:—Bun Feroli, N. of Shebeyli River (West Somaliland),
June 10-20, 1895, 1 ¢; Odewein, Haud District (dry river-bed),
June 21, 1897, 4 o, 2 2; Bally Maroli, Haud District (open
plain), June 25, 1897, 1 ¢. The males, even those caught on the
same day, vary much in size; in a few of the smaller examples the
freckling of the underside is very faint or absent. The three
females are all of the yellow form.

Contas MARNOANA Rogenh.
One female. Hargaisa, April 25-28, 1895.

TeRIAS HAPALE Mab.
Two males. Hargaisa, April 25-28, 1895.

TERACOLUS EUPOMPE Klug.

Seven specimens; 4¢,3 9. These were captured as follows :—
a “dry season” 3, and “intermediate” 3 & ? at Hargaisa, April
25-28, 1895; a “ wet season” ¢ in the dry river-bed at Odewein,
Haud, June 23, 1897; a “wet season” ¢ & @ on the plateau of
Edegan, July 9, 1897.

A remarkable form, apparently of the female of this species,
was caught on the Sule River (West Somaliland), May 29, 1895.
The wings have no marginal black except at the apex of the fore
wing, and no biack at the bases except a slight dusky powdering
like that of the “dry season” male. The crimson apical patch
has no chain of submarginal dark spots, but an ill-defined inner
dark border to the patch is present, widened between the second
and third median nervules. Beneath, there is only a very faint
indication of the crimsen apical patch, but the submarginal chain
of dark spots in the fore wing is well-developed, except that there
is no spot between the first and second branches of the median.
The hind wings show a faint drab irroration, and the submarginal
spots are ill-developed. The first submarginal spot of the fore
wing, and the first two of the hind wing, together with the discoidal
spot of the hind wing, are pale-centred, looking like incipient ocelli.

(12)

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND. 15

The “intermediate ” female shows in some respects an approach to
this condition, and the specimen just described may perhaps be
considered as an extreme “dry season” form, though this wouid
scarcely be expected in view of the recorded date.

TERACOLUS OMPHALE Godt.

Two specimens; ¢ & 9. These were both caught at Hargaisa,
April 25-28, 1895. They are of the “ intermediate” form, and
are smaller than the average size of the species.

TERACOLUS PHILLIPSI Butl.

Teracolus phillipsi, Butl. Proc. Zool. Soc. 1885, p. 772, pl. xlvii.
figs L.

Nine specimens; 6 5, 3 9. The males are all of the “ wet
season” form; four were taken on July 4, 1897, at a pool in the
open plain of Eyk, in the Haud District; the remaining two on
the dry plain of Edegan in the same district, on July 9, 1897.
Two ‘“ wet season” females were taken at Odewein, Haud, on June
21, 1897; and a “wet” or “intermediate” female at Hargaisa,
April 25-28, 1895.

TERACOLUS HELVOLUS Butl.

Teracolus helvolus, Butl. Proc. Zool. Soc. 1888, p. 94.

Seven specimens; 5 ¢,2 9. Four males and one female at
Hargaisa, April 25-28, 1895. These are of the “dry season”
form. <A ‘wet season” ¢ & 2 were taken in the river-bed at
Odewein, June 23, 1897, and at the pool of Eyk on July 4, 1897,
respectively.

TERACOLUS HELIOCAUSTUS Butl.

Teracolus heliocaustus, Butl. Proe. Zool. Soc. 1885, p. 768,
pl. xlvi. figs. 8 & 9.

One male. Odewein, June 21, 1897.

TERACOLUS PROTOMEDIA Klug.

One male. Sibi (West Somaliland), May 27, 1895.
HERPENIA MELANARGE Butt.

Herpena melanarge, Butl. Proce. Zool. Soc. 1885, p. 774.
Herpenia iterata, Butl. Proc. Zool. Soc. 1888, p. 96.

One male. Sibi (West Somaliland), May 27, 1895. A “wet
season ” form.

BELENOIS PEELI, sp. nov. (Plate I. figs.5 ¢,6 9.)

Types (¢ & 2) in Hope Museum, Oxford.

Exp. al., ¢ 50 mm., 2 62 mm.

Male. Above white, with a pearly lustre at the base of the
wings as in B. gidica, B. abyssinica, &c., best marked in the fore
wing. Costa of the fore wing with a very thin edging of black.
A black marginal spot at the termination of each of the following

[13]

16 MRC. V. A. PEEL AND OTHERS ON [Jan. 23,

veins :—in the fore wing the 2nd and 8rd subcostal, the two radial
and the three median branches; in the hind wing the 2nd subcostal,
the radial, and the three median. ‘These marginal spots are oval
in the hind wing, the long axis being parallel with the border of
the wing; in the fore wing they are more or less triangular, with
their bases at the margin. The first two at the apex of the fore
wing are more or less fused; the remainder in both wings are
distinct. There is a slight submarginal powdering of black scales
in the fore wing between the 2nd and 38rd subcostal, and also
between the two radials. A much fainter powdering occurs
between the 3rd subcostal and upper radial, and the slightest
possible trace of a similar powdering between the 2nd and 38rd
median. The marginal spots and the submarginal powdering mark
out between them a very indistinct series of white subapical spots,
the first three more or less wedge-shaped, the fourth nearly
circular ; these are barely to be separated from the general white
surface. Beneath, the wings are white; there is a thin dusky line
along the costa of the fore wing, and dark marginal spots, like
those of the upper surface but smaller, occur at the extremities of
the same veins in both wings. There is an orange-vellow patch at
the base of the fore wing, occupying about one-fifth of the cell, and
somewhat prolonged along the course of the subcostal vein. The
same orange-yellow colour occurs in the hind wing as a streak along
the costa, as a well-defined patch between the roots of the median
and submedian veins, and as a median chain of spots crossing the
disc of the wing from the costa to the inner border; these latter
are seven in number, one occupying each interspace posterior to
the costal vein except the space between the second subcostal and
discoidal, and that between the second and third median. The
third of the series surrounds the discocellular venule. This and
the four succeeding spots are fairly distinct; the other two are
faint. There is also a very slight indication of a submarginal
series of dull yellow spots running parallel with the hind border
of the wing.

Female. Above, like the male, but with the marginal dark spots
somewhat larger. These are triangular in the hind wing, and in
the fore wing become fused towards the apex into a narrow marginal
band. The dark subapical powdering forms more definite and
larger patches than in the male, and the white subapical spots are
consequently more distinct. There is a trace of a marginal dark
spot at the termination of the submedian vein of the fore wing,
which is hardly if at all visible in the male. The basal pearly
gloss is well marked, and the basal orange-yellow shows slightly
through from the lower surface. Beneath, as in the male ; but the
basal orange-yellow occupies from one-third to one-half of the cell
instead of only one-fifth. The submarginal series of spots is some-
what more distinct than in the male, and appears to be of the same
orange-yellow as the median series.

In both sexes the first subcostal branch of the fore wing coalesces
with the costal. The wings of the male are slightly more pointed,

[14]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND. iY

as in B, gidica, &c., than those of the female. This species can
readily be distinguished from any other of its genus by the entire
absence of a dark discoidal spot from both surfaces of both wings
in each sex, and by the presence of a double row of yellow or
orange spots, unaccompanied by any dark markings, on the under-
side of the hind wing. It is probably a “dry-season” form, and
perhaps most nearly recalls the dry-season B. abyssinica Luc., but

it is far less heavily marked.
Two specimens; ¢ & 2. Sule River, West Somaliland, May
29, 1895. Both, especially the female, somewhat worn.

BELENOIS MESENTINA Cram.

Nine specimens; € ¢,3 2. Dates and localities as follows :—
Sule River, May 29, 1895, 2 ¢; Odewein, Haud, June 21, 1897,
26; 12; pool at Eyk, Haud District, July 4, 1897,2 5,19;
summer or autumn of 1897 (Central or East Somaliland), 1 9.

NYcHITONA MEDUSA Cram.
1 2. Hargaisa, April 25-28, 1895. This is of the form alcesta
Cram.

MYytLoraris AGATHINA Cram.
1 2. Sibi, May 27, 1895.
PAPILIONIN &,
PAPILIo DEMOLEUS Linn.
Four specimens; 2 ¢, 29. WebbiShebeyli, near Mt. Kuldush,
June 27, 1895.
H&ESPERIID®.
SARANGESA ELIMINATA Moore.

Two females, one at the Webbi Shebeyli, June 28, 1895; the
other at Odewein, Haud, June 21-23, 1897.

RHOPALOCAMPTA ANCHISES Gerst.

4 $,2 9. Two males and a female at Odewein, Haud, June
21-23, 1897; the remainder in the summer or autumn of 1897, in
Central or East Somaliland, the exact locality being uncertain,

4, LEPIDOPTERA HETEROCERA.
By Hersert Drvcs, F.L.S., F.Z.S.

Mr. Peel’s collection of Moths includes specimens of four new

species.
ARCTIADS,

SEcusio srrieata Wk,
North-west Somaliland, Hargaisa, April 25-28, 1895. One

example.
Proc. Zoon. Soo.—1900, No. II. 2
[15]

18 MR. C, V. A. PEEL AND OTHERS ON (Jan. 23,

DEIOPEIA PULCHELLA Linn.

N. Central Somaliland, Odewein, in the Haud District, June
21-23, 1897. In dry river-bed with thickly wooded banks. Two
specimens.

Nocruip.

HELIOTHIS ARMIGHRA Hubn.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

Eupbexta oppostra W1k.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

AMYNA SELENAMPHA Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. ‘Two specimens.

AMYNA octo Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. ‘Two specimens.

TARACHE OAFFRARIA Cram.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. Two specimens. Somaliland, 1895 or 1897. One
example.

CosMOPHILA SABULIFERA Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

EUTELIA SUBAPICALIS WIk.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. Two specimens. North-west Somaliland, Berbera,
June 5-9, 1897. One example.

PSHUDOPHIA OPPIA, sp. n. (Plate I. fig. 1.)

Male. Head, antenne, and collar white ; thorax and tegule pale
greyish brown; abdomen pale brown, each segment edged with
white ; the underside of the thorax, abdomen, and legs white.
Primaries brown, the outer margin grey ; a reddish-brown band
edged with a metallic line crosses the wing about the middle from
the costal to the inner margin; a short yellowish-white band at
the end of the cell. Secondaries white, broadly bordered with black
from the apex to the anal angle; the fringe black and white.
Expanse | inch.

Hab. E. Somaliland: Joh, in the Haweea Country. Three
specimens captured September 20, 1897. Type and co-type in the
Hope Collection, co-type in the British Museum.

[16]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 19

PsEUDOPHIA LINEATA, sp. n. (Plate I. fig. 7.)

Head, collar, thorax, and tegule pale greyish brown; abdomen
wanting ; antenne whitish ; legs and underside of the thorax white.
Primaries greyish brown, palest at the base and on the costal and
inner margin; a zigzag creamy-white line at the end of the cell
extending to the base; a zigzag yellowish submarginal line ex-
tends from the costal margin near the apex to the anal angle; the
marginal line bluish grey; the fringe dark brown. Secondaries
greyish brown; the outer margin broadly bordered with dark brown;
the fringe alternately brown and white. Expanse 12 inches.

Hab. North Central Somaliland: Haud, Odewein, June 21-23,
1897. Indryriver-bed. One example. Type in Hope Collection,
Oxford.

CEROCALA MUNDA, sp. n. (Plate I. fig. 3.)

The head, tegulz, and thorax pale reddish brown ; abdomen and
legs reddish brown ; antenne and palpi brown. Primaries reddish
grey, with a large reddish-brown spot near the base, one at the end
of the cell and one on the inner margin close to the anal angle ; the
apex dark brown; a submarginal reddish line extends from the
apex to the anal angle; the outer and inner margin greyish; the
fringe white at the apex, the rest dark brown. Secondaries pale
reddish brown; the outer margin spotted with black; a dark
brown spot at the end of the cell; the fringe pale brown.
Expanse 1 inches.

Hab. North Central Somaliland: Haud, Odewein, June 21-23,
1897. In dry river-bed. One example. Type in the Hope Col-
lection, Oxford.

GNAMPTONYX VILIS WIk.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One broken example.

PLECOPTERA REFLEXA Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. Two specimens.

TRIGONODES HYPASIA Cram.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

POLYDESMA OTIOSA Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

CYLLOGRAMMA LATONA Cram.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry

river-bed. Eleven specimens.

Somaliland (1895 or 1897). Four specimens.
Q*

(17]

20 MR. ©. V. A. PEEL AND OTHERS ON [ Jan. 23,

OPHIUSA MELICERTE Cram.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. Two specimens.

Somaliland (1895 or 1897). Four specimens.

SPHINGOMORPHA CHLOREA Cram.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

Somaliland (1895 or 1897). our specimens.

ReEMIGIA REPANDA Fabr.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. Two specimens.

ZETHES HESPERIDOIDES Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

HYPENA ABYSSINIALIS Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. ‘Two specimens.

HYPENA ABYSSINIALIS var. JUSSALIS W1k.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. Three specimens.

SPHINGID A,

DBILEPHILA LIVORNICA Esp.

N. Central Somaliland, Haud District, Eyk, July 3, 1897, In
the open plain, by thick bush. One 92 example.

PHLEGETHONTIUS CONVOLVULI Schauf.

N. Central Somaliland, Eyk, in the Haud District, July 3, 1897.
In the open plain, by thick bush. One 2 example.

GEOMETRID&.
TEPHRINA DISPUTARIA Guen.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

LASIOCAMPID 2.
CHILENA SABRINA, sp. n. (Plate I. fig. 2.)

Male. Head and front of the thorax white ; antenne yellow; the
collar and thorax pale brown ; tegule pale brown edged with white;
abdomen and legs white. Primaries pale brown, with two white
bands extending from the base almost to the outer margin, the
first line straight, the second curved; the costal margin edged with

[18]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 21

white; the fringe white. Secondaries yellowish white. Expanse
17 inches.

Hab. West Somaliland (1895). One example. Type in Hope
Collection, Oxford.

LiMACODID”.

PARASA FULVICORPUS Hmpsn.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

ARBELA QUADRINOTATA WI1k.

N. Central Somaliland, Odewein, June 21-23, 1897. In dry
river-bed. One example.

5. COLEOPTERA.
By C. J. Ganan, M.A., and Ginserr J. Arrow.

With the exception of three species of Curculionide and two or
three other species belonging to genera at persent undetermined,
all the species of Coleoptera of which examples were collected by
Mr. C. V. A. Peel in Somaliland are enumerated in the following
list. Mr. Arrow has contributed that portion which deals with the
Lamellicorn beetles and has assisted also in the determination of
some of the species belonging to other groups. Mrs. M. Kk. Thomas
has been good enough to determine the species of Mylabris and
to describe one new form belonging to that genus. One or two
species have been described from African specimens other than
those collected by Mr. Peel. Full reference to the history of such
specimens will be found in the descriptions. [C. J. Gawan. |

CARABIDZ.

CaLosoMA RUGOSUM De Geer.

Central or East Somaliland. Oneexample. June 5 to Oct. 29,
1897. <A widely distributed species, being found in South and
East Africa, Abyssinia, Nubia, and in the island of Socotra.

ANTHIA FEROX Thoms.

N.W. Somaliland, Galadi in the Mijertain Country. One
example, Oct. 4, 1897. Occurs also in Abyssinia.

PoLYHIRMA CALLIAUDI Casteln., var.

N.W. Somaliland, Galadi, Oct. 4, 1897. One example. The
sutural vitta behind the scutellum is longer, and the elytra some-
what less strongly punctured than in normal examples from
Abyssinia. Type of variety in British Museum.

CHLAENIUS sp.

Central or East Somaliland. One example (1897).
[19]

i)
bo

MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

HYDROPHILID®.
Hi yYDROPHILUS SENEGALENSIS Perch.

Fast Central Somaliland, Sinnadogha in the Haweea Country.
One example, in water-tank, Sept. 8, 1897. This species is
widely distributed throughout Africa.

HisTHRIpDӎ.

HisveR MEMNONIUS Erichs.

Central or East Somaliland. One example (1897). Occurs also
in Senegambia, Nubia, Abyssinia, and Hast Africa.

HistHR GEHINI Mars.

Central or East Somaliland. One example (1897). Found also
in Senegambia, Abyssinia, and East Africa.

SAPRINUS SEMIPUNCTATUS Fabr.

Central or East Somaliland. Six specimens (1897). (One
perhaps 1895 Expedition.)

SAPRINUS CHALCITES II].

Central or East Somaliland. Four specimens (1897). Occurs
also in Egypt, Abyssinia, and East Africa.

DERMESTID A.
DERMESTES VULPINUS Fabr.
Central or East Somaliland. Ten specimens (1897).

SCARABZIDZ®.
Hyposorvs ILLIGERI Reiche.

Somaliland. Three specimens (1895 or 1897). This insect
appears to be of almost world-wide distribution.

PHxocHRovS BECCARIE Har. Coleopt. Hefte, vii. p. 26.

Somaliland. Four specimens (1895 or 1897), apparently
belonging to this species, which was described from N. Abyssinia.

TROX SQUALIDUS Oliv.

Somaliland (1895 or 1897). Two specimens were found of this
insect, which is distributed generally throughout Africa.

Trox pxpansus Arrow, sp. n. (Plate I. fig. 16.)

Breviter ovatus, modice convexus, ferrugineo-tomentosus ; capite
bituberculato ; prothoracis latitudine quam longitudine duplo
majore, lateribus fortiter explanatis, serratis, angulis posticis
fere rectis, disco quadri-carinato, carinis post medium conver-
gentibus ; elytrorum marginibus late explanatis, biseriatim
tuberculiferis, dorso confluenter seriato-tuberculato, interstitiis

[20]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 23

parce granulatis ; antennis ferrugineis, articulo primo nigro,
fusco-hirto ; tibiis anticis acute quadridentatis, alus serratis.
Long. 21 mm., lat. 13 mi.

Type in British Museum.

The single specimen was obtained in Central or Eastern Somali-
land (1897). ‘his species is allied to 7’. denticulatus Oliv., which
also occurs in Somaliland, but it is considerably larger and relatively
broader, the widely dilated margins of the elytra making their
outline almost circular. The elytral costz consist of irregular
tubercles which are separate at the base, more or less confluent
on the dise, and small and scattered towards the apex; the alter-
nate rows only of these reach the basal margins of the elytra.
The anterior tibia is armed with four acute teeth placed at right
angles to its axis and regularly increasing in size towards its
extremity, the last being very sharp and prominent. 7’. funestus
Lansb., a W. African species, appears to be nearly related to this
insect, but is smaller and narrower, 7. evpansus being one of the
largest at present known in the genus.

SCARAB EUS ISIDIS Cast.

North Central Somaliland, Bally Maroli in the Haud District.
Two examples, June 25, 1897. This occurs throughout the
Nile Valley and also in 8.W. Asia.

ScaraBaus Berront Waterh. Ann. & Mag. Nat. Hist. (6) xx.
p- 553 (1897).

One specimen found July 26, 1895, at Whardi Datal, North-
west Somaliland. It was recently discovered by Mr. Betton in
British East Africa.

GYMNOPLEURUS LEVIS Arrow, sp. 0.

G. splendenti valde affinis, sed colore, et sculptura minus evidente,
distinctus, fusco-niger, levis, opacus ; pedibus antennisque fusco-
rufis, harum clavis ferrugineis ; capite subtiliter granulato,
antice acute quadridentato; prothorace convexo, subtiliter
coriaceo, linea media lev angusta, basi bi-impressa, lateribus
parum grosse foveolatis, marginibus lateralibus anguste reflexis
postice obtuse angulatis ; elytris subtilissime disperse granu-
latis et viw striatis, absque plicatulis ; pedum anteriorum tibiis
serratis, dentibus tribus terminalibus acutis et longissimis
munitis, femoribus emarginatis ct dentatis. Long. 16 mm.

Type in British Museum.

One example (1895 or 1897). This insect is almost black
without a trace of metallic lustre. The surface is very smooth,
with a fine granulation which is most apparent on the head and
exceedingly minute on the elytra. The striation of the latter is
almost obsolete and only visible under a lens. The three teeth
upon the anterior tibie are very long and acute.

G. lugens of Fairm. is very near this species, but is more nearly
related to G. splendens Cast., if not merely a variety of it.

[21]

24 Mi. 0. V. A. PEEL AND OTHERS ON (Jan. 23,

Copris HaARRISt Waterh. Ann. & Mag. Nat. Hist. (6) vii.
p. 515 (1891).

Two female specimens of this insect, described from Abyssinia,
were found at Whardi Datal (July 26, 1895).

ONTHOPHAGUS GERSTAECKERI Har.
A single specimen was brought from Central or East Somaliland

(1897).
ONTHOPHAGUS GAZELLA Fabr.

One example (1895 or 1897). This insect occurs throughout
Africa and even extends to Madagascar.

ONTHOPHAGUS sp.

A single female of another small species, probably new (1895
or 1897).

SCHIZONYCHA SQUAMOSA Raffray.

Three specimens (1895 or 1897). This species appears to be
common in Somaliland, having occurred in several collections from
that country.

SCHIZONYCHA NIGROFUSCA Arrow, sp. n.

Ovato-cylindrica, fere nigra, antennis pallidioribus, pectore flavo-
hirto ; clypeo equaliter punctato, margine arcuato vie sinuato
fortiter reflexo, fronte acute bicarinata, vertice spatioque post
carinam anteriorem lavibus ; prothorace antice et postice valde
contracto, grosse et confluenter punctato, punctis squameferis,
lateribus crenatis et piliferis, ante marginem posteriorem utrinque
elevato et levigato ; scutello fere levi punctis nonnullis grossis
sulcaque obsolete mediana ; elytris cum pygidio dense sat grosse
punctatis, punctis squamas setiformes vix perspicuas ferent-
ibus ; tibits anticis tridentatis, dente superiore parvo. Long. 18-
19mm. @.

Type in British Museum, co-type in the Hope Coll., Oxford.

Two specimens, both females (1895 or 1897).

This insect is readily distinguishable from the majority of its
congeners by its dark colour. It has some resemblance to the
common W. African S. crenata, but the prothorax is relatively
much smaller.

Single specimens of two other apparently new species were
found (1895 or 1897), but in view of the large number of closely
related forms described, it seems advisable to leave them for the
present unnamed.

ANOMALA, sp. n.

Two specimens, ¢ and 2, were collected (1895 or 1897), ex-
hibiting a difference in the structure of the claws, and probably
allied to A. transvalensis Arrow. As the anterior tarsi of the male

[22]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 25

are imperfect, the affinity of the species cannot be decided with
certainty, and it will therefore not be described.

There are two more single specimens (1897) of species of this
genus, which can only be properly described from individuals of
both sexes.

PHYLLOPERTHA HORTICOLA Linn.

One example (1897). The distribution of this insect is very
remarkable. It is properlya northern insect, occurring throughout
the North of Europe and Asia from the British Islands to Japan.
In the British Museum, however, are specimens from South
Africa, and Mr. Peel has now brought a specimen from Somaliland.
It seems possible that these African examples have been artificially
introduced.

HomMoruyrRBA HELEN® Schaum.

Two specimens (1897). This species has also been found in
Abyssinia, in East Africa, and at Aden in South Arabia.

RHABDOTIS SOBRINA G. & P.

Fifteen specimens, found by Mr. Peel on mimosa bushes at
Arigumeret, Farfanyer District of Central Somaliland.

BUPRESTID”.

JULODIS LATICOLLIS Gahan, sp. n. (Plate I. fig. 15.)

Viridi-metallica, supra fere glabra, infra subaurata sat dense
cano-pilosa ; capite dense punctato ; prothorace brevi, lato,
supra valde convevo, dense minus fortiter punctato et subopaco,
lateraliter rugoso-punctato, basi utrinque fortiter sinuato, medio
acutangulatim producto ; elytris quam prothorace via latioribus,
dense wrregulariterque foveolatim impressis ; foveolis subauratis,
dense punctatis et plus minusve pubescentibus, interstitiis elevates,
angustis, reticulatis et impunctatis. Long. 18, lat. 8 mm.

Hab. Somaliland (1895 or 1897). One female example.

Type in British Museum.

This species seems to be allied to J. vittipennis Fahr., and
J. subvittata Saund. (Aimblysterna), but is readily distinguished
from either by its broader form, its shorter, broader, and more
convex prothorax, and its differently sculptured elytra.

STERASPIS sp.
Central or East Somaliland (1897). One example.

PsILOPTERA SOMALICA Gahan, sp. n. (Plate I. fig. 10.)

Purpureo-violacea, elytris ( foveis pubescentibus propter 40 cupreatis,
pretermissis) ngris, tarsis supra viridibus ; capite inter oculos
sat lato, dense fortiterque punctato, versus medium sparse, ad
latera dense sat longeque, pubescente ; prothorace quam longiore
sesqui-latiore, antice a medio angustato, supra sat dense forti-

[23

26 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

terque punctato, area parva triangulare ante scutellum levi ;
elytris punctato-striatis, apice acuminatis, interstitiis antice fere
planis, versus apicem angustis et conveais. Long. 25, lat.
9 mm.

Hab, Central or East Somaliland (1897). Oneexample. Type
in British Museum.

Head, prothorax, legs, and underside of a purplish-violet colour ;
tarsi metallic green above. Elytra black, each with about twenty
shallow cupreous pits, which are densely punctured and more
or less completely covered with greyish-white pubescence : six or
seven of the smallest of these pits are placed at irregular intervals
along the third elytral interstice, three of the larger pits on the
fifth, three on the seventh, and about seven, diminishing in size
from before backwards, along the ninth interstice ; one of the larger
pits on the fifth interstice, a little in front of the middle, and one at a
short distance behind the middle, encroach upon the sixth interstice
and nearly join two correspondingly large pits upon the seventh
interstice.

This species differs from all other African species of the genus
known to me in the disposition of the pubescent pits on the elytra,
and is further distinguished by having each of the elytra narrowed
to a single point, and not truncate at the apex.

ELATERID.

AGRYPNUS LONGICORNIS Gahan, sp. n. (Plate I. fig. 11.)

Rufo-brunneus, luteo-pilosulus ; antennis nigris, basin elytrorum
paullo superantibus, articulo 3° quam 2° haud longrore, articulis
47 ad 10%" angulatim sat valde productis ; prothorace quam
latiort via longiort, supra sat valde convexo, lateribus medio
paullo rotundatis, angulis posticis tenuibus, acutis, divaricatis ;
elytris tenuiter punctato-striatis. Long. 17, lat. 5 mm.

Hab. Central or East Somaliland (1897). One example.
Type in British Museum.

In general appearance this species resembles A. bocandei Cand.,
but is easily to be distinguished by the black colour and greater
length of itsantenne. The third joint of the antenne is a little
broader, but not longer, than the second, and the joints from the
fourth to the tenth are each produced antero-distally into a strong
angulate process; the disc of the prothorax is more convex and
somewhat less densely punctured, and the sides are more rounded
in the middle than is the case with A. bocandet.

DASCILLID®.

GENECERUS NEBULOSUS Gahan, sp. n. (Plate I. fig. 9.)

Piceo-brunneus, cinereo- sat dense pubescens; elytris brunneo-
nebulosis, longitudinaliter subcostatis ; antennis (3) flabellatis
vel (9) serratis. Long. 14-16, lat. 4-43 mm.

Hab. Brit. E. Africa: Samburu, Voi and Ndi (C. S. Betton);
[24]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND, 27

Central or East Somaliland (1897), two males (C. V. A. Peel).
Type in the British Museum, co-type in Hope Coll.

Dark brown to reddish brown in colour ; covered with an ashy-
grey pubescence. Elytra with a number of dark-brown blotches,
which are, however, absent in some specimens, so that the elytra
have a nearly uniform grey colour. Head with a Y-shaped glabrous
mark reaching from the antennary condyles to the occiput; eyes
rather large, hemispherical, finely facetted. Prothorax about one
half broader than long; its antero-lateral angles rounded and
obtuse ; the postero-lateral very slightly projecting and acute ; the
disc marked with a faint median impressed line extending from
the anterior margin almost to the base. Elytra with some feebly
raised and obtuse longitudinal cost, along which the grey
pubescence seems somewhat more dense than over the rest of the
surface. Mesosternum with a small projecting process near the
middle of its anterior margin, this process being fringed with
fulvous hairs at its sides and apex. Posterior margin of the fifth
abdominal sternite of the male bisinuate.

This species, though very distinct from Genecerus cervinus Walk.,
appears to be truly congeneric with it, agreeing as it does in all
essential points of structure. The genus Genecerus, stated by
Walker to be allied to Plastocerus, and by subsequent authors
placed in the family Cebriontde, seems to me to belong to the
family Dascillide, in which I should place it near Anorus Lee.

Lycip &.
Lycus amMpiiatus Fahr.

North-west Somaliland, Hargasia, April 25-28, 1895. One
example. ‘This species is found also in East Africa as well as in
Natal and the Cape of Good Hope.

CLERID2.
NeEcropia RUFIPES De Geer.
Central or East Somaliland (1897). Three specimens.

BostRYCHID®.

APATE TEREBRANS Pall.

West Somaliland, Bun Jijjiga, July 15,1895. Three examples.
This species occurs also in West Africa from the Gold Coast
to Angola, in Natal, Nyasaland, and East Africa.

BostRYCHUS sp.

Central or East Somaliland (1897). Two specimens.

TENEBRIONIDA.
ZOPHOSIS AROMATUM Gestro, Ann. Mus. Ciy. Gen. (2) xy. p. 258.

Central or East Somaliland (1897). The two examples obtained
by Mr. Peel are somewhat larger than the type from Archeisa

28 MR. C. V. A. PEEL AND OTHERS ON (Jan. 23,

described by Dr. Gestro, but in other respects fully agree with the
description.

Homata AGona Fairm.
Central or East Somaliland (1897). ‘Two specimens.

RHYTIDONOTA DELICATULA Fairm.
Central or East Somaliland (1897). Two specimens.

Ruyriponora Ropusta Gahan, sp. n. (Plate I. fig. 13.)

Capite supra subtilissime punctato, clypeo utrinque leviter im-
presso ; prothorace quam longitudine paullo latiort, lateribus
marginatis antice arcuatim convergentibus, versus basin minus
fortiter convergentibus, angulis posticis sat latis et retro paullo
productis ; elytris quam prothorace paullo latioribus, bast mar-
ginatis et ad humeros angulatis. Long. 19-20, lat. 73 mm.

Hab. Central or East Somaliland (1897), two specimens ; and

West Somaliland (1895), one specimen. ‘Type (Central or East
Somaliland) in British Museum, co-type in Hope Collection.
Prothorax nearly one-fourth broader than its length ; widest
across the middle, with sides converging strongly towards the
apex, less strongly towards the base ; basal margin straight in the
middle, oblique towards each side and there forming with the
lateral margin a subacute angle slightly projecting backwards.
Elytra widest a little in front of the middle, narrowed slightly
towards the base, and strongly towards the apex; completely
margined at the base, and with a small projection at each of the
humeral angles. Third joint of the antenne twice as long as the
second, and nearly equal in length to the fourth and fifth united.

HIMATISMUS sp.
Central or East Somaliland (1897). One example.

OCNERA sp.
Somaliland (1895 or 1897). One mutilated specimen.

PIMELIA HILDEBRANDTI Har.

P. cenchronota Fairm.

Central or East Somaliland (1897). Four specimens.
PSAMMODES sp.

Somaliland (1895 or 1897). Two specimens.

AMIANTUS sp.
Central or East Somaliland (1897). One mutilated specimen.

AMIANTUS SEXCosTaTUS Gahan, sp. n.

Niger ; prothorace supra valde convexo, dense fortiterque punctato,
lateribus paullo rotundatis, postice leviter marginatis ; elytris
ad suturam haud elevatis, utroque carinis tribus sat valde

[26]

1900.1} INSEOTS AND ARACHNIDS FROM SOMALILAND. 29

elevatis instructo, carinis omnibus fere equilongis, a basi ad
dechivitatem apicalem extensis ; segmentis 2° 3°que abdominis
in medio rufulo-pilosis. Long. 19, lat. (ad med. elytrorum)
103 mm.

Hab, Central or East Somaliland (1897). One example. ‘T'ype
in British Museum,

Black ; prothorax strongly convex above, and strongly and
rather thickly punctured ; its sides slightly rounded in the middle
and feebly marginate posteriorly, the margins becoming obsolete
in front. lytra depressed along the suture, each witb three
well-marked carine reaching from the base to the posterior
declivous portion, the intervals between the carine being rather
strongly concave ; the concave sutural area between the innermost
carine is sparsely granular in its anterior half. Femora rugosely
punctured. Second and third abdominal segments with a patch
of reddish pubescence in the middle.

SEPIDIOSTENUS ERINACEUS Fairm.

North Central Somaliland: Bally Maroli in the Haud District,
June 25, 1897. Nine specimens, captured in the open plain.

SEPIDIUM MAGNUM Gahan, sp.n. (Plate I. fig. 12.)

Pube pallide cervina dense obtectum: prothorace utrinque plaga
subnuda nigra, elytris rugis elevatis glabris nigro-fuscis, et
plagis adspersis cretaceis ; prothorace basi apiceque constricto,
lateraliter carinato, carina antice obtusa, pone medium tuber-
culatim dilatata, disco in medio carinato, antice valde tuber-
culato, tuberculo crasso, apice rotundato supra sulcato-impresso
haud bifurcato ; elytris utrisque longitudinaliter bi-carinatis,
transversim reticulatumque rugosis. Long. 29-35, lat. (ad med.
elytrorum) 12-15 mm.

Hab. Central or East Somaliland (1897). Four examples.
Type and co-type in British Museum, co-types in Hope Collec-
tion.

Brownish black, with the head, prothorax, underside, legs, and
antenne thickly covered with a pale fawn-coloured pubescence.
Prothorax furnished with a large prominent tubercle, directed
obliquely forwards from the anterior part of the disc; this tubercle
is rounded at the extremity, impressed along the middle with a
linear groove, and has on each side a naked and rugose black patch;
from the base of the tubercle a median carina runs along the dise
to the base of the prothorax. On each side of the prothorax there
is a carina, obtuse in front, but more acute and prominent behind
the middle, there forming a flattened tubercle, behind which the
prothorax is abruptly constricted. Elytra each with two longitu-
dinal and more or less sinuous carinz, each of which gives off
short transverse or reticulating ridges on both sides.

This species is one of the largest of the genus, being equal in
size to S. ruspolw Fairm., from which it differs chiefly in havin
the lateral tubercles of the prothorax placed behind the middle

(27]

30 MR, 0. V. A. PEEL AND OTHERS ON (Jan. 23>

and in the presence of short transverse ridges running from the
dorsal carina of each elytron towards the suture.

SEPIDIUM BILOBATUM Gahan, sp.n. (Plate I. fig. 14.)

Pube rufulo-cervina dense obtectum: elytris supra plus minusve
cinereis ; prothorace antice valde tuberculato, tuberculo crasso
untrorsum directo et bilobato ; elytris dense fortiterque punctatis,
utriusque disco tuberculato, tuberculis in seriebus duabus irregu-
laribus ordinatis. Long. 17, lat. 6 mm.

Hab. Somaliland (1895 or 1897). One example. Type in

British Museum.

Closely covered with a reddish-fawn-coloured pubescence, with
the disc of each elytron ashy grey along the middle. Prothorax
with a large tubercle directed almost horizontally forwards from
the anterior part of the disc; this tubercle is distinctly bilobed
in front, each lobe being rounded at the extremity ; each side of
the prothorax carinate, the carina dilated to form a tubercle just
behind the middle. Elytra densely and strongly punctured ; the
disc of each with a number of sharply raised tubercles forming
two irregular, longitudinal series.

SEPIDIUM CRASSICAUDATUM Gestro.
Somaliland (1895 or 1897). Nine examples.

VIELA sp.
Central or East Somaliland (1897). One example.

VIBTOMORPHA FOVEIPENNIS Fairm.
Central or East Somaliland (1897). One example.

MICRANTEREUS sp.
Central or East Somaliland (1897). One male example.

MICRANTEREUS ASIDOIDES Fairm.
Central or Hast Somaliland (1897). One example.

AMARYGMUS sp.
Somaliland (1895 or 1897). One mutilated example.

PRAOGENA NIGRA Gahan, sp. n.

Nigra ; capite prothoraceque crebre punctatis et opacis ; elytris
nitidis, seriatim haud fortiter punctatis, tnterstitis planis, sparse
minutissime punctulatis. Long. 18, lat. 45 mm.

Hab. Somaliland (7. Greenfield and C. V. A. Peel, 1895 or 1897).

Type in British Museum, co-typs .n Hope Collection.

Entirely black, with the elytra somewhat glossy and the
rest of the surface more or less opaque. Head and prothorax
closely punctured, the latter a little broader than long, slightly
rounded at the sides. Elytra each with nine rows of rather small
punctures, the first row (that next the suture) being very short ;

[28]

1900. ] INSECIS AND ARACHNIDS FROM SOMALILAND. 31
intervals between the rows flat and sparsely and very minutely
punctulate.

This species is larger and less convex than P. gagatina Mikl.,
its prothorax is relatively a little broader, the rows of punctures
on its elytra are much finer, and it is distinguished turther from
that species by its darker coloration and its glossy elytra.

CANTHARID®.
MYLABRIS HYPOLACHNA Gestro.
Central or East Somaliland (1897). Two examples.

MYLABRIS SENN” Gestro.
Central or East Somaliland (1897). Three examples

MYLABRIS LATEPLAGIATA Fairm.
Central or East Somaliland (1897). One example.

MYLABRIS SOMALICA Thomas, sp. n.

Black, opaque, rugose, clothed with short yellow pubescence
interspersed with longish black hairs. The head (which has an
elevated shining median longitudinal line), prothorax, and elytra
all thickly and coarsely punctured ; the elytra dilated and widening
towards the apex.

Each elytron ornamented with two long oval basal yellow mark-
ings, the margiual one joining a transverse yellow band extending
to suture, and with a second transverse band, starting from but not
quite touching the suture, extending down margin and curving
round across the apex of elytron back to the suture: these markings
are all margined by a fine brownish-red line. Antenne red, with
the exception of the first and second joints, which are black. The
underside, legs, and tarsi are all black with long yellow hairs ; the
nails and spurs red.

This species is very similar to Mylabris hypolachna of Gestro'
in size, form, and general characteristics, but it differs in having
the second transverse band continued along the outer margin and
thence across the apex to the suture.

Length 12 millim., breadth 4 millim.

Hab. Central or East Somaliland (1897). Type in the British
Museum. [M. K. THomas. |

EprcauvTa AMErHystina Mikl.
Somaliland (1895 or 1897). One example.

CERAMBYCID2A.
MacroroMa PALMATA FE abr.

West Somaliland: South-west Haud, Owari, East of Milmil
(March 16, 1895). One example of this widely distributed African
Prionid.

' Ann, Mus. Genoy, (2) xy. p. 393 (1895.
[29]

32 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

PLOCEDERUS MELANCHOLICUS Gahan, var.

Central or East Somaliland (1897). This variety occurs also in
South Arabia and in British East Africa.

PLoceDERUS PEELI Gahan, Ann. & Mag. Nat. Hist. (7) 11. p. 42
(1898).

Somaliland (1895 or 1897). One example. Co-type in Hope
Collection.

This species has been found in British East Africa as well as in
Somaliland.

PacHyYDISsUS (DEROLUS) SOMALICUS Gahan, sp. n.

Piceo-brunneus, griseo-pubescens ; prothorace supra rugoso, late-
raliter paullo rotundato, area opaca evcisa inter notum pleurum-
que; elytris absque punctis (punctulis minutissimes pubescentiam
gerentibus exceptis); femoribus subtus leviter bicarinatts ; antennis
(2) quam corpore paullo brevioribus. Long. 17, lat. 4 mm.

Hab. Somaliland (1895 or 1897). One example. Type im
British Museum.

Head and prothorax dark brown, with a greyish pubescence.
Head with a suleate impression on the vertex between the upper
lobes of the eyes. Prothorax a little longer than broad; trans-
versely wrinkled above; slightly rounded in the middle on each
side, with a small excised space, bare of pubescence, just below
the pronotum. Elytra parallel-sided throughout the greater part
of their length, rounded at the apex ; clothed with a short closely
laid grey pubescence, and devoid of all punctures, excepting those
very minute ones from which the hairs of the pubescence spring.
Femora feebly carinate along each side near the ventral border.
Prosternal process subvertical behind; the small antero-lateral
processes of metasternum almost completely shutting off the
epimera from the intermediate cotyloid cavities.

PHYLLOCNEMA SEMIJANTHINA Fairm.
Central or East Somaliland (1897). Two examples.

ALPHITOPOLA PEELI Gahan, sp. n.

Fusca ; capite, prothorace, scutello et corporis inferioris lateribus
pube fulvo-ferruginea obtectis ; elytris fuscis, vitta sutural et
vitta postica submarginali connamomers, utroque elytro puncta
basali et maculis quatuor—duabus (quarum interna elongato-ovali,
medio fusco-punctata) ante medium, duabus (elongatis et postice
conjunctis) pone medium—niveis ornato. Long. 17, lat. 53 mm.

Hab. Galadi in N.W. Somaliland. One example, Oct. 4, 1897.
Type in British Museum.

Head and prothorax dark brown, covered with a short reddish-
tawny pubescence. Prothorax transverse; with a small blunt
tubercle at the middle of each side; with two transverse sulcate
impressions near the base and two near the apex. Elytra dark
brown, somewhat thickly punctured, with the punctures rather
large near the base and diminishing in size posteriorly ; a sutural

[30]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND. 33

vitta, and a submarginal vitta on each elytron reaching from the
middle to the apex, where it joins the sutural vitta, cinnamon-
coloured ; a small punctiform spot at the extreme base, two spots
in front of the middle and two behind the middle of each elytron,
snow-white ; the inner spot of the anterior pair larger than the
outer, elongate- oval in shape, and marked in the middle with a

narrow brown spot; the outer spot of the same pair emarginate
in front; the two spots of the posterior pair elongate, shah the
inner one commencing before the outer and coalescing with it
behind. Body underneath with an ashy-grey pubescence along
the middle, fulvous brown towards the sides ; : legs brown, more or
less suffused with grey at the base and on the ventral side. Inter-
coxal process of the mesosternum very feebly tubercled in the
middle. Last abdominal segment feebly and sinuately emarginate
at the apex.

Antenne longer (by the last three or four joints) than the body,
third joint half as long again as the fourth.

CEROPLESIS REVOILI Fairm.

West (April 16 to Aug. 7, 1895) and North-west Somaliland,
Galadi (Oct. 4, 1897). Three ex xamples, two from the latter locality.

CERATITES JASPIDEUS Serv.
Somaliland (1895 or 1897). Ten specimens. This species occurs
also in West and Hast Africa and in Abyssinia.

CALOTHYRZA PAULI (Fairm.).

Anoplostetha pauli Fairm. C. R. Soc. Ent. Belg. 1884, p. 124;
Ann. Soc. Ent. Fr. 1887, p. 338.

West Somaliland, Bularli (May 24, 1895). One specimen.

Tn this species and in the closely allied South-African 4. gardiner
White the claws of the tarsi are divaricate, and the scape of the
antenne is entirely devoid of a cicatrix. Both species are out of
place in Anoplostetha and should be referred to the genus Culothyrza
Thoms., with which they agree in all essential points of structure.
A third African species of ¢ ‘alothyrza has been described by
Dr. Gestro (Ann. Mus. Civ. Gen. (2) xv. p. 423), the remaining
species of this genus being the two Indian forms—C. sehestedi
Fabr. Ent. Syst. Suppl. p. 146 (=C. margaritifera Thoms.) and
C. margaritifera Westw.

CRrossoTUS PLUMICORNIs Nery.

North-west Somaliland, Hargaisa (April 25 to 28, 1895). One
example. This species is found in uae in Hast Africa and
Natal, an example from the last-mentioned locality forming the

type of White’s C. natalensis.

CROSSOTUS sp.

Central or East Somaliland (1897). One somewhat rubbed
female specimen.

Proc. Zoou. Soc.—1900, No. III. 3

[31]

oF MR. C. V. A. PEEL AND OTHERS ON (Jan, 25,

6. NEUROPTERA.
By Rosert McLacunan, F.R.S. &e.

PLANIPENNIA.

PALPARES PAPILIONOIDES Klug, var.

West Somaliland, Sule River, May 29, 1895. One female.

It appears to me to be safer to regard this single specimen as a
variety of P. papilionoides rather than to describe it as new,
Klug’s species was from Arabia Felix. When compared with
Klug’s description and figures, this female is somewhat larger and
the dark bands of the anterior wings are more distinctly fenestrated
in consequence of the dark colour being restricted to margining of
the network. Some examples of P. tristis Hagen, diverge from
the type form in the same manner, and I have a female from
Somaliland that at first I thought was specifically identical with
that from the Sule River, but there is a slightly different form of
wing, and in papilionoides the abdomen has black longitudinal
bands which are wanting in ¢ristis.

PALPARES WALKERI McLachlan.

Sule River, May 24, 1895. One female.

The male of this species was described by me in the Ent.
Monthly Mag. for August 1894 from two examples taken by Mr.
J.J. Walker, R.N., F.L.S., at Aden, which are in my collection.
Subsequently Col. Yerbury, R.A., F.Z.S., found examples of both
sexes at the same place and presented them to the British Museum.
Upon comparing the female from Somaliland with those from Aden,
Isee nothing that can be considered of specific difference. The
female has never been described. It is’ larger (expanse about
120 mm.) and the wings are broader (19 mm.), the isolated black
markings on the anterior wings are larger ; on the posterior wings
the fascie are both broader and longer, the second of them extending
to the dorsal margin, and very broad and strongly angulate in the
middle; the third is also very broad and connected more or less with
a series of spots towards the dorsal margin. (In no two specimens
do the dark bands and other markings precisely agree nor are they
symmetrical on the opposing wings.) ;

MYRMELEON VARIEGATUS Klug.

West Somaliland (1895). One male, without indication of
further locality. Described originally from Arabia Felix. I have
what appears to be exactly the same species from the Sinai Penin-
sula. Probably widespread.

ODONATA.

CACERGATE LEUCOSTICTA Burm.

East Central Somaliland: Haweea Country, Sinnadohga, by a
water-tank, Sept. 8, 1897. One female.
A widespread African insect.

[32]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 30

GYNACANTHA sp. ?

One immature female from the same locality as the preceding.
This in all probability represents a new species, but I am not dis-
posed to describe and name a species of this very critical genus
from an immature female. It appears to be quite distinct from
any West-African species.

7. ORTHOPTERA.
By Matcoim Borg, F.Z.S., F.E.S.
(Except Phasmatodea by Dr. C. BRuNNER VoN WATTENWYL.)
The collection of Orthoptera made by Mr. C. V. A. Peel in
Somaliland is not large, but contains examples of a good number

of genera and species new to science. These are distributed as
follows :—

Nuivber
of species. New Genera. New Species.
Blattodeas 420.5 22. 4 = =
MantodeGs sss 6 ]
Phasmatodea ...... 3 i g
Acridiodea ........ 1155 il 2
Wiocustodeéa’ .% 3 4s <5 1131 1 ff
Gryllodea, ..5.2).2% + - =
Totalleeig.coone 43 4 12

It will be noticed that there are no less than seven novelties
out of the eleven Locustodea captured, a high percentage, which is
probably due to the stationary habits of these insects. Most of
these new species are Phaneropteridw, for there are only one
Conocephalid and one Stenopelmatid, both of which are flightless
forms.

The collection contains no Forficularia.

An account of a collection of Orthoptera made in Somaliland by
the late Prince Ruspoli has been published by Herr Dr. Schultess
Rechberg-Schindler. They are mostly of the same genera, but the
species are different. This is especially noticeable among the new
forms.

I take this opportunity of acknowledging my deep indebtedness
to Herr Dr. C. Brunner von Wattenwyl, for his invaluable assist-
ance in the identification of many of the more difficult and obscure
species, especially for the determination and descriptions of the
Phasmatodea, upon which group his knowledge is unrivalled.

BLATTODEA.
PHYLLODROMIID&.

ISCHNOPTERA ATRA Walk.

Somaliland (April 16 to Aug. 7, 1895, or June 5 to Oct. 29,
1897). One male.
3%
[33]

36 MR. C. V. A, PEEL AND OTHERS ON (Jan. 23,

PERISPH ERID #.
DEROCALYMMA BOTTEGIANA Sauss.

Central or East Somaliland (1897). Two female specimens.

1 cannot distinguish D. analis Sauss. from D. bottegiana Sauss.
The former is merely referred to and included in the synoptical
table, which is very difficult to follow. Mr. Peel’s two examples
appear to agree entirely with the description of D. bottejiana
Sauss., which is described from Somali specimens. One of these
specimens still retains the yellowish velvet coat which is so easily
lost in handling. They are both a shade smaller than de Saussure’s
types.

PERIPLANETID&,

Buavra ORIENTALIS L.

Somaliland (1895 or 1897). One female.

PERIPLANETA AMERICANA (L.).
Somaliland (1895 or 1897). One female.

MANTODEA.
ORvTHODERID”,

CHARIEIS, gen. nov.

Oculi conict, apice pedunculatt. Caput haud valde depressum.
Colore griseo. Ocelli in trianyulum positi ; antennee elongate,
graciles, testacee ; occiput convecum. Pronotum breve, suleulo
wrregulart transverso instructum, margine antico subsinuato,
dilatato, margine postico sainnodato. postice angustatum. Elytra
et ale perfecte explicate. Illa membranabea, remote venosa,
hyalina, venis infuscatis, venulis transversis ramorum vene
radialis incrassatis, bicoloribus ; membrana analis brevis, sed
longior quam ihn Hu hyaline, venulis aree antice infus-
catis. Pedes gr actles ; femora anteca gracilia, spins discoida-
hibus 3 par vis armata, spinis marginis externo irr eqularibus,
prime nunimo, 2° longiori, 3°-6u parvis, dehine margine
crenulato. Tibi antice spinis brevibus armate. Lamina
subgemitalis 2 magna, fornicata, rotundata. Cerct Q sat longi,
laminam subgenitalem haud superantes. Lamina supraanalis
2 brevis, transversa. 9. oS rgnotus.

This genus is a link between Brunner’s two sections of the
Orthoderide. In the form of the pronotum it agrees with Chetessa
Burm., while the presence of small discoidal spines on the anterior
femora would place it in the second division. The conical and
pedunculated eyes, the spines of the anterior tibie, the discoidal
spines, the somewhat long anal membrane of the elytra separate it
from Chetessa Burm.; the form of the pronotum satisfactorily
divides it from the Hremiaphile, Humberticlle, and Chiropache.

In general appearance and coloration it resembles Chiropacha

and 7 ‘arachodes, but it differs in the depressed head.
[34]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND, 37

CHARIEIS PEELI, sp.n. (Plate II. fig. 4.)

Parva, grisea, Elytra hyalina, venulis infuscatis. Ale hysline,
venulis partis antice infuscatis. Pronotum inerme. &.

2.
Wong: COMrporiss —.. sen ae ae 2+ mm.
a) PRONOUN. «c/o eeac ree Sy ramen
3) velytrorum 57.528 ey Sh ee
Vat. max. promot, 4. sees ee 7S
Ay Tanna LMAO B ES Wisi hE 2
Long. fem. anticorum ........ SH)
tiblarum, ee 2-5

Patria. West Somaliland ; North-west Haud, Abriordi Garodi,
May 4, 1895. One female example. Type in Hope Collection,
Oxtord.

(In the figure, the antenne and apex of the abdomen are
wanting.)

CHIROPACHA DIVES Sauss.

West Somaliland: North-west Haud, Abriordi Garodi, May 4,
1895. Three male specimens.

CHIrROPACHA GILVA Charp.

West Somaliland: North-west Haud, Abriordi Garodi, May 4,
1895. ‘Two specimens, male and female,

MAntiIp ®.

Popa unpULATA (Fabr.).

West Somaliland: North-west Haud, Abriordi Garodi, May 4,
1895. ‘I'wo specimens, one male and one nymph.

MiIoManTIs FENESTRATA (Fabr.).

West Somaliland: North-west Haud, Abriordi Garodi, May 4,
1895. One male.

HIERODULA sp.

North-west Somaliland, Berbera, April 16,1895. One mutilated
specimen.

PHASMATODEA.
By C. BrunNER von WATTENWYL.
CLITUMNID&®.
Burrta, gen. noy.’

2. Caput elongatum, Antenne tertiam partem femorum anti-
corum haud attingentes, articulo sccundo moniliformi. Thorax
granulosus, Femora omnia acute carinata, mutica. Segmentum
abdominale octavum longum, segmenta bina terminalia unita.
superans, Segmentum anale parum compressum, minime emar-

1 Tn honorem serutatoris assidui Maleolm Burr.

[35]

ginatum, lamina supraanali minima apposita.
Operculum longissimum, segmenta abdominalia

rect,

tervetes.

MR. C. VY. A. PEEL AND OTHERS ON

[Jan. 23,

Cerci breves,

ria terminalia triplo superans, compressum, acuminatum.

Dispositio specierum.

1. Mesonotum et metanotum irregulariter granulosa.
Stabura Major .......-..:seceeeseceeseeeeeeecereeenneeeoee

- Mesonotum in margine laterali dense granulosum,
medio granulis raris obsitum. Metanotum lve.
Shatlira MIN OF ).22.cc5 eco 5-snese sere scsewecseesceeescss

1. BURRIA LONGIXIPHA, sp. n.

(Plate IT. fig.

longixipha, sp. n.

Jarinosa, sp. 0.

6.)

2. Mesonotum et metanotum irregulariter granulosa.

Long. corporis (operculo excluso) ......

”

2)

33

23

Patria. West Somaliland (1895).

mesonoti
metanoti cum segmento mediano. .
sepmenti mediani 3... =... ..
femorum anticorum .........
- imtermeds 2 cece. -
fs posticornmM. ~--..- = on
opercull cme. Sehege er iesn eens

Collection, Oxford.

2. BuRRIA FARINOSA, sp. n.

2. Caput totum farinosum.
ornatus, necnon apex abdominis farinosus.

One Peale

104 mm.

20
15

3
25

21

24
32

Type in Hope

Thorax vitta mediana longa farinosa
Mesonotum maryin-

ibus dense granulosis, medio in vitta farinosa raro granulosum.

Long. corporis (operculo excluso)

39
3°
93
39
39
39

39

Patria. West Somaliland, Bun Figen (July 15, 1895).
Type in Hope Collection, Oxford.

female.

MCSONOM ose T eee eee ene eet
metanoti cum segmento mediano..
segmenti mediani ..... Becks Hic :
femorum anticorum 3.22.22. --

i Intermed... eee

> POsiicorumMy......j2- 2) Ae
Operculitteees sera ee ene rleneee .

LEPTYNIA sp.

West Somaliland, Bun Jijjiga (July 15, 1895).

specimens.

ACRIDIODEA.

TRUXALID®.

TRUXALIS NasuTA (L.).

North-west Somaliland, Whardi Datal, July 26, 1895,

females.
[36]

Ds
68 mm.
1
Tal
2
19°5
14:5
18
26

One

Five mutilated

Tl.ree

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 39

TRUXALIS UNGUICULATA Ramb.

North-west Somaliland, Whardi Datal, July 26,1895. Seven
females, one male.

MACH#RIDIA BILINEATA Stal.
North-west Somaliland, Whardi Datal, July 26,1895. I refer
a mutilated larva to this species.

EPacRoMIA THALASSINA (Fabr.).
West Somaliland (1895). One example.

CEDIPODID &.

HUMBERTIELLA TENUICORNIS (Schaum).

West Somaliland: Boholo Deno, near R. Shebeyli, June 24, 1895 ;
three male specimens, one female. Somaliland (1895 or 1897);
one male and one female.

This species is purely a native of tropical Africa.

GASTRIMARGUS VERTICALIS Sauss.

West Somaliland (1895). One male and one female specimen.

Var., fusca, fascia fusca alarum viv persprcud.

North-west Somaliland, Hargaisa, April 25-28, 1895. One
female specimen.

CEDALEUS INSTILLATUS, sp. n.

Pronotum haud valde elongatum, fusco-testaceum, indisiinete fusco-
notatunr. Tempora acuta, trigonalia. Statura minore.
Gracilior. Frons subreclinata ; costa frontalis latwr, margin-
ibus a vertice subidivergentibus ; vertex ad frontem haud rotun-
datus. Pronoti crista humilis, acuta, sinuata, margine postico
acutangulo. Elytra venis spuriis completis, area ulnare w-
regulariter reticulata, fusco-testacea, dimidia basalt fascits
3 latis fuscis, 2 latis pallidis, opacis, dimidia apwali plus
minus hyalina, maculis nebulosis infuscatis ornata, Ale bast
flavicantes, fuscia fusca lata semilunari, marginem posticum
attingenti ornate, ultra hance hyaline, apice macula fusca, 2

lobos includenti ornata, Pronotum metazona quam prozona haud

brevior. Femora postica testacea, extus punctulata, vel wrregu-
lariter fasciata. Tibiw postice rufescentes. SQ.

che fone
Long. corporis ....... Peel) hii 29 nm.
Pamie EONOUMed Sac. .0sie Sevas 12 oko 6
eee ClviGronulnivy te se eee ae 26°5
», femorum posticorum .. 15

Patria, West Somaliland (1895). Twospecimens. Types, male
and female, in Hope Collection, Oxford.

This new species falls into the second division of de Saussure’s
subgenus @daleus sensu stricto. It is u great deal smaller than

[37]

40 MR. C. V. A, PEEL AND OTHERS ON [ Jan. 2é

CE. nigrofasciatus de G., and @. infernalis Sauss., and falls between
. senegalensis Kr. and (@, abruptus Sauss. From both of these
it differs in the acute angle of the posterior border of the pronotum,
at least in the male (this angle is more obtuse in the female).
I have drawn up the description on the model of de Saussure’s
synoptical table, in which he distinguishes these two species.

ACROTYLUS LONGIPES (Charp.).

Somaliland (1895 or 1897), one female. West Somaliland
(1895), one male and two females.

These specimens are the red variety, the same as recorded by
me from Sokotra (P. Z. 8. 1898, p. 384).

PYRGOMORPHID #&.

CHROTOGONUS LUGUBRIS Blanch.
Somaliland (1895 or 1897). One male, two larve.

PHYMA'TEUS STOLLII Sauss.

North-west Somaliland, Whardi Datal, July 26, 1895. Two
males and two females.

PH. MORBILLOSUS Nerv. ?

eae ae Somaliland: Hand, Odewein, June 23,1897. Two
larve. (‘ Dry river, thickly w ooded banks.” —C. V. A. es)
il: ane these leer with some hesitation to P. morbillosus.

PYRGOMORPHA GRYLLOIDES (Latr.).

North-west Somaliland: Whardi Datal, July 26, 1895. One
female.

ACRIDIIDE.
ScHISTOCERCA PEREGRINA (Oliv.).

North-west Somaliland: Gooban District, between Hargaisa and
Berbera, August 6, 1895. Six males, one female.

North Central Somaliland : Haud, Odewein, July 23, 1897. Four
nymphs. (“ Dry river, thickly wooded banks.”—C. V. A. P.) Also
five very young larve.

ACRIDIUM SUCCINCTUM Serv.
(nee Linn. = ruficorne, Serv. et Stal).
West Somaliland (1895). One male and one female.

SAURACRIS, gen. nov.

Corpus apterum, depressum, granulatum, witidum. Ocul pro-
minuli, a supero visi, subcontigui. Frons reclinata ; antenne
filiformes. Pronotum deplanatum, carinis nullis, suleis 3
instructum, maryine postico subsinuato, lobis defleais marginibus
antico et postico obliquis, inferiore subrecto. Elytra minima,

[38]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND, 41

rudimentaria vel nulla, Ale nulle, Abdomen segmentis 1° et 2°
deplanatis, ceteris compressis, carinulatis. Pedes crassiuscult.
Femora postica valde incrassata, haud serrulata ; tibie postice
margine externo 6—S , interno 8-9 spinis armate ; tarst postice
segmentis 1° et 3° elongatis, 2° parvo. Cerci g breves, compress,
conici, apice obtusi. Lamina supraanalis 3 obtusa, apice
paullo bi-emarginata, medio plus minus sulcata. Lamina
subgenitalis 3 mayna, fornicata; lamina subgenitalis 9?
simplex. Valvule ovipositoris breves, yranulate. Corpus
glabrum, partibus genitalibus 3 Q tibits posticis hirsutis,
This new genus falls into the group Coptacre. The absence of
elytra, wings, and of the carinz of the pronotum separate it from
Cyphocerastes Karsch, Epistaurus Bol., Acridoderes Bol.

SAURACRIS LACERTA, sp.n. (Plate I. fig. 3, 3.)

Statura mediocri, testacea, nigro varia, Caput pallide testaceum ;
oculi globosi, badii. Pronotum pallide testaceum, fasciis 3
Susctoribus longitudinalibus ornatum. Elytr al minutissima,
granulati, lobiformia, segmenti abdominalis primi dimidium vi
attingentia, vel nulla. Abdomen testaceum, vittis et maculis
nigris ornatum. Pedes testacei ; femora postica extus fusco vel
atro ornata ; tibie postice spinis extus 6-8, intus 9 armate,
Abdomen subtus nigro variegatum. Femina mare major,

juscor. 36 &.

os Oe
hong COrporis' «42 seyoce oe 255mm. 389 mm.
oe CONROE cs. Cee ee 6 8
» elytrorum (si adsunt). . are) ‘75
PP femorum posticorum .. 13-13°5 17

Patria. North-west Somaliland, Hargaisa, April 25-28, 1896 ;
1 male, 1 female and 1] nymph. Central or East Somaliland (1897) ;
2males. Female, type trom Hargaisa, in Hope Collection, Oxford.
Male type from Central or East Somaliland, in Hope Collection.
‘o-types at Oxford and in the collection of Malcolm Burr.

LOCUSTODEA.

PHANEROPTERID ®,
PrRONURA sp. ine.

North-west Somaliland, Whardi Datal, July 26, 1895. One
female nymph.

It is impossible accurately to determine this immature specimen.

PHANEROPTERA PUNCLULATA, sp. 0.

Parva, lete viridis ; pronoto, cruribus, abdomine nigro-punctulatis.
Pronotum lobis deflexis wque altis ae longis 3 carine haud
valde expresse, flavide. Elytra breviora, venulis transversis
haud prominulis, parte anali usque ad apicem nigro-punctulata.
Ale elytris longiores. Femora postica gracilia, apicem elytrorum

[39]

42

MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

superantes, basi incrassata. Lamina supraanalis 3 oblonga.
Cerci 3S longi, teretes, acuminati, apice mucronati. Lamina
subgenitalis 3 elongata, trianc gular iter emarginata, cercos hawd
superans. .
3.
Wong Bpronotiaasectyaseio uae -.-1tenele oman

COFPONISH news Jrcrcterata es
ALAM eat eiaisenecie see toes

yu ClyYCrOrlumiss . ae eeeeee re
Lat. elytrorum medio ..........
Long. femorum posticarum......

9

”

me ho

rE
~T -1 ~]
Or Oi Or

e

Patria. North-west Somaliland, Whardi Datal, July 26, 1895.
One male. Type in Hope Collection, Oxford.

This species comes nearest to Ph. nana Charp., but may be
distinguished by the absence of the basal spots of the elytra, which
also separates it from Ph. quadripunctata Br. The minute black
dots all over the body resemble those of Leptophyes punctatissima,
Bose.

MILIvITSA, gen. nov.

In tribum Terpnistriarum locandum. ¢ ignotus.
Fastigium verticis sulcatum et compressum. Frons brevis, per-

pendicularis. Oculi valde globosi. Pronotum selleforme, disco
antice elevatum, obtusum, postice deplanato, margine ipso sub-
elevato, rotundato, carinis lateralibus nullis; lobis deflexis
altioribus quam longioribus, marginibus rotundatis. Elytra
angusta, paullo ante medium latiora, dehine attenuata, margine
postico sinuato, apice oblique truncata ; vene 2 radiales sub-
contiguee, apicem versus divergentes, ramis 1-3 vene ulnaris
conjunctis, ramo ultimo in marginem cprenlem elytri ecurrente ;
vena ulnaris a vena radiali valde remota, in marginem posticum
elytri defleca, ramos nonnullos furcatos gee circumdatos venam
radialem versus emittens, quorum postremi ramis vene
radiahs conjuncti ; campus analis basi latus, dehine angustis-
simus, venulis rectis transversis plurimis instructis, in tertia
parte apicali elytri evanescens ; campus marginalis latus,
ad conjunctionem vene radialis anterioris cum maryine antico
productus ; imtra marginem per totam longitudinem elytra
anguste decolor, hyalinus, Ale elytris longiores. Coww antice
spina nulla instructe. Femora omnia teretes, postica subtus
spnulis 3. armata, lobis genicularibus acuminatis ; tibie antice
et imtermedie compress, margine postico spinulis minimis
armate. Tibie antice utringue foramine aperto, oblongo,
instructe ; tibiw postice per totam longitudinem utrinque supra
spines albidis, apice rufescentibus, armate. Meso- et meta-
sternum lobis obtusis instructa. Ovipositor satis magnus, sensim
incurvis, apice obtusus, valvulu superiore minime crenulato,
valvulam inferiorem muticam superans. 2.

This genus may be at once separated from Verpnistria St&l by
the unspined pronotum and unlobed legs. The unarmed cox:

[40]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 43

and open tympana of the anterior tibie distinguish it from
Diogena Br.

MILITITSA SOMALIENSIS, sp. n. (Plate II. figs. 5 & 7.)

Corpus ferrugineum, Caput et pronotuin viridia, Elytra viridia,
venulis transversis hyalino-circumdatis, imargine antico hyalino,
margine postico sordide ferrugineo, Ovipositor nitidus, viridis,
apice infuscatus, Ceret recti, conict. Lamina subgenitalis 2
parva, apice angusta, truncata. 2.

Q.
Wong? corperis. «.)2%/s-5..2c sheets eee ee ee 20°5 mm.
55 pronoti... ..). Soa. ake mc ie to eere color
pan OLY bROTEMNS st tyrone Bb PLE nn ee 27
», femorum posticorum ....... nips ae = Pate
sot LOVIDOSIUOIIS: © -cchieus cts gers Oy eicieretati sen 5, ld

Patria. North-west Somaliland, Whardi Datal, July 26,
1895. One female specimen. Type in Hope Collection, Oxford.

TYLOPSIS PERPULCHRA, Sp. Nn.

Viridis, elongata, Pronotum dorso deplanatum, viride, minutis-
sume fusco-punctulatum, marginibus antico et postio subsinuatis,
carinis lateralibus rectis, parallelis, albis, antice subtus atro-
marginals ; lobi deflewi angulato wmserti, margine inferiore
recto, angulo postico retroproducto, margine postico valde
simuato, antico perpendiculari, Elytra elongata, angusta,
lanceolata, ramo primo vene radialis apice furceto, viridia,
antice late albo-marginata, vitta pallide purpurea intra, apicem
versus attenuata ornata. Ale elytris valde longiores, venis
radialibus meds valde infuscatis, (Pedes antict et intermedia
desunt.) Pedes postici gracillimi, longissimi, fusco-testacei.
Cerci longi, validi, apice valde undulato-incurvi, apice
decussati et infuscatt. Lamina subgenitalis 3 elongata, apice
triangulariter evcisa. S.

3
Hones CORPOFIS v.10 tape aes Peace, 2d varie
Se) PROMOUI Sareea ae eset ay ean ee)
Sra CLV UROLFUM sce ts. cst no at ae ase

» femorum posticorum........ 28

Patria. North-west Sowaliland, Whardi Datal, July 26,
1895. One male specimen. Type in Hope Collection, Oxford.

This species agrees with 7’. bilineolata Serv. in the straight
inferior borders of the side flaps of the pronotum, but may be
distinguished by the variegated elytra.

OTIAPHYSA ANGUSTIPENNIS, sp. n. (Plate II. fig. 2.)
Flavo-ferruginea ; elytra, alarum apices, virides ; elytra venulis
transversis albido-circumdatis, margine antico valde sinuato,
margine postico recto, paullo ante medium latiora, pone medium
valde angustata, apice oblique truncata. @.
[41]

44 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

Wong. corporis 0. =e 5 eee eee ee OnE,
son VEDEOHODE o/s gry aseretere te tears ainaer )
ee WUEORUMUS eetatereiret err 40
Lat. max. elytrorum ante medinm Bretes 2

a as prone medium.... 6
Tiong. OVIPOSIbOLIs —..0. 2 oe O75

Patria. North-west Somaliland, Whardi Datal, July 26,
1895. One female specimen. Type in Hope Collection, Oxford.

This species differs from O. hebetata Karsch in its smaller size,
narrowed elytra, and slightly longer ovipositor. In 0. hebetata the
elytra attain their greatest breadth (12 mm.) just before the apex ;
in this species, in the first half of the elytra.

RHEGMATOPODA PEEL, sp. un. (Plate II. fig. 1.)

Pallide virescens. Pronotum disco infuscato, Elytra translucida,
longiora. Ale elytris longiores. Pedes graciles ; femora antica et
intermedia subtus utrinque spinulosa; tibie antice et intermedie
triseriatim spinulose. (Femora postica desunt.) Elytra vene
radiales basi remote, dehine appropinquuntes, subcontigue,
deinde divergentes, vend postica ramum furcatum in apicem
elytra emittens ; vena ulnaris sinuata; area analis mages
explicata ; venule transverse omnes recte, parallele ; campus
tympanalis valde prominulus. Lamina supraanalis 3 brevis,
rotundata. Cerct $ laminam subgenitalem multo breviores,
acuminati, mucronati. Lamina subgenitalis 3 elongata,
deplanata, apice triangulariter emarginata, lobis obtusis. ¢.

Q agnota.

(oii
Tieng. corporis,...2.;..-- £2) min.
Spe PLOMObl ner. ones UO

5 elytrorum’.)...... 34

Patria. North-west Somaliland, Whardi Datal, July 26, 1895.
One male specimen. Type in Hope Collection.

Tabula specierum generis Rhegmatopode.

1. Elytra campo marginali sensim angustato, tertia parte

apizali ev anescenti, vena ulnari decurva ; area ulnaris

(pone venam ulnarem) sat lata. Pronotum lobis

deflexis, margine postico vix sinuato, margine infero

conyvexo. Femora MUEICA “223i... -seossesenesitecescceseer 1. leptocerca Stal.
2. Elytra campo marginali latiori, apicem See attin-

genti; vena ulnari sinuata, undulata, apice recurya,

area ulnari latissima. Pronotum lebis deflexis,

margine postico valde sinuato, margine infero

obliquo. Femora auntica et intermedia subtus

SPEC GT SB ceregacgsta bone bon GRbcgs SASSO SDOKOO ECOG CAG oOEE 2. peeli, n. sp.

[42

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 45

EurycorypHa VARIA Brunner.

Patria. North-west Somaliland, Whardi Datal, July 26, 1895.
One female. Recorded from Kilimanjaro by Brunner.

CoNOCEPHALID &.
XIPHIDIUM SOMALI, sp. n.

X. natalensi Redt. vicinum. Fastigium verticis ab antico visum
lateribus haud distincte divergentibus. Caput et pronotum flava,
fasciis tribus, media lata, lateralibus angustis fuscis ornata ;
pronotum inerme, lobis deflevis obtuse triangularibus, marginibus
obliquis, viv sinuatis, callo convexo nullo. Elytra abbremiata,
lohiformia, pronoto breviora, testacea, margine antico infuscato.
Ale. obliterate. Abdomen fusco-testaceum, utrinque flavido-
bilineatum. Femora postica inermia, genibus inermibus. Pedes
testaceis. Ovipositor longissimus, rectissimus, Lamina sub-
geutalis 2 apie truncata. 2. J tynotus.

Wong corporisiy. sae aas se ieee) Lonmin.
+» pronoti 5)
PL NAA KGIADIIN Bi Om Rion o me
5 OV IPOSILOB Serene! aerate 8

Patria. North-west Somaliland, Whardi Datal, July 26, 1895.
One female specimen. Type in Hope Collection, Oxford.

This species differs from X. natalense Redt. in its unarmed
knees and differently coloured head and pronotum. From
X. brevicercus and X. armaticeps Karsch it may be distinguished
by its unarmed head and pronotum.

eel te (eel ie) eo) 8) Leleliele nse

Herropip#.
EUGASTER LORICATUS Gerst.
North Central Somaliland: Haud District, Eyk (“in open plain
by thick bush”), July 2-4, 1897. One male and one female
specimen.

SPALACOMIMUS TALPA (Gerst.).

North Central Somaliland: Haud District, Eyk, July 2-4,
1897. (‘Open plain by thick bush.”—C. V.A.P.) One male and
one female specimen.

SrENOPELMATID #.
MAGRETTIA OBSCURA, sp. n.

Fusco-testacea, nitida. A M. abominata Br. differt femoribus
tantum spinula minima unica apicem versus armatis, tibus
anticis inflatis. A M. mutica Br. differt femoribus spinula
united, neque 4-5 armatis, pronoto antice quam postice paullo
latius. .

[43]

46 MR. C. V. A, PEEL AND OTHERS ON [Jan. 23,

Gia
Wong corporis. ieee 22 mm,
Sj DLOMOULUbcesvaisenvor ore ry)
pro Les pOStay me ieee 16°5

Patria. North Central Somaliland: Haud District, Eyk (“ on
open plain by thick bush”), July 2-4,1897. One male specimen.
Type in Hope Collection, Oxford.

This species seems to be a link between M. abominata Br.
(African), which has many-spined femora and compressed anterior
tibie, and the pronotum slightly broader anteriorly than posteriorly,
and M. mutica Br. (Asiatic), in which the femora have only 4-5
minute spines beneath, while the anterior tibia are sub-inflated
and the pronotum is cylindrical.

GRYLLODEA.
GRYLLID 2.

LIOGRYLLUS BIMACULATUS (De Geer).

North-west Somaliland, Hargaisa, April 25-28, 1895. Four
female specimens.

GRYLLUS sp.?

North-west Somaliland, Hargaisa, April 25-28, 1897. One
female, one nymph. Both specimens are too mutilated for
determination. The elytra are abbreviated, and the ovipositor is
as long as the body. The general colour is dull black.

GRYLLUS MELANOCEPHALUS Serv.

North-west Somaliland, Hargaisa, April 25-28, 1895. One
male specimen. Possibly only a dark variety of G'. domesticus.

GRYLLUS LUGUBRIS Stal (=G. afer var. ?).

North-west Somaliland, Hargaisa, April 25-28, 1895. One
male specimen.

8. INSECTS OF OTHER ORDERS.

By various Contributors.

{ Mr. Peel also brought several Hymenoptera and a few Hemiptera.
The known species of the former have been kindly identified by
Mr. W. F. Kirby, of the latter by Mr. W. L. Distant.

E. B. Pourron. |

HYMENOPTERA.
Terebrantia. Entomophaga,

BRACONID®.

Bracon sp. A single specimen (1897).
[44]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND, 47

Aculeata. Heterogyna,
ForMIctp&.
PALTOTHYREUS PESTILENTIUS (Smith).

Three specimens from Bularli (West Somaliland), May 25, 1895.
Mr. Peel describes the species as excessively abundaut at this
locality.

Fossores.
Muviniipé&.
MoctTILUA ARENARIA (Fabr.).
A single female specimen (1897).

PoMPILID&.
SALIUS MEGAERA (Smith).
A single specimen from the Webbi Shebeyli, near Mount Kul-
dush, June 28, 1895,
LARRIDE.
LARRADA sp. allied to Z. diabolica (Smith).
A single specimen from Sibi (West Somaliland), May 27, 1895,

Diploptera.
EUMENID&.

EUMENES LEPELLETIERI (Sauss.).

Seven specimens from Eyk, in the Haud District of North
Central Somaliland, July 2-4,1897. ‘ Open plain by thick bush ”
(Cav. ALP),

EUMENES DIMIDIATIPENNIS (Sauss.).
A single example from West Somaliland (1895).

RHYNCHIUM sp.

Two specimens from the Bun Feroli, north of the Webbi She-
beyli, June 10-20, 1895.

Anthophila.

APID#

XyYLOCOPA sp. near to flavilabris.

A single example (1897).

APIS LIGUSTICA (Spin.

Eight specimens from the Haweea Country in East Central

Somaliland, Sept. 8-27, 1897. CW EK)
[45]

48 MR. ©. V. A, PEEL AND OTHERS ON [Jan. 25,

HEMIPTERA.

Two species of Rynchota Hemiptera were obtained by Mr. Peel.

ODONTOPUS SEXPUNCTATUS (Lap.).

One specimen from Bularli in West Somaliland, May 25, 1895 ;
and others from the 1897 Expedition in Central and Hast Somali-
land (no further locality).

Two specimens of another species in bad condition were also
obtained in 1897, (W. L. D.)

9. CHILOPODA AND ARACHNIDA.
By R. I. Pocock.

The Arachnida collected by Mr. Peel proved on examination to
be exceptionally interesting. Of the six species of Acari, two of
the parasitic species of the genus Llipicephalus seem to be well-
marked new forms. One of them is represented by both males
and undistended females; the other unfortunately by a single
male, but this specimen differs so strikingly in colour from all
the species of the genus recently recorded in Neumann’s useful
monograph of the group, that 1 have not hesitated to describe
it as new. Mr. Peel was even more fortunate with his Scorpions.
He collected only five specimens ; but they represent four species,
three of which have never been previously described. Two of
these, Buthus calviceps and Pandinus pugilator, are exceptionally
well-marked forms; while the third species, which L have dedicated
to Mr. Peel, belongs to a section of the genus peculiar to Somali-
land and hitherto ‘represented by a single species.

The working out of this material has entailed a revision of
the Scorpions of Somaliland, based upon those contained in the
collection of the British Museum. The results have been incor-
porated in a supplement to the report upon Mr. Peel's collection,
in the hope that sportsmen and naturalists who visit this country
may see at a glance what is known of these animals and may be
induced to follow, so far as collecting is concerned, the examples
of Messrs. Lort Phillips, Donaldson Smith, and Peel,

Class CHILOPODA.
Family SCOLOPENDRID®.

Two representatives of this family were obtained, April 25-28,
1895, at Hargaisa, North-west Somaliland, namely Pihiosnorens
trigonopoda, a species which is distributed throughout. tropical
Atrica, and a damaged example of a species of 7th ysida probably
referable to R. paucidens, Pocock’, originally procured at Loga in
the Arnec Galla country, but the absence of the anal legs makes
the determination doubtful.

The specimens of these species are in the Hope Museum at
Oxford.

1 Tn Donaldson Smith’s ‘Through Unknown African Countries,’ p. 404 (1897).

[46]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND. 49

Class ARACHNIDA.
Order ACARI.

Family TRomBIpIID #.
Genus TROMBIDIUM.

TROMBIDIUM TINCTORIUM (Linn.), Trouess.

Trombidium tinctorium, Trouessart, Ann. Soc. Ent. France, 1894,
pp. 89-91 & 94, fig. a.

Loc. Hargaisa in North-west Somaliland (April 25-28, 1895).
Two specimens in the Hope Museum at Oxford.

Family ArGasip2.
Genus OrniTHODOROS Koch.

ORNITHODOROS SAVIGNYI (Aud.).

Argas savignyi, Aud., Description de l’Egypte, Hist. Nat. i.
pl. iv. fig. 5; Explanation of Plates, p. 183 (1827).

Ornithodoros savignyi, Neumann, Mém. Soc. Zool. France, x.
p. 26 (1897).

Loc. Bularli in West Somaliland (May 1895). Six specimens,
two in the British Museum, the rest in the Hope Museum at
Oxford.

Family [xop1p#,
Genus Hyatomma C. Koch.

HyatomMa G@Rossum C. Koch.

Hyalomma grossum, C. Koch, Arch. Natur. x. i. p. 220, no. 2,
Uebersicht etc. iv. p. 34, pl. ii. fig. 8 (1847).

Loc. Bularli in West Somaliland (May 1895).

A single distended female, probably referable to this species ; in
the Hope Museum at Oxford.

Genus RuipicerHatus C. Koch.

RHIPICEPHALUS SANGUINEUS (Latr.).

Ixodes sanguineus, Latreille, Hist. Nat. Crust. Ins. i. p. 157
(1804),

Lthipwephalus sanguincus (Latr.), Neumann, Mém. Soc. Zool.
France, x. p. 385 (1897).

Loe. Bularli in West Somaliland (May 1895).

Three specimens, one in the British Museum and two in the
Hope Museum at Oxford.

Proc, Zoot. Soc.—1900, No. LY. 4
[47]

50 MR. C, V. A. PEEL AND OTHERS ON [Jan. 23,

RHIPICEPHALUS MARMOREUS, sp. n. (Plate IIL. figs. 1-1 d.)

3g. Colour. Capitulum yellowish brown; dorsal surface of body
yellowish white, with five large deep red-brown patches, one on
each side extending backwards from the cervical groove, one on
each side longitudinal and sending off towards the middle line a
broad transverse bar, and a large posterior median patch with
convex hinder border and anterior border produced forwards in the
middle line, also a narrow brown band running round the margin
of the dorsal scute; legs yellowish brown, with a broad longi-
tudinal white band on the upperside of the femora, patelle, and
tibie; ventral surface of body whitish, with chitinous sclerites
deep brown like the legs.

Posterior border of capitulum evenly concave, the angles
moderately produced; lateral margins moderately, not strongly
diverging, the anterior border only a little wider than the posterior.

Dorsal plate almost entirely covering the sides of the body,
leaving merely a narrow marginal membranous rim; cervical
grooves distinct, short ; marginal groove represented by a series
of coarse punctures ; three posterior punctured grooves on the
posterior dark patch; for the rest the dorsal plate is without
grooves but is pitted with coarse scattered punctures; the posterior
rim divided by short sulci into eleven festoons. Ventral area with
a single long piriform adanal plate, which is narrowed in front
and extends backwards almost to the posterior cox; its external
border slightly convex, internal border concave in the middle,
convex in the anterior and posterior third, posterior border oblique
and ending internally in a rounded rectangular prominence; a
single small median ‘sclerite on the postanal membranous area.
Ooxe of anterior legs produced into two strong spiniform pro-
cesses ; of 2nd to 4th with a neo e distal spiniform process on the
posterior side.’

Total length of dorsal plate 3°3 mm.

Loc. Bularli in West Somaliland (May 1895).

A single specimen ( ¢ type) in British Museum.

RHIPICEPHALUS ARMATUS, sp.n. (Plate III. figs. 2-2 f.)

3. Colour a tolerably uniform deep brown, with black markings
on the grooves of the dorsal plate; legs deep brown ; ventral area
yale.

Capitulum with posterior border straight in the middle, its
angles more abruptly spiniform than in &. marmoreus, its lateral
margins more obliquely diverging in their posterior two-thirds,
then abruptly converging. Dorsal scute not quite covering the
lateral area, with normal cervical grooves ; marginal groove ‘deep,
strongly pitted, extending from a point on a level with the cervical
groove to the beginning of the festoons of the posterior border ;
posteriorly there is a pair of deep pitted grooves, and between

' Owing to the dried state of the specimen, satisfactory examination of the
mandibular armature was impossible,

[48]

1900. | INSECTS AND ARACHNIDS FROM SOMALILAND, 51

them and the marginal eueoe on each side another similar but
eurved groove which extends from in front of the middle of the
dorsal area back as far as the posterior end of the marginal groove;
in addition to the punctures in the grooves there are a few coarse
punctures scattered here and there ; 5 posterior border with eleven
festoons. Adanal plates narrowed in front, but not extending far
forwards beyond the anus; their inner neatlee nearly straight,
external border convex ; posterior border produced into a short
external and a long spiniform internal tooth, which bees
nearly as far back as the posterior border of the dorsal plate, the
apex of this spine is obliquely truncate and subbifid, above it is a
second strong spine; a pair of postanal sclerites. Cove of 1st
leg strongly bidentate; of 2nd to 4th also bidentate, the two teeth
being on ‘the posterior border, one proximal, the other distal and
thinner. Total length 4:8 mm.

Two undistended female specimens, probably referable to this
species, have the cephalic plate about as wide as long, reddish in
the middle, blackish at ihe sides, the cervical grooves “deep, a row
of large punctures representing the marginal grooves on each side,
some large punctures along the lateral edge i in front of the eye, a
few between the cervical grooves, and a few large ones and many
smaller ones on the middle of the posterior area. The abdomen is
impressed posteriorly with three shallow grooves and is obsoletely
festooned. Cow of the legs posteriorly weakly bispinate, the
external spine being the larger.

Loc. Bularli in West Somaliland (May 1895).

Six specimens. Type (male) and two co-types (male and
female) in the British Museum. Three co-types (2 males and
1 female) in the Hope Museum at Oxford.

The inales of the three species of this genus obtained by
Mr. Peel may be determined as follows :—

(dg) a. Dorsal plate marked behind with a pair of moderately
long grooves, one on each side of the middle line;
between these and the deep marginal groove there is a
third long and deep groove extending from the second
or third suleus of the festooned border past the middle
of the dorsal plate; adanal plate produced behind into
a long avd strong spiniform process, immediately above
which there is a second strong spine; a pair of chiti-
MOUSHPOStanalyscleritesmunerececeeeeescacacce se sore ceneateees armatis.
6. Dorsal plate with a longish posterior median suleus and
a pair of shallower impressions, one on each side of it,
with only a very short shallow and inconspicuous groove
between the marginal groove and the middle line ; adanal
plate not produced behind into a long spiniform process ;
no spine above the termination of this plate.
a’. Dorsal plate yellowish white, with large symmetrically
disposed reddish-brown patches ; legs brown, painted
with white above; dorsal plate sparsely punctured ;
marginal groove represented by a series of punctures. marmoreus.
5'. Dorsal plate deep brown with blacker patches and
yellow lateral margin ; legs uniformly brown; dorsal
plate closely and irregularly punctured; marginal
STOOVE CAISUIMCHE savecsceseessascmcasceeacices souonddosaccosanac sanguUineus.

[49]

52 MR. 0, V. A. PEEL AND OTHERS ON [Jan. 23,

Order ARANE &.

Unfortunately the few Spiders obtained by Mr. Peel were not
all preserved in such a manner as to make their specific deter-
mination possible, being dried, pinned, and for the most part very
much shrivelled. The following forms, however, are recognizable.

1. ARANEUS HOPLOPHALLUS Poc. (Bull. Liverpool Museum, 1.
p. 40, 1899.)

An adult male certainly belonging to this species and a female
doubtfully referred to it, the former from Berbera in North-west
Somaliland, the latter from East Central Somaliland (1897). The
type was procured in Sokotra and is preserved in the British

Museum ; the specimens procured by Mr. Peel are in the Hope
Museum.

29, ARANEUS NAUTIcUS L. Koch.

A single female, most likely of this species, from Berbera, in
the Hope Museum.

3. CEBRENNUS ZTHIOPIUS Simon.

A single immature female, probably referable to the Abyssinian
species, was taken in Eastern Central Somaliland. In the Hope
Museum.

4 & 5. OXYOPES sp. ?

Two specimens from Western Somaliland, each representing an
indeterminable species. In the Hope Museum.

Order SCORPIONES.

Family ScorPionip&.
Genus Panpinus Thorell.

PANDINUS PUGILATOR, sp.n. (Plate IV. figs. 1, 1 a.)

Colour of chelx, tail, and upperside of body olive-brown, the
hands rather paler ; legs and vesicle of tail clear pale yellow.

Carapace smooth, sparsely punctured above, granular at the
sides, its length exceeding that of the movable digit and almost
equalling that of the 3rd and 4th tail-segments.

Tergal plates granular laterally ; the last more or less granular
throughout. Sternal plates smooth, the last weakly crested.

Tail short and slender, less than three times the length of the
carapace, considerably narrowed posteriorly: the 4th segment
about twice as long as wide, the 5th considerably more than twice
as long as wide; Ist segment weakly granular above, for the rest
smooth, all its keels smooth; 2nd segment entirely smooth, 3rd
with the four inferior keels strong, rugose, the intervening spaces
granular; 4th with the infero-lateral keels strong and denticulated,

[50]

1900. | INSECTS AND ARACHNIDS FROM SOMALILAND. 59

the inferior median heels not differentiated amongst the coarse granules
covering the lower surface of the segment; 5th segment with the
inferior keels denticulated, the area between them coarsely granular;
upperside of the 5rd, 4th, and 5th segments smooth, the keels
also smooth or at all events only roughened with pores ; vesicle
granular, narrow, its width equal to its height and only about
one-third the length of the vesicle and aculeus.

Chele: humerus coarsely granular above, its anterior surface with
strongly granular crests; lower surface granularly crested behind ;
brachium finely granular in front, also roughened with pores and
granules behind ; hand wide, its width about equal to the length
of the movable digit, its upper surface smooth posteriorly on the
lobe, the area just above external keel coarsely granular, the rest
covered with low more or less anastomosing tubercles, which are,
however, more distinct towards the base of the immovable digit ;
the inner edge smooth, though the granules of the lower surface
run right up to it or even project slightly beyond it; the lower
surface sparsely granular, with two weakly defined keels; keel
defining the hand-back above very strong and prominent.

Legs smooth; protarsal segment of Ist and 2nd with a single
external apical spine; tarsi armed with eight spines, two on each
side being on the lobe, one on its lower angle, the other in the
middle as in P. colei, P. bellicosus, &e.

Peetinal tecth 17.

Measurements in millimetres. Totallength 93 ; length of carapace
18, of tail 48; width of Ist segment 5°8, of 4th 3:5, of vesicle 3-2 ;
width of hand 16, length of movable digit 17.

Loc. North-west Somaliland (Berbera or Hargaisa). A single
specimen (type) in the British Museum.

Recognizable by the granulation of the lower side of the 4th
caudal segment and the obsoleteness of its inferior median
crests, &e.

PANDINUS PEELI, sp. n. (Plate IV. fig. 2.)

3. Closely allied to P. colei (Pocock), but differing in the
characters pointed out in the table given below (cf. p. 62).

Carapace quite smooth above, sparsely punctured, much less
closely granular laterally than in P. colei. Terga weakly granular
at the sides only, not closely granular throughout the posterior
half as in P. colet. Chele larger than in P. colei; upper crest of
brachium smooth ; upperside of hand externally granular as in
P. colet, but internally much smoother, the granules anastomosing
and running together into ridges which become almost obsolete
on the posterior lobe of the hand. Fourth abdominal sternum
obsoletely granular in the middle. Pectinal teeth 15.

Measurements in millimetres. Total length 81; length of carapace
14:5, of tail 38, of underhand 10; width of hand 13:5.

Loc. North-west Somaliland (Berbera or Hargaisa), A single
specimen (type) in the British Museum.

[51]

54 MR. C. V. A, PEEL AND OTHERS ON (Jan. 23,

Family Burnin”. P

Genus Uropnectes Peters.

UROPLECTES FISCHERI (Karsch).

Lepreus fischeri Karsch, Mitth. Miinch. ent. Ver. 1879, p. 124.

The two specimens of this species that were obtained in North-
west Somaliland (at Berbera or Hargaisa) agree closely in
characters with those collected by Dr. Donaldson Smith at Lummo
and Turfa, and discussed on pp. 400-401 of that author’s account
of his expedition. One of Mr. Peel’s specimens is in the British
Museum, the other in the Hope Museum at Oxford.

Genus Buruus Leach.

PUTHUS CALVICEPS, sp. n. (Plate IV. figs. 3-3 a.)

Colour (dry specimen): trunk blackish yellow; appendages
uniform pale yellow.

Carapace weakly granular ; keels almost entirely obsolete, only
the anterior median distinct but failing to attain the front border
of the carapace. Tergal plates rather coarsely granular; keels
normal and granular. Sternal plates smooth and polished; the
last very finely granular at the sides, polished and rather coarsely
but sparsely punctured in the middle; the lateral keels almost
obsolete, represented by about three larger eranules.

Tail of medium thickness, nearly ane ace the length of the
carapace, posteriorly narrow ed; the sides and upper surface of the
segments normally crested, the intercarinal spaces finely granular ;
the median lateral keel complete on the 2nd and almost “complete
on the 3rd segment ; lower surface of Ist, 2nd, and 3rd segments
rather coarsely but strongly punctured with setiferous pores; the
median keels of the 1st obsolete, those of the 2nd and 3rd strong
and denticulated, increasing in strength posteriorly, the inferior
laterals of these segments also strong and denticulated, converging
posteriorly and fusing, like the medians, with a transverse granular
crest; lower surface of the 4th w eakly granular, without median
keels ; 5th segment without a trace of superior lateral keels, finely
granular below, with a few coarser granules intermixed, the median
keel denticulated; the laterals strongly denticulated, with at least
one large lobate tooth behind the middle of their length, ending
behind in a big subdivided lobe on each side of the anal aperture ;
vesicle of medium size, smooth, punctured.

Chele weak ; humerus granular above, normally crested ; bra-
chium smooth, weakly crested, anterior surface with two weakly
granular keels ; ; hand smooth, small, narrower than brachium,
movable digit with eight rows of teeth along the middle line, the
basal row long and rising right at the extremity of the segment,
the lateral teeth forming short oblique rows of three each, two

[52]

1900. | INSECTS AND ARACHNIDS FROM SOMALILAND, 50

outer and one inner ; finger a little more than twice the length of
the underhand.

Legs with granularly crested femora ; the feet with two rows of
hairs below.

Pectinal teeth 21.

Genital operculum considerably longer than sternum.

Measurements in millimetres. Total length 31 ; length of carapace
3°5, of tail 17°5; length of movable digit 3.

Loc. North-west Somaliland (Berbera or Hargaisa). A single
specimen (type) in the British Museum.

In size and some points of structure this little Buthus approaches
Nanobuthus andersoni Poe., obtained at Duroor to the north of
Suakim (Journ. Linn. Soc., Zool. xxv. p.314). The dentition of the
mandible and of the digits of the chele, however, is quite normal
for the genus Bu‘hus. The most striking structural peculiarities
of the species are : (1) the obsoleteness of all the cephalothoracic
keels with the exception of the anterior median: (2) the coarse
but sparse punctuation of the last sternite and of the lower side of
the first caudal segment, accompanied as it is by the disappearance
of the median keels ; (8) the disappearance of the median keels on
the lower side of the 4th caudal segment. These characters do
not co-exist in any species known to me. Judging from the
structure of the inferior lateral keels of the 5th caudal segment
and of the inferior keels of the 2nd and 8rd segments, this species
belongs to the same category as B, occitanus.

10. GENERAL List oF THE SCORPIONS OF SOMALILAND AND
THE Boran Country. By R. I. Pocock.

Family Burn.

Genus Uropiectss Peters.

UROPLECTES FISCHERI (Karsch).

Lepreus fischeri, Karsch, Mitth. Miinch. ent. Ver. iii. p. 124
(1879).

Loe. Barawa (Karsch), Turfa and Lummo (Donaldson Smith),
and Berbera and Hargaisa (C. V. A. Pecl).

To the south of Somaliland the typical form of this species is
replaced by two subspecies, one paler, the other darker than
U. fischeri typicus. The three may be contrasted as follows :—

a, Hands entirely pale, body banded above as in
UEHOGE UBJIOGDS » ooncscoonaccsteseesacsbooogaenacoonce fi. flavimanus, subsp.
b. Hands wholly black or at least lined with black [noy.
and black at base of fingers.
a’, Terga yellow, with a pair of black spots ; cara-
pace and terga broadly yellow at the side... fi. typicus.
b'. Terga mostly black, with a narrow median
and lateral marginal band, much less yellow
AISI EHOt CATA PACE. .cnaccseeneeasseceeeneaienss ae ji. nigrimanus.

[53]

56 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

The subspecies nigrimanus was based upon a single example
from Mombasa (see Proc. Zool. Soc. 1890, p. 130, pl. xiv. fig. 2).
U. flavimanus is based upon a specimen in the British Museum
obtained by Mr. J. Wilson at Mombasa in British East Africa.

It is interesting to note that the two subspecies most distinct
from each other, namely, U. f. flavimanus and U. f. nigrimanus,
occur in the same locality, whereas the subspecies intermediate
between them is found elsewhere.

Genus Parapurnus Pocock.

PARABUTHUS GRANIMANUS Pocock.

Parabuthus granimus, Pocock, Journ. Linn. Soe., Zool. xxv.
p- 311 (1895).

Loc. Zeyla in North-west Somaliland (4. W. Oates), Goolis
Mountains (Lort Phillips). In the British Museum.

PARABUTHUS HETERURUS Pocock.

Parabuthus heterurus, Pocock, in Donaldson Smith’s ‘ Through
Unknown African Countries,’ p. 402 (1897).

Loe. Hargaisa, Silul, Shebeli River (A. Donaldson Smith) ;
Goolis Mountains (#. Lort Phillips). In British Museum.

The two species of this genus may be recognized as follows :—

a. Hand and brachium closely granular ; 5th segment of

tail strongly infuscate at least below ..............-.6-.-+ P. granimanus.
6. Hand and brachium smooth, punctured, hairy; 5th

caudal segment clear yellow throughout, 4th segment

andiwvesicle black: cicccc.c.pessesssesee acsadeneesccesecmerees P, heterurus.

Genus Buruus Leach.

Boruus occrranus (Amoreux), subsp. nov. BERBERENSIS.

Colour yellow, with black lines along the keels of the upperside
of the trunk, humerus, brachium, and hand, also the distal half of
the femora infuscate and the base and keels on the patella. Body
and tai crested and granular as in the typical form, the granules
of the inferior keels of the 2nd and 3rd caudal segments about
as much enlarged as in the Spanish form; external surface of
hand with a few granules, a pair of strong granular finger-keels
running along its upperside and inner edge.

Total length 38 mm.

Somaliland (Miss Gillett). In British Museum.

Subsp. nov. ZEYLENSIS.

Colour a uniform reddish yellow, the appendages and _ tail
clearer than the trunk. Frontal intercarinal area of carapace

covered with granules; a median row of granules running along
ix
[54]

1900. ] INSECTS AND ARACHNIDS FROM SOMALILAND, 57

the middle of the ocular tubercle. erga closely and finely
granular throughout. Inferior keels of 2nd and 3rd caudal
segments strongly dentate. Hand smooth, with a pair of weak
and weakly granulate crests on its upperside.

Total length 50 mm.

Loc. Zeyla in North-west Somaliland (Z. W. Oates). In British
Museum.

BurHuwus CALVICEPS Poe.
Cf. supra, p. 54.

BUTHUS ACUTECARINATUS Simon.

Buthus acutecarinatus, Simon, Ann. Mus. Genova, xviii. p. 245,
pl. vii. fig. 13 (1883).

Loc. Leyla (EL. W. Oates). In British Museum.

Occurs also in Egypt and Arabia.

Buruus potysrictus Poe.

Buthus polystictus Pocock, Ann. Mag. Nat. Hist. (6) xviii. p. 178,
pl. xi. fig. 1 (1896).

Goolis Mountains in Somaliland (4. Lort Phillips). In British
Museum.

Buruts EMINI Pocock.

Buthus eminii, Pocock, Ann. Mag. Nat. Hist. (6) vi. p. 98
pl. i. fig. 2 (1890) ; 2d. op. cit. (6) xviii. p. 179 (1896).

Loc. Aimcea in the Boran Country, 3000 ft. (A. Donaldson
Smith). In British Museum. *

This species and perhaps also B. polystictus may prove to be
subspecies of B. trilineatus Peters, described from Tete.

The species and subspecies of Buthus mentioned above may be
diagnosed as follows :—

a. Inferior lateral keels of 5th caudal segment posteriorly
lobate ; inferior median keels of 2nd and 3rd with the
granules enlarged towards the posterior end.
a’, Carapace without distinct median, lateral, and
posterior keels, inferior keels on 4th caudal segment
OWSOLELE eee rae tee ac eciseceioose cc setes aston ont omase calviceps,
b', Carapace with distinct median posterior and lateral
keels, the posteriors turning outwards and uniting
with the laterals, inferior keels on 4th caudal segment
eyelopedinr aca. ssecctinsatheccmsactet soenarccesenesucnscoase oceitanus.
a*, Legs, chele, and body uniformly yellow, not
lined with black ; intercarinal ocular area closely
@ranllanysy.c+sehosgsdiw see sacs ce seadaracsencdeeasteedees subsp. ceylensis.
6?, Chele and body with black-lined keels, femora
of legs distally infuscate ; patella also infuscate
basally and along keels; frontal area of carapace
sparsely granular in the middle ..................... subsp. berberensis.

[55]

58 MR. C. V. A. PEEL AND OTHERS ON (Jan. 23,

b. Inferior lateral keels of 5th caudal segment and inferior
median of segments 2 and 3 uniformly granular
throughout.
a3, Hand carinate and densely granular; dorsal ab-
dominal keels posteriorly strongly spiniform......... acutecarinatus.
}°, Hand smooth, not carinate ; dorsal abdominal keels
not strongly spiniform posteriorly.
a‘, Less coarsely granular; tail thinner and lower,
height of 4th segment barely half its own length
and distinctly less than length of Ist; upper
surface of caudal segments much less strongly
COGAN, CMOS Goancdcdecoentasddecodacuspocousacogos00c polystictus,
b4, More coarsely granular; tail thicker, its superior
keels more strongly elevated; height of 4th
segment more than half its length and equal to
length of Ist ..... sobenne yadadnoaccanapodcocadsppuconsade emini.

Family ScoRPIONID#.
Genus Panprnvs Thorell.

PANDINUS MEIDENSIS Karsch.

Pandinus meidensis, Karsch, Mitth. Miinch. ent. Ver. iii. p. 127
(1879); Kraepelin, Das Tierr., Scorpiones, &ec. p. 119 (1899),
Loc. Meid in Somaliland. In Berlin Museum.

PANDINUS SMITHI (Pocock).

Scorpio smithii, Pocock in Donaldson Smith’s ‘Through Unknown
African Countries,’ p. 198 (1897).

Lee. Hargaisa, Silul, Abdeh, and Turfa in Somaliland (A.
Donaldson Smith). In British Museum.

PANDINUS PALLIDUS (Kraepelin).

Scorpio pallidus, Kraepelin, Mitt. Mus. Hamburg, xi. p. 60 (1894).

Pandinus pallidus, id. Das Tierr., Scorpiones, &e. p. 120 (1899).

Loc. Barawa in Somaliland. In Hamburg Museum and British
Museum.

The typical form of this species was based upon immature
individuals measuring only up to 75 mm. long. Until the adult
is known it seems to me impossible to classify the species with
certainty. It is undoubtedly nearly alhied both to the following
species, P. phillipsi from North-west Somaliland, and to the
more southern Masailand form P. gregorii, but it will probably prove
to be at all events subspecifically different from both.

PANDINUS PHILLIPSI (Pocock).

Scorpio phillipsit, Pocock, Ann. Mag. Nat. Hist. (6) xvii. p. 101
(1896).

Pandinus phillipsti, Kraepelin, Das Tierr., Scorpiones, &e. p. 120
(1899).

Loc. Dooloob and the Goolis Mountains, inland of Berbera
(KZ. Lort Phillips). In British Museum.

[56]

—

1900. | INSECTS AND ARACHNIDS FROM SOMALILAND. 59

This form is evidently allied to the typical P. pallidus ; but
until adults of the latter come to hand for comparison, it is im-
possible to say what the exact relationship between the two may
be. The original examples of P. phillipsi are a pair of females
obtained at Dooloob. Mr. Lort Phillips subsequently procured an
adult male and a young female on the Goolis Range of mountains.
The former has 17-18 pectinal teeth, a longer tail and larger vesicle
than the female, and lobate movable finger on the chela. The
young one is as large as a co-type of P. pallidus, the carapace in the
two measuring 11 mm. Moreover the posterior tarsal lobe is tipped
above with bristles as in P. pallidus, not with a spine as in the adult
P. phillipsi. But the shape of the hand in the young P. phillipsi
is different, this organ being very noticeably narrower, and the
tubercles on its upperside are much sharper and more strongly
defined.

The following actual measurements (in millim.) of the two
examples may be advantageously compared :—

Total length Length of Length of Length of Width of

(without Carapace. brachium. underhand. movable hand.
: vesicle). finger.
eee
D haee \ 68 1 75 7 Wl 85

As will be seen, there is practical identity of measurements '
except where the width of the hand is concerned.

PANDINUS COLEI (Pocock).

Scorpio coler, Pocock, Ann. Mag. Nat. Hist. (6) xviii. p. 180,
pl xstigs. 2, 2.0 (1896).

Pandinus colet, Kraepelin, Das Tierr., Scorpiones, &c. p. 120
(1899).

Loc. Berbera and Goolis Mountains (2. Lort Phillips). In
British Museum.

This species was based upon a subadult specimen from Berbera.

Mr. Lort Phillips subsequently procured in the Goolis Mountains
and kindly sent to the British Museum three additional examples,
an adult male and female and a young specimen considerably smaller
than the type. The characters upon which the species was based
prove perfectly constant. The adult male and female are much
alike ; the former, however, has the terga of the abdomen finely and
closely granular posteriorly, whereas in the female they are nearly

' The difference in total length in this and in many other cases is due to the
degree of distension of the abdominal region. The length of this region is su
very liable to alteration in accordance with the mode of preservation of the
specimen after death, and depends so largely upon the fasting or full-fed,
pregnant or not pregnant condition of the Scorpion, that the relative lengths
of the tail as compared with the trunk, which Kraepelin, Karsch, and others so
frequently quote, have but little importance. The length of the carapace, which
does not vary, should be taken as standard for comparison.

[57]

60 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

smooth. Again, the 2nd and 4th abdominal sterna are finely
granular in the middle; and, lastly, the vesicle is more strongly
inflated, its width considerably exceeding its height and being
equal to that of the 4th caudal segment.

In the female the width of the vesicle is scarcely greater than
its height and less than the width of the 5th segment.

Pectinal teeth 11-13(3 9).

Total length (3 )82 mm., carapace 18°5, tail 38, underhand 8-5,
width of hand 12.

PANDINUS PEELI Pocock.
Cf. supra, p. 53.

PANDINUS HAWKERI, sp. nD.

Colour of carapace and palpi yellowish brown ; tergal plates and
tail reddish brown ; vesicle brown with yellow lines ; legs entirely
pale yellow, abdominal sterna testaceous.

Carapace smooth, polished; terga also smooth, punctured
along the posterior margin, the last very weakly granular laterally,
the crests obsolete. Sterna, with exception of the last, smooth ;
the last obsoletely crested, but mesially closely granular.

Tail short, only a little more than two and a half times the length
of the carapace, which is almost as long as its first three segments;
the inferior median keels absent on segments 1-5, the inferior
laterals present, smooth on segments 1 and 2, granular on 3 to 5,
the area of the lower surface of the tail between them granular as
in P. colet; superior and superior-lateral keels of tail weak,
punctured, but not granular; sides of tail smooth ; upper surface
at most very sparsely granular, except along the posterior edge of
segments 1-4, where there is a series of denticuliform granules ;
vesicle strongly punctured and setose beneath, but scarcely
granular, its width exceeding its height. Chele: humerus smooth
below and behind, the crests on its upper and anterior surfaces
coarsely granular, its upperside sparsely granular in the basal
half ; brachium smooth, except for some minute granules in front
and some coarser ones along the anterior inferior crest: hand
moderately wide, its width equal to three-fourths the length of
the carapace ; upper surface granular on the external slope : above
the strong keel of the underhand; the rest of the upper surface
nearly smooth and polished, beset with a fine reticulation of ridges
which are almost obsolete in adult, coarser in young; inner edge
almost smooth, punctured, lower surface sparsely and w eakly
granular towards the base of immovable digit; immovable digit
with its basal width less than half the length of is biting- -edge ;
movable digit shorter than carapace, exceeding width ot hand,
equal to length of 3rd and 4th caudal segments.

Legs smooth; protarsi of Ist and 2nd with one posterior apical
spine; tarsal lobes with two spines ; lower surface of tarsi with
one anterior and three posterior spines; anterior claw much

[58]

1900. | INSECI'S AND ARACHNIDS FROM SOMALILAND, 61

weaker than posterior. Sternwm long, about one-third longer
than wide.

Pectinal teeth 12-15 in @.

Measurements in millimetres. Total length 81, carapace 15,
tail 50; width of hand 12; length of movable digit 13, of hand-
back 8.

Loc. Jifa Uri inland from Zeyla (2. M. Hawker). In British
Museum.

Differing from P. colet and P. peeli in the characters pointed
out below (p. 62).

PANDINUS PUGILATOR Poe.
Cf. supra, p. 52.

PANDINUS MILITARIS, Sp. 0.

Scorpio bellicosus, L. Koch; Pocock, in Donaldson Smith’s
‘Through Unknown African Countries,’ p. 397 (1897) (nee P. belli
cosus L. Koch).

2. Colour yellowish brown; legs paler yellow, hand reddish
brown with black fingers. Carapace granular laterally, entirely
smooth above except for a few granules in the anteocular groove.
Terga finely granular laterally. Tail a little more than two and a
half times as long as the carapace, the inferior keels on segments
1-3 quite smooth, those on 4th at most slightly rugose; superior
and superior-iateral keels of all the segments granular or weakly
denticulated ; superior surface of Ist granular, of 2nd less so;
width of Ist exceeding length of 3rd, almost equal to that of 4th ;
5th about twice as long as wide. Chele: humerus granular above
at least on its basal half, smooth below; brachium almost entirely
smooth, its anterior side weakly granular above, more coarsely
below ; hand wide, its width in adult exceeding length of 3rd and
4th caudal segments, inner edge smooth, upper surface smooth,
finely reticulated, a few low tubercles just above the keel of the
underhand and at the base of the immovable digit; external
portion of upper surface rising vertically above keel of underhand ;
thickness of hand at the front equal to length of 4th caudal
segment. Lower side of hand granular distally, scarcely crested.

Pectinal teeth 12-15.

Measurements in millimetres (of type). Total length 112,
carapace 19, tail 51 ; width of hand 16-8, underhand 10-5.

Loc. Aimola in the Boran Country (Donaldson Smith); also Ndi,
on the Weiss Road inland from Mombasa (C. Stewart Betton). In
British Museum.

Nearly allied to the East-African species P. cavimanus, but
differing in the following characters :—the carapace in the female is
longer than the 4th and 5th caudal segments, and the basal width
of the immovable finger is only about half the length of its free
margin ; whereas in P. cavimanus (2 ) the carapace is shorter than
4th and 5th caudal segments, and the basal width of the immovable
digit is about two-thirds the length of its free margin.

[59]

62 MR. C. V. A. PEEL AND OTHERS ON [Jan. 23,

T at one time supposed this species to be the female of the
Abyssinian P. bellicosus L. Koch, but judging by Kraepelin’s recent
diagnosis of the latter species (Das Tierr., Scorpiones, p. 121,
1899), P. malitaris certainly differs in having the superior caudal
keels denticulated, the hand finely punctulate above and. below,
and the last abdominal sternite scarcely visibly crested.

Synopsis of the Somali Species of Pandinus.

a. Median eyes always some distance behind middle of cara-
pace; tarsi more numerously spined, the lobes with 3-4
spines, total number on lower side of tarsi 9 behind,
6 in front.
a‘, Tarsal lobes with 4 spines, a strong spine being on the
tip of the lobe; humerus of chela furnished below
with two short rows ef denticles.................0..000-0++ meidensis.
. Tarsal lobe with 3 strong spines, the spine on the tip
smaller and usually filiform distally; humerus of
chela smooth below.
a. Ornamentation of hand consisting of conical
tubercles ; inner margin of hand distinctly tuber-
cular and denticulate from base of finger to carpal
articulation, width of hand greater than length of
carapace; pectinal teeth 18-21 ........................ smithi.
6?, Ornamentation of hand consisting of low rounded
or irregular shaped, often anastomosing tubercles ;
lobe of hand with smooth posterior edge; length
of carapace exceeding width of hand; pectinal teeth
PO =AWS Peo eras ne sacs dabsncacese comes teciaencutseecoensseciet phitlipst.
4. Median eyes in middle of carapace, rarely a little behind
the middle; total number of spines on tarsi 5 behind,
3 in front, 2 only being situated on each lobe, the tip of
which is furnished with bristles.
a®. Lower surface of all the caudal segments and the
middle of at least the last abdominal sternite closely
granular and not keeled.
a‘. Upper surface of hand from inner edge to crest
of anderhand uniformly covered with coarse
granules which do not anastomose; upper crest
of brachium distinctly granular,............/..........- colei.
b*. Upperside of hand coarsely granular only above
crest of underhand, the rest of its upperside either
almost smooth or ornamented with low rounded
tubercles which run into ridges; upper crest
of brachium quite smooth.
a°. Upperside of hand covered with ornamentation
of low, more or less anastomosing tubercles ;
its lower surface distinctly granular; hand
larger, carapace equal to length of underhand
+ one-third of movable digit, and only as
long as the hand from the posterior edge of
the lobe to the base of the immovable finger ;
immovable finger not twice as long as its basal
WLC Ge nciecstelectisesiasemcue te mshiasitracceoreseceocctscctst peel.
6°. Upperside of hand smooth, at most ornamented
with a network of low ridges, lower side very
sparsely granular; hand smaller, carapace as
long as underhand + half the movable finger,
and as the hand measured from the posterior
edge of the lobe + half the immovable finger ;
immovable finger twice as long as its basal width. hawkert.

[60]

1900.] INSECTS AND ARACHNIDS FROM SOMALILAND. 63

53, Last abdominal sternite and lower side of Ist and

Fig.

Fig.

Fig.

Fig.

a’.

2nd caudal segments, and usually also of 3rd,
smooth ; caudal segments 1-5 keeled below.
Lower surface of 4th caudal segment coarsely but
sparsely granular, the median keels obsolete, the
lateral keels strongly denticulate; superior and
superior-lateral crests of tail smooth; protarsal
segment of Ist and 2nd legs with a single external
apical SUING): sndapaneddecnodeciGboras daaapeoccus coco cHOn Gane pugilator.

6°. Lower surface of 4th caudal segment not uniformly

and coarsely granular, furnished with four normal,
subequally strong keels; superior and superior-
lateral crests of tail denticulated ; protarsi of Ist
and 2nd legs with at least two external spines...... militaris.

EXPLANATION OF THE PLATES.
Prats I,
Lepidoptera, Coleoptera, &e., from Somaliland.

1. Pseudophia oppia, p. 18. Fig. 9. Genecerus nebulosus, p. 28;
2. Chilena sabrina, p. 20. 10. Pstloptera somalica, p. 25.
38. Cerocala munda, Palo: 11. Agrypnus longicornis, p. 26.
4, Acrea mirabilis, yop Able 12. Sepidium magnum, p. 29.
5. Belenois peeli, 3, p. 15. 13. Rhytidonota robusta, p. 28.
6. Dato: 14. Sepidium bilobatum, p. 30.
(f Pseudophia lineata, p- 19. 15. Julodis laticollis, p. 25.
8. Pangonia tricolor, p. 7. 16. Trox expansus, p. 22

Prate II.

Orthoptera from Somaliland.

1. Rhegmatopoda peeli, p.44. Left elytron. ¢.
2. Otiaphysa eS p.43. Left elytron. Q.
3. Sauracris lacerta, 3, Ihe
4. Charieis peeli, 2, p. 31. (Antenne and apex of abdomen missing.)
5. Milititsa somaliensis, p. 43. Right elytron.
6. Burria longixipha, 2, p. 38.
7. Milititsa somaliensis, p.43. Head and pronotum,

Prats IIT,

Ticks from Somaliland.

1. Rhipicephalus marmoreus, p.50. $. Dorsal view. X11 times linear.
Ia is 45 Ventral view.
1d, _ ze Capitulum and mouth-parts from below.
ives * oe Coxe of 1st and 2nd legs.
hah, Stigma, adanal plate, and anus of left side.
Zz Rhipicephalus ar matus, p- 50. G. Dorsal view. X11 times linear.
ide 5% 5 Ventral yiew.
2b, = ‘ Capitulum and mouth-parts from below.
Jie: % 3 Coxa and trochanter of Ist leg.
2d. i 7 Adanal plate of right side.
2e. 3 os Lateral view of posterior end of abdomen show-

ing adanal plate, spike above it, and stigma,
QF. + ss ©. Dorsal view.

Puate IV.

Scorpions from Somaliland.

1. Pandinus pugilator, p. 52. Nat. size.
la. A Fe Underside of tail.
2. Pandinus peeli, p. 53. Underside of tail.
3. Buthus calviceps, p. 54. Carapace.
Oils = . Underside of 3rd, 4th, and Sth caudal segments.
SiO ees R Chela.

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LEPIDOPTERA, GOLEOPTERA, ETC, FROM SOMALI-LAND

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SCORPIONS FROM SOMALI-LAND.

97

[Reprinted from The Entomologist’s Record, &c., Vol. XII., April, 1900.]

On the British Orthoptera in the Hope Museum, Oxford.
By MALCOLM BURR, F.Z.S., F.E.S.

In the Hope collection, in the University Museum, at Oxford, there
are a number of old specimens of considerable interest, as many are
labelled in the handwriting of the late Professor Westwood, and some
in the handwriting of Stephens. Professor Poulton has been kind
enough to permit me to examine and rearrange them. ‘The following
notes on some of the older specimens may be of interest.

Forricutaria.—Labidura riparia, Pall.—One g, two 9. “ Eng-
land.” No further information. on the labels. They are very probably
some of the original specimens taken near Christchurch. Anisolabis
maritima, Bon.—‘‘ Northumberland. G. Wailes. 1857, end of Sep-
tember.’’ These are evidently not some of the original examples taken
by Bold at South Shields, but those captured a few years later by
George Wailes, when he took a number of specimens for his careful
observations of the development of the antenne and abdominal
segments, recorded by him in the Zoologist, vol. xvi., p. 5895, in a
paper that has to a large extent been overlooked, in spite of its great
interest. The insect was taken in heaps of ballast emptied by ships
returning from abroad, and it is practically certain that it was an
importation. It was first captured in 1856, and was still to be found
in 1860. It is unknown whether the. species still exists there. Labia
minor, L.—Six g, ten 2. ‘Snowdon.’ Forficula auricularia, L.—
Numerous examples. Two are labelled ‘ borealis”? in what I believe
to be Stephens’ handwriting; these are what I have always regarded
as the variety forcipata. The difference between the two is merely a
matter of the comparative length of the forceps. I. lesnei, Fin.—One
9. ‘Kingstone,’ an old specimen. Apteryyida albipennis, Meg.—
One pair, old, and in bad condition. There is no locality attached,
but the specimens are probably some of the originals taken at Ashford
by Westwood. In his handwriting they are labelled ‘‘/’. centralis,
Westw., MS.”

Buatropra.—Lctobia lapponica, L.—Ten gf. ‘Sunning Hill,
Berks.” . livida, Fabr.—‘‘ Rudd. N. F.” This species has been
previously recorded from the New Forest. EH. panzeri, Steph.—Several
examples. ‘Two males labelled ‘‘ J. C. Dale, 1865.” One specimen, a
male, is labelled ‘‘ Blatta nigripes, Mus. Steph., in Notes on habits,
B. G. C.,” but it is too pale in colour to be the true form nigripes, and
does not agree with Stephens’ own description. It is the ordinary
typical form. There are several specimens considerably darker in
colour, approaching more nearly to the true niyripes. One dark speci-
men is labelled ‘“ Rev. W. Kirby, lapponica.” There is one ? with
ootheca labelled ‘‘ Black Gang Chine.”’ Phyllodromia germanica, L.—
This species is usually considered to have been introduced into England
by the soldiers returning from the Crimea, but we have distinct
evidence here that it was numerous in parts of England before that
date. There are several old specimens, one of which is labelled
‘“‘Infesting the kitchens of houses at Kildare, Ireland, living behind

98

the skirting and abounding in the crevices of the kitchen table,
destroying all kinds of paper and in many ways very troublesome,
March 1852,” and there is another specimen labelled in the same
handwriting, ‘“‘ Mr. Gray Dudley ; taken by him in his garden at the
foot of Castle Hill.”’ Reports of this species taken out of doors are
rare. It occurs in a wild state in woods in Prussia and Russia, so
there is no reason why it should not become settled wild in this
country, as it has become firmly established in a domesticated condi-
tion. latta orientalis, L.—Several specimens unlabelled ; one @?
nymph ‘“‘ under bark of tree, 10ft. up, K. G.”” (?=Kew Gardens). It
is rarely that this species is taken out of doors, for it seldom leaves the
warm kitchens and hot corners that it loves. It would probably be
unable to exist in the natural state in this country. There is one
crushed and mutilated nymph, mounted on card with the label ‘‘ Larva
of a Blatta, sent by Mr. Backhouse from Gateshead, as a flea, and
exhibited by J. O. W. at Ent. Soc. as Pulea imperator.” This is the
original type of Westwood’s famous mistake. He exhibited this
specimen at the Mntomological Society (Proc. Hunt. Soc., 2nd series,
iv., p. 70) in 1857, as a new species of flea, twenty times the size of
the largest flea hitherto known! He corrected his mistake in the
following year (op. cit., v., p. 60). There is also a broken ootheca with
the label ‘‘ Kge-pouch of B. orientalis? Mr. W. Baird in acct.
amongst ent. scraps. For the parasitic Chalcididae bred see them
among Chalcididae.”’ Ihave carefully searched through the drawers
of the British Chalcididae and have been through Walker’s Monoyraphia
Chalciditum but cannot find the specimens referred to.
AcripiopEA.—Mecostethus yrossus, L.—F ive g and three ?. With-
out locality, labelled ‘‘ Locusta flavipes.” Stenobothrus lineatus, S.
viridulus, S. rufipes.—Several specimens with no interesting labels. S.
bicolor, Charp.—Many specimens without labels. One ¢ labelled
‘““erucigera,’ and another g ‘Isle of Purbeck, 1830. L. rubicunda.”
This is apparently the pink form known as purpurascens, Fieb. The
name rubicunda has usually been referred to S. viridulus, of which a
reddish form is sometimes taken. 8S. parallelus, Zett.—Several
nymphs, and one empty nymph-skin, ‘‘ Blenheim Park, 1832.”
Gomphocerus stbiricus, L.—One 3 unlabelled, which is most probably
Stephens’ type, said to have been taken at Netley. G. rufus, 4.—One
S$ unlabelled. G. maculatus, Thunb.—One g ‘ Devil’s Ditch,
Newmarket Heath, July 2nd, 1833. J.C. Dale.” Pachytylus miqra-
torius, L.—There are four specimens of undoubted miyratorius, showing
that this species has really occurred so far west as this country. One
male is unlabelled, another ‘‘ Littlehampton, Sussex, 1846, J. O. W.”
One @ ‘alive near Oxford, J. O. W.” and a second @? ‘Ct. alive near
Chepstow, September 16th, 1857.” [There are no specimens of P.
danicus, L.|. Schistocerca peregrina, Oliv.—T wo specimens, one labelled
“F. Bond, Esq., near Falmouth, a number of specimens near
Plymouth,” the other ‘ Cornwall, F. Bond, Esq., 1870.” A good
many examples of this locust were taken in Britain in 1869, but I do
not think any have been recorded for the following year. Tettia
subulatus, L.—‘* Cambridge.” Some are labelled “ bipunctatus.” T.
bipunctatus, L.—About 380 specimens without labels.
Locustopra.—Leptophyes punctatissima, Bose.—A number of
unlabelled specimens. One female ‘“ caught October 1st, Coombe

99

1806, clypeata, Pz., Mr. Neale.’’ One 3, two @, ‘‘ Miss Badcock, on a
pear tree,” another ‘‘ Hphippisora virescens, Steph., /U., vi., 16, vili., 33.”
Ephippigera virescens, Steph., is known to be a synonym for L.
punctatissima, but ‘‘ clypeata, Panz.,” is usually regarded as a synonym
of Thamnotrizon cinereus, Lu. Meconema varium, Fabr.—One male
‘«« Sept. 16th, 1833, Foxley Wood, Norfolk,” another ‘‘ Taken in second
week in November, ’66, Mr. Briggs, St. John’s Coll.” This species
is one of our latest Orthoptera, and may be taken well into the winter.
One female, ‘‘ Miss Badcock, on a pear tree.”” Xiphidiwm dorsale, Latr.
—One male labelled ‘ Acrida fusca,’’ and a male ‘ Norfolk, August
Ist, A. fusca, Pz.,” also four 3, seven ? and two nymphs, all labelled
“« fuscum.”’ Stephens obviously confused X. dorsale with X. fuscum, a
very distinct species, which is far more common and widely distributed
on the continent than YX. dorsale. Locusta viridissima, L.—One male
« Parks, Oxford, 1848 ;”’ several unlabelled specimens. Thamnotrizon
cinereus, Li.m—One male ‘ 1). apterus,’ one @ ‘near Croydon,” one 2
“‘ enhippiger’’ and four 3 and four ? unlabelled. Platycleis yrisea,
Fabr.—Several specimens without labels, one nymph queried ‘ viridis-
sima, ex Museo D. Hill.” P. brachyptera, L.—One 3 labelled
‘* brachyptera’’ and one ‘‘ kirbit”” also a male labelled “ clypeatus”’ and
several nymphs “ kirbit.”” Decticus verrucivorus, L.—A male ‘‘ bingleit,
Rey. D. Bineley,”’ probably one of Curtis’ types, but of the green form,
also one female ‘ verructvorus.”

GryLLopEA.—Nemobius sylvestris, Fabr.—Several specimens of both
sexes, unlabelled. Gryllus campestris, L.—One male ‘‘ Weaver, N. F.”
and several specimens without labels, including one of the very rare
variety with fully developed wings, projecting well beyond the elytra.
[There is also a specimen with wings projecting far beyond the apex
of the abdomen, which I refer with some doubt to G. bimaculatus, de
Geer, a meridional species. It may be an extreme form of G. campes-
tris, or may have been accidentally introduced.] G. domesticus,
several examples, unlabelled. Gryllotalpa gryllotalpa, L.—Five 3,
four @, unlabelled.

?

THE METHODS OF SETTING AND LABELLING
LEPIDOPTERA FOR MUSEUMS.

By E. B. Poutton, M.A., F.R.S., Hope Professor of Zoology in the
? University of Oxford.

| See quite recent years the method of setting butterflies

and moths adopted in this country differed very widely
from that which prevailed and still prevails on the Continent.
Furthermore, the continental method has been uniform, while
the British has varied in many directions, according to the
taste of different distinguished collectors, each of whom may
be said to have founded a fashion which endured for a time
and over a more or less limited area.

In continental setting very long (1$ inches or more, and
inconyeniently flexible for use with a cork lining to the drawer)
foreign pins are used ; the insect being pierced so that the pin is
as nearly as possible upright. The insect is raised on the pin so
that a length of only from 3, to } of an inch projects above
the dorsal surface of the thorax. The wings are extended on
flat boards, the object being to render them as horizontal as
possible. ‘The margin of the fore wing which overlaps the hind
wing (inner margin) is placed at a right angle to the long axis
of the body, and therefore in a line with the corresponding
part of the opposite fore wing. This arrangement involves, in
the great majority of Lepidoptera, a strained or unnatural
appearance, owing to the fore wing being drawn so far

forward—an effect which is sometimes increased by drawing

eed

the hind wings backward so as to leave a considerable gap
between them and the fore wings.

It has been already stated that there is no one British
method, but a great variety, differing widely from each other and
from the continental. The pins used are of British make and
superior in stiffness to those supplied by the continental makers,
There is no attempt at uniformity in length, but the size of the
pin is regulated by that of the insect. The insect is pierced so
that the pin slopes forward, but there is no uniform or standard
slope; and, indeed, such a standard would be far more difficult to
attain than under the continental method, in which the upright
pin is required. Even the longest pins (13 inches) used are
inferior in length to the continental pins. In the older collec-
tions set in England the insect was placed very low on the pin, so
low in fact that the edges of the wings (hind margins), which
were made to slope downwards, rested on the paper lining of the
cabinet drawer. The obvious danger and injury involved in
this position—the tendency to curl up—the ready accumulation
of dust—the convenient access afforded to mites, etc.—the
want of a space on the pin below the insect for the attach-
ment of labels—gradually caused its abandonment; so that, in
most modern English collections, the insects are fixed higher
on the pin; but here again there is no standard, some collectors
being satisfied if the insect be just clear of the bottom of the
drawer, others raising it until only from ,%, to } inch of pin
projects above it, as in the continental method, others again
adopting an intermediate position. The wings are set less
forward than in the latter method, and care is exercised to
preserve the natural amount of overlap of the fore over the
hind wing. Instead of the flat setting the aim has been to
make the wings slope downwards away from the body of the
insect, and to increase the slight, natural curvature of the
insect’s wings. This is usually accomplished by curved
setting-boards, but it has been brought about by means of
cardboard slips placed de/ow the central part of the wings,
together with other slips adove their outer part, the elasticity

ese]
of the wings keeping them in place. It has even been
suggested that a curvature from back to front in addition to
that from side to side would be peculiarly elegant, and collec-
tors have been advised to make setting-boards out of portions
cut from a sphere.

Apart from the relative advantages and disadvantages offered
by these methods, the fact of the difference is much to be
regretted. One of the most serious difficulties which have
been brought about is the want of uniformity in the depth
of the drawers of British and continental cabinets; and _ here,
too, the want of a standard has produced inconvenient results
within the limits of this country—the British cabinets varying
from an effective depth from glass to paper of 14 inches
to one of 134 inches, as against a depth of 14 inches
in the continental drawers. This is, of course, excessively
inconvenient when any large exchange of specimens takes
place. In fact, it is not too much to say that the uniformity of
the continental method would render its adoption expedient even
if it were less suitable in other respects. But a careful com-
parison of all the various points will show that in almost all
points it is superior to the English method. I will take the
various points separately.

Choice of Pin.—The largest English pin (No. 16 or No. 11,
with a length of 12 inches) is stiffer and more convenient than
the continental article, and is sufficiently long to permit the
advantages mentioned in the paragraph below. All insects,
large and small, should be mounted so that they are at an
equal distance from the glass.

Height of Insect on the Pin.—The immense advantages of
a high position have been already mentioned: furthermore,
it may be added that it is of advantage to bring the insect
near to the glass, so that it can be examined with a lens
without removing the cover. The high position also enables
the observer to read two or three labels placed at different
heights on the pin below the insect without disturbing them
or the insect, and even without removing the cover of the

tae

drawer. In these days of careful cataloguing, and full and
accurate data, this abundant space for small labels is of the
utmost importance. It should be noted that the proximity of
the insects to the glass necessitates care in the removal of the
covers, because of the injury to the wings which is apt to be
caused by a sudden rush of air.

Direction of the Fin.—The upright position has the great
advantage that it favours uniformity, which, on the other hand,
is very difficult to secure over a wide area with the sloping
position. The latter method, furthermore, tends to hide the
labels on the pin, inasmuch as it brings them further under
the body of the insect.

Plane of the Wings.—The flat position also favours uniformity,
whereas the slope always tends to vary according to the taste of
the collector. Another, and perhaps the chief, advantage is that
the flat wings being at right angles to the line of vision are more
clearly and distinctly seen. To see the sloping wings perfectly
the insect must be turned first to one side and then to the other,
and doth sides cannot be seen perfectly at the same time. The
flat continental boards do not, as a rule, produce perfect
flatness because the wings are naturally curved a little, and
this requires special correction. Mr. H. J. Elwes has devised
boards, now much used in this country, which are curved
slightly in the opposite direction, viz., upwards, and these are
very effective in producing the desired flatness. ‘The wings are
usually held in place by strips of engineers’ tracing cloth secured
by the ordinary milliners’ steel, glass-headed pins, which can
be easily driven into soft wood.

Forward direction of the Wings.—Here, too, the continental
method of placing the inner margin of the fore wings at nght
angles to the body has the great advantage of uniformity,
and when the setting is performed mechanically, with no
knowledge of or interest in the insects themselves, it would
probably be the most expedient system to adopt. If, however,
the setter has something of the feeling of a naturalist and
artist, far more pleasing results may be obtained by placing

regal

the wings in as natural a position as is consistent with their
proper display, the latter being, of course, the paramount con-
sideration. Some judgment will be necessary in dealing with
insects of which the habits are unknown. I think it may be
assumed that the natural amount of overlap—easily distin-
guished by the characteristic texture and appearance of that
part of the hind wings which is naturally concealed—should
always be retained.

The feeling in favour of this or that form of setting, apart
from the considerations of utility, is largely, if not entirely,
a question of fashion. As a boy I looked on the continental
setting with horror; now the sight of the sloping wings in
contact with the paper gives me a cold chill. It is probably
the case that a very large number of our collectors are still
strongly in favour of some form of the English method. ‘To
these I would urge the advantage of uniformity, and would
point out that the method they still follow is disappearing
with all possible speed. The British Museum, the Godman-
Salvin Collection, Mr. Herbert Druce’s Collection, and the
Hope Collection are all being rapidly transformed in the
direction I have indicated. The method was first shown
on a large scale in this country in the fine collection of
Mr. Elwes. Whatever our preferences may be, the question
is being rapidly settled on grounds of expediency, and
the sooner the change is effected the better it will be for
everybody.

Before concluding I must say a few words about labelling.
The most important fact is, of course, the locality, and this
should be precise and exact: next in importance is the date
of capture; next the name of captor; then the date of
presentation and the name of donor. Of some _ interest
for future workers is a statement as to the authorship of
the manuscript label. As regards locality, at Oxford we
make a point of printing the most important word first and
in capitals; then follows the more minute locality and the
height above the sea, if attainable. Examples of the labels

tea

of specimens presented to the Hope Collection in 1897 are
printed below :-—

In the much rarer cases in which notes of experiments or of
natural history observations accompany the specimens, these are
incorporated in an epigrammatic form in the label. Examples
are given below :—

NE DURBAN, NATAL, es
Until, NE DUEBAN, NATAL,
= 5 feet |
One of 12 Exrals : ——
mis asd wabibes | 6d 20h Imsgo,
at on fern breed April 20, 1897.
34 px. June & V7. | aad pres. i297 by
= a Guy A K. Marshall
by @ AK Marshall =ayGaAE™M |
mabGAEK M

As regards date of capture the importance of the day of the
month is at once seen in relation to the recent work upon
seasonal forms and as giving a solid fact in the natural history
of the species. Many excellent collectors neglect the date
altogether. This is a serious error; for the study of the
gradual changes in the distribution of forms will demand these
very data from the specimens now being added to our
Museums. To take a concrete case. Mypolimnas muisippus
has of late years invaded America, and the dates of its
arrival and gradual spread are of very great interest. There
is a Godman-Salvin specimen in the British Museum captured
by Belt in the Montes Aureos, Maranhdo, Brazil; but without
a date. In this case, no doubt, the date can be fixed approxi-
mately by indirect means, although I have not as yet been
able to ascertain it; but in many other cases it cannot be
recovered. What a splendid history of the gradual spread of
Anosia plexippus from its home in North America could be
deciphered from our great collections if only the collectors had
troubled to put dates on their captures. When labels are
printed it is almost as easy to print two or three as one only.
It is of great advantage to pin a second label Jdeside the
specimen where it can be read with the greatest ease, while

a sal

a third may be found useful for cataloguing. Diamond type
will be found the most satisfactory for this purpose, while a

small and simple press, which permits easy and rapid alteration
of type, is the most convenient.

Mr. PLATNAUER asked if anything could be done to the pin to
render it more difficult for the predaceous intruder to climbit. He
also questioned the wisdom of putting labels, and especially the
original or collectors label, on the pin under the insect. Not
only did it make the collection more unsightly, but it exposed
the label to the risk of sharing the fate of the specimen. He
would suggest that the specimen bear a number only, and that
this number refer to a catalogue which could contain full references,
and into which the original label could be pasted. In this case,
everything would not be lost at one blow; even if the specimen
were destroyed, the reference would remain.

Dr. PETRIE asked whether labels or pins could not be treated
with protective substances such as white arsenic or corrosive sub-
limate and glycerine.

ee 2 eae

Mr. POULTON said that the labels under the pin formed an
excellent barrier against intruders. The use of strong antiseptic

; or protective substances, such as arsenic and corrosive sublimate,
was objectionable on account of their tendency to make the pin
; brittle. The question of pins was important enough to constitute

a separate subject for consideration ; he himself knew of no satis-
factory substitute for the pins ordinarily used by entomologists,
| although some authorities favoured platinum, some silver, and
some steel pins. The best way to preserve Lepidoptera was to
keep the drawers containing them thoroughly disinfected with
a good antiseptic. Nothing was better adapted for this purpose
than naphthalene. As to putting labels into a catalogue, the objection
seemed to him that this exposed the collection to the risk of total
| loss of all references by one mishap. If the catalogue were lost
or destroyed, a// information with regard to the specimens would
be gone, whereas, with separate labels on the specimens, there
was not nearly the same chance of an extensive loss. He instanced
the Burchell collection of South American Lepidoptera which was
Fi almost useless in consequence of the loss of its catalogue. [Since
the above was written this catalogue has been recovered, but it is
impossible at present to decide whether it is complete.—E. B. P.,
1898. ]

iy
tas, iy
Lik ye

7 "
et,

Fune, 1898.

Report of the Hope Professor of Zoology.

The year 1897 was marked by very large accessions to the
Hope Collections, chiefly of specimens obtained from Canada
and some of the adjacent States (Illinois and Wisconsin).
The British Collections were also very largely increased.
Mr. G. A. K. Marshall’s gift of South African butterflies,
described in detail below, must also be specially mentioned.

Much work has been done in labelling the Godman-Salvin
specimens and classifying the existing manuscript labels
for printing. The W. W. Saunders Collection of Sphingidae
have received printed labels giving the geographical data and
the historical account of the specimens. An example of the
form which has been adopted is reproduced below.

)ssSse Se
| NEW ZEALAND |
Cape Campb ell
Ex Coll. (1830-73)
W. W. Saun Jers.
Presented 1873 by |
Mrs, F. W. Ho ype.
ms. locality by
W.W.S.

|

Similar labels have been added to a part of some other
groups and will eventually be extended to the whole of the
historic collection of moths which the University owes to the
generosity of Mrs. F. W. Hope.

In March, Mr. H. R. Smith, the Junior Assistant, left this
country for Canada. Before leaving he presented a large
number of insects for the British Collections, and he has since
sent a number captured in Canada.

After waiting a few months this place was filled by the
appointment of Mr. A. H. Hamm of Reading. Three-fourths
of his time is devoted to the Hope Department, the remaining
fourth being occupied by the Delegates in printing for the
Museum. His experience as a printer has proved of great

B

2

value in the immense amount of labelling required by the
Department.

At the close of the year the thorough revision of the Hope
Library was commenced and has occupied Mr. Holland’s time
ever since.

The Danaine butterflies, with the exception of the /thomiinae
(Neotropinae), have been finally arranged by Mr. Holland in
two of the new cabinets. In this group, so far as it is at
present arranged, the Godman-Salvin specimens are incor-
porated, and the whole re-set according to modern principles,
with coloured labels to show the geographical distribution of
the species and genera, and with printed locality labels to
a large proportion of the specimens, constituting an extremely
instructive and valuable Collection. It is much to be regretted
that the imperative necessity for classifying the Library has
for a time interrupted the progress of this work.

In the course of this arrangement and in the preliminary
work upon other groups preparatory to their final arrange-
ment, it has been necessary frequently to consult the Insect
Department of the National Collection and the Godman-
Salvin Collection. In this work the greatest courtesy has
been shown, and much kind help afforded.

The Department has, as in previous years, received much
invaluable help in the determination and arrangement of the
specimens.

Mr. M. Jacoby has continued his kind assistance in working
out the large Collection of Phytophaga (Coleoptera). Dr.
Dixey has worked at the Pzerzvae preparatory to their final
arrangement. Col. Swinhoe has continued and has nearly
completed the arrangement of the Oriental Heterocera ;
Mr. Arthur Sidgwick and Mr. Pogson Smith the arrange-
ment of the British Heterocera; the Macro-Lepidoptera and
the Deltoids being now finished and contained in about 100
drawers in two cabinets. Mr. Malcolm Burr has done much
work upon the Collections of Forficulidae and Acridiidae.
Miss Cecil Vernon Harcourt has prepared the Collection of
British Aculeate Hymenoptera for final arrangement, printed
labels for the specimens, and has begun the arrangement.

3

Mr. G. C. Champion has worked out the 77izg7tidae (Rhynchota
Heteroptera) in the Department and described one new genus
and three species (Trans. Ent. Soc. Lond. 1898, p. 55).

Several distinguished entomologists have visited the Hope
Collection in the course of the year, and have given much
needed advice and assistance as regards the groups which they
have specially studied. Col. Bingham made a rapid survey
of the very large general Collection of Aculeate Hymenoptera,
Mr. Edward Saunders of the Palaearctic Aculeate Hymen-
optera. Mr. Alexander Fry gave valuable information about
the Miers Collection. Dr. F. Moore, after visiting the Depart-
ment, worked through the Eastern species of Vefzzs and allied
genera of butterflies. Mr. R. H. F. Rippon examined the
butterflies of the genus Orzzthoptera, and has since worked
out the more obscure forms. Mr. W. F. Kirby identified
many types of Homoptera.

The Council of the Entomological Society were invited to
visit Oxford, July 3-5. The President, Mr. Roland Trimen,
F.R.S., the Treasurer, Mr. R. McLachlan, F.R.S., the Secretary,
Mr. W. F. H. Blandford, Professor Meldola, F.R.S., Mr. O.
Salvin, F.R.S., Sir George F. Hampson, and Mr. M. Jacoby
were present, while Mr. F. D. Godman, F.R.S.,and Mr. Herbert
Druce also joined the members of Council and the Oxford
entomologists. Many friends of the Department kindly
assisted in entertaining our visitors. The Hope Collection
has greatly benefited by the experience of these distinguished
authorities in so many groups of insects.

The state of the fund for the purchase of new cabinets
which are so greatly needed by the Department was shown in
the Report of 1896. It has now been raised to over £400,
by 410 kindly subscribed by the Warden of All Souls.

New cabinets containing 200 drawers were delivered in
November, and another equal consignment has since arrived
(March 1898). These 400 drawers have been excellently
made by O. E. Janson and Son, and are in every way
satisfactory. The fund is now exhausted, but it is much
to be hoped that £200 for another consignment may be
raised at least six months before the arrangement of these

B 2

4.

400 drawers is complete; for it has been found that at least
this period must elapse between the order and delivery.

The first volume of the Hope Reports appeared on
June 14. This volume contains copies of the various
original memoirs which were written in connexion with the
Hope Department between June 1893 and June 1897. During
these four years much time and attention has been devoted
to securing increased space and many necessary conveniences
for work in the Department; so that it is confidently to be
hoped that the succeeding volumes will appear at much
shorter intervals. Indeed the materials for Volume II are
already accumulating rapidly. The volume has been or will
be distributed to Hope Curators, Delegates of the Museum,
College and University Libraries, and to the majority of those
interested in the progress of Science in Oxford. Outside
Oxford they are sent to the chief Scientific Libraries of
England, the Continent and America.

Volume I contains 16 memoirs by W. Garstang, M.A.,
W..Schaus, F.Z.S., F.A. Dixey, M.A: M.D. and Professor
Poulton, M.A., F.R.S.

ADDITIONS TO THE COLLECTION.

Fifty-six specimens of butterflies, captured April 5, 18y6,
near Durban, Natal, by D. Chaplin, Esq., were presented by
F..D., Godman,, Msq:, and ©. Salvin, Esq:) (Many sommnese
are of especial value in providing material for the separate
series illustrating Mimicry and Common Warning Colours.

A Dipterous insect from the Isle of Wight was presented
byab.S. Ocles Bsa:

A very valuable set of 13 Forficulidae from Lombok,
W. Java, and S. Celebes, and 3 Orthoptera from Surinam and
W. Java, was presented by Malcolm Burr, Esq. (New
College, Oxford), together with 14 Neuroptera captured by
him at Esher.

A fine set of larvae of Vanessa urticae, var. ichnusa, was
brought from Corsica by H. C. Playne, Esq., M.A. (Univer-
sity College), and bred in the Department together with one

eee

5

Vanessa atalanta. Of the former 13 selected specimens were
added to the Collection. Furthermore 27 specimens of insects
of various orders from N.W. Finland were presented by H. C.
Playne, Esq., and A. F. R. Wollaston, Esq.

Thirty-one specimens of Lepidoptera from the Italian Alps
were presented by H. M. Wallis, Esq.

Two Rhopalocera from the shores of Lake Erie, Ontario,
were presented by Dr. G. S. Ryerson of Toronto.

A moth and a beetle, bearing a close resemblance to each
other, both captured in June 1897, by H. G. Hubbard, Esq.,
in the Chiricahua Mountains, Arizona, were presented by
Dr. L. O. Howard of Washington. These will prove a
valuable addition to the series on Mimicry, &c.

Two Rhopalocera from Corsica, and 8 from Civita Vecchia,
were presented by A. B. Trevor-Battye, Esq.

A large and valuable collection of 215 Rhopalocera cap-
tured in Trinidad about 1874, was presented by Prof. R.
Meldola, F.R.S. The collection includes a specimen of
Hypolimnas misippus, thus proving that this oriental species,
now established in many of the W. Indian Islands and in
parts of tropical America, had reached Trinidad by or before
1874. Several specimens from this Collection will be included
in the separate series on Mimicry.

A collection of 119 specimens of Lepidoptera from Deme-
tara was presented by A. Waterfield, Esq. In this collection
the same species (//. mzsippus) is included, and Mr. Waterfield
informs me that he has observed its gradual increase during
recent years.

Nineteen Lepidoptera from Tasmania were presented by
M. Jacoby, Esq.

A specimen of Limnas chrysippus from Greece was pre-
sented by Rev. G. D. Allen, B.A.

Eight specimens of 2 species of Forficulidae, new to
Britain, from Queenborough, Kent, together with a rare
Coleopterous insect from Chatham, were presented by J. J.
Walker, Esq., R.N.

Fourteen Lepidoptera from Mercedes, B.A., Argentina,
were presented by Rev. F. D. Tubbs.

8

6

A set of 5 Avraschnia prorsa-levana, bred from European
pupae, was presented by Dr. F. A. Dixey.

A very fine set of 78 Lepidoptera from the neighbourhood
of Brisbane, 31 from Ecuador, and 1 from Colombia, was
presented by G. C. Griffiths, Esq.

A fine set of 80 Lepidoptera and 1 Coleopteron from the
neighbourhood of Oaxaca City, Mexico (6,100 ft.), was
presented by Osbert H. Howarth, Esq.

Further invaluable additions to our series of African
butterflies were made, as in last year, by Guy A. K.
Marshall, Esq. The collection shows the same accurate
labelling which renders it so especially valuable to us. The
collection contains 416 Rhopalocera, 2 Heterocera, and 2
Ichneumonidae from various localities (all precisely recorded,
and now copied on printed labels) in Natal; and 143 Rhopa-
locera from various localities in Mashonaland.

In addition to these important accessions, Mr. Marshall
has presented several groups of the utmost interest and value
for the series on Mimicry, &c. Some of these captures
prove that the species which resemble each other in the same
locality also occur at the same time of the year. Thus the
Hope Collection has received 5 specimens of butterflies of
the group which resembles Amauris echeria, all captured at
Malvern, Natal, on March 6, 1897; 8 butterflies of the Lzmnas
chrysippus type of pattern and colouring, captured at Malvern
on the same date, and again 6 of this latter group on March
30. Also 5 specimens of butterflies and moths which appear
to resemble one of the black and white species of Amauris,
probably A. ochlea, captured at Malvern on March 27. Thirty-
three other specimens in part illustrated the same principle,
in part certain interesting observations on the habits of
Euralia mima and E£. wahlbergt, and in part formed the
material of some important experiments on the effect (on
the perfect insect) of dry heat and moisture upon the pupae
of various species of butterflies. A detailed account of these
experiments will be published shortly.

The absence of Professor Poulton in America, from the
middle of July to the middle of September, caused an

rf

‘immense accession of specimens captured by him. and the
friends who accompanied him, and also kindly presented by
many naturalists who reside in Canada and the United States.
The work thrown on the Department in setting, printing, and
cataloguing, has consequently been so excessive that it has
been found impossible to include a complete account of these
and many other additions in the present Report. A brief
account will now be given, to be followed next year by
a complete description giving the numbers of specimens.

Professor Poulton’s captures (of insects of nearly all orders)
were chiefly made at Quebec, Montreal, Toronto, Chicago,
Detroit, Niagara Falls, Hartland (Wisconsin), Rat Portage,
Rush Lake, Regina, Lake Louise (Laggan), Glacier House
(Selkirks), Vancouver City, and Victoria (Vancouver Island).
Small numbers of captures were also made at a very large
number of localities on the Canadian Pacific Railway.

Professor Meldola, F.R.S., also captured and presented
a very large number of specimens from the Canadian
localities.

In Chicago, Professor Wheeler and Professor Watasé cap-
tured many specimens for the Hope Department, and the
former also presented some valuable specimens from his own
collection for the series on Mimicry, &c. In Hartland,
Professor Poulton was kindly helped by Dr. G. W. Peckham,
who also presented specimens, chiefly of Aculeate Hymen-
optera, he had previously captured.

At Victoria he received much kind assistance from Dr. E.
Crompton (of Victoria), from E. A. Carew-Gibson, Esq. (of
Victoria), from Dr. Kirker (of Esquimalt), and from Professor
Fitzgerald, F.R.S. Furthermore a fine collection: of insects,
chiefly Lepidoptera, from the neighbourhood of Victoria was
presented by the captors, Dr. E. Crompton and F, Crompton,
Esq. These will be of the greatest value to the Hope Col-
lection. A small set of insects from the neighbourhood of
Victoria was presented by E. Baynes Reed, Esq., of Victoria,
and another from Vancouver City, by C. A. Ellis, Esq., of
Vancouver,

Valuable collections from Ottawa and from Virden (Mani-

8

toba), were made during the Summer and Autumn of the year
by Miss M. G. Holmes and presented to the Department.
The data accompanying the specimens were unusually full
and precise. Mr. H. R. Smith has also sent a large collection
from the neighbourhood of Russell, Manitoba.

A small but very valuable collection from the far north—
Hudson’s Strait—was presented by the captor, F. F. Payne,
E'sq., of Toronto. Miss E. C. Ramsay and W. Ramsay also
presented many specimens captured by them in Montana,
U.S. A., and at localities on the Canadian Pacific Railway.

Specimens from Canadian localities were also presented by
FE. H. Chapman, Esq., B.A. (Magdalen College).

From this preliminary list of presentations it will be seen
that the North American insects of the Department have
been very greatly augmented during 1897. This addition will
be especially welcome, as the insect fauna of this interesting
region is poorly represented in the University Collection,
while the specimens we already possess are very poorly and
insufficiently labelled. The large collection now added is
especially strong in this respect.

The Professor also desires to acknowledge the kindly
interest and sympathy which he everywhere received through-
out the whole of his visit, in his efforts to add to the Hope
Collection.

The extra strain thrown on the Department has also pre-
vented the cataloguing of many other valuable accessions
from other sources. These, too, will now be briefly men-
tioned, the full record being deferred till next year.

A small but very valuable collection of insects, made in
Sokotra in the winter of 1896-7, by E. N, Bennett, Esq. (of
Hertford College, Oxford), was presented by him. Although
the insects are in poor condition they are of great interest,
and include 2 new species of Rhopalocera, 1 of Orthoptera,
and 4 of Arachnida, which have since been described.

Colonel Swinhoe has presented a very large number of
Heterocera, many Rhopalocera, especially Pierinae, and also
some specimens for the series on Mimicry, &c.

A valuable collection of insects of many orders captured

‘
||
Bh

9°

upon two visits to Somaliland by C. V. A. Peel, Esq., was
presented by him. Several new species are probably in-
cluded, which will be worked out as soon as_ possible.
Mr. Peel also presented a few Rhopalocera from various
localities.

A large collection of Lepidoptera from many localities
has been presented by H. Druce, Esq. Many of these
will be of the highest value to the Hope Collection.

A set of insects, chiefly Lepidoptera, from British Guiana,
has been presented by Rev. O. Pickard-Cambridge, F.R.S.

Several European Rhopalocera and Coleoptera have been
presented by Rev. G. D. Allen.

A set of Swiss Lepidoptera was presented by Mrs. Bassett.

A few Rhopalocera, especially needed by the Collection on
account of their rarity or the localities at which they were
captured, were presented by Mr. A. H. Hamm.

The British Collections have been greatly strengthened
during the year by very large accessions, with excellent
data, presented by Mr. W. Holland, Mr. H. R. Smith, and
Mr. A. H. Hamm. The specimens are principally Lepi-
doptera, but other orders are also represented; the speci-
mens were principally captured in the neighbourhood of
Oxford and Reading.

Dr. F. A. Dixey presented a valuable set of insects of many
orders captured by him at Morthoe, N. Devon. The data
accompanying the specimens are excellent.

Miss Thoyts presented a valuable set of Aculeate Hymen-
optera captured at Sulhampstead, Reading, the data being
especially complete. British specimens of the same order
were presented by Edward Saunders, Esq., from his col-
lection,

The British Collections were also increased by specimens
presented by Arthur Sidgwick, Esq. (Lepidoptera), Miss
C. V. Butler (a specimen of the rare moth, Drymonia chaonia,
captured near Oxford), Professor Poulton (chiefly Lepidop-
tera from Freshwater, Isle of Wight), and his son, Edward
P. Poulton (two rare moths from St. Helen’s, Isle of Wight).

A moderate number of specimens, chiefly intended for the

10

Series on Mimicry, &c., were purchased from Watkins and
Doncaster.

In addition to the gifts fully recorded or briefly mentioned
above, the Hope Department has received from Canon
Tristram, F.R.S., a valuable set of 291 insects of many
orders from Palestine. The history of the collection is
remarkable. These insects, with others, occupied four
drawers in the Department which were labelled Palestine.
They probably reached Oxford about 30 years ago, but
nothing was known of them except the statements in Professor
Westwood’s handwriting that all came from Palestine and
that a part had been captured by the Rev. O. Pickard-
Cambridge. There was also a label on one drawer with
these words, “ Zzzeae named by Stainton,” and on another,
“Oxford and Cambridge Exploration Fund. Hymenoptera
to F. Smith.’ It was known that Professor Westwood was
very careful of the insects and would never allow them to
be placed in the general collection. Enquiry not only failed
to ascertain the existence of an “Oxford and Cambridge
Exploration Fund,” but showed that these insects were
certainly unconnected with any fund of the kind. <A _ label
from one of the Z7zezza was then sent to Lord Walsingham,
who not only identified it as the handwriting of the late H. T.
Stainton, but also as the label of a long-lost type described
in “The Tineina of Syria and Asia Minor” (London, 1868).
Lord Walsingham furthermore stated that all these 7zneina
had been given to him by Canon Tristram, and the Canon
himself wrote in confirmation. The matter was brought
before the Hope Curators and the Professor was authorized
to forward the 56 specimens (36 species and 31 types and
co-types) of Zizeiwa to Lord Walsingham. This result
did not encourage further researches into the history of the
collection. Nevertheless Canon Tristram was written to
about the remaining specimens, and he explained that
Professor Westwood had borrowed the whole collection
from him for the purposes of study. The specimens had
been principally captured by Rev. O. Pickard-Cambridge,
F.R.S., in 1865, but a large number by Canon Tristram

it

himself in 1863-4, and the whole collection was undoubtedly
his property. He has now very kindly presented the whole
of them to the Hope Department in which they have rested
so long.

MODITIONS TO" THE HOPE LIBRARY.

The Smithsonian Institution (United States National Mu-
seum) presented their publications during the year, which
dealt with the subjects of the Hope Department.

The Boston Society of Natural History and the Bombay
Natural History Society presented their publications for the
year.

The University of the State of New York presented three
volumes of their Report, 48 (1, 2, and 3).

The Secretary of State for India in Council presented the four
volumes on “ The Moths of India” by Sir George F. Hampson.

The Director of the Royal Indian Marine presented two
parts of “Illustrations of the Zoology of the Royal Indian
Marine Surveying Steamer Investigator.”

The Linacre Professor of Comparative Anatomy, Oxford,
presented “ The Linacre Reports,” vol. ii.

“ The Hope Reports,” vol. i, was deposited in the Library.

The Radcliffe Librarian, Oxford, presented the Catalogue
of Books added during 1896.

The Transactions of the Entomological Society, and the
Transactions and Journal of the Linnean Society for the
year 1897, were presented by Professor Poulton.

Canon W. W. Fowler presented the parts of his monograph
in the Biologia Centrali-Americana (Rhynchota Homoptera,
vol. ii), published during the year.

Copies of original papers were also presented by the fol-
lowing authors :—Dr. L. O. Howard, Sir George F. Hampson,
Malcolm Burr, Esq., Hamilton Druce, Esq., F. T. Howard,
Esq.,and T. H. Thomas, Esq., Miss Eleanor A. Ormerod, Baron
Osten Sacken, Herbert Druce, Esq., Rev. T. R. R. Stebbing,
Games Carpenter, Esq. PR: Ps Calverty Esq: and. Gy WwW:

12

Kirkaldy, Esq. Two plates of “uxploetnae were presented
by Ff. -G- Moore; Esq:

An extremely valuable copy of W. J. Burchell’s “ Travels
in the Interior of Southern Africa” (London, 1822), containing
MS. notes and corrections by the author, was presented by
J. P. Mansel Weale, Esq. The presence of the Burchell Col-
lection in the Hope Department renders this gift especially
welcome.

Several second-hand works of reference and original papers
were purchased during the year, together with the Zoological
Record, the parts of Barrett’s “ British Lepidoptera,” the Ray
Society volume for the year, and the parts of Rippon’s “ Icones
Ornithopterarum.”

EDWARD B. POULTON.

Report of the Hope Professor of Zoology.

The additions to the Hope Department made during the
year 1898 are numerous and valuable, although they do not
approach those of 1897. Mr. Guy A. K. Marshall’s beautifully
collected specimens from South Africa, accompanied by full
and precise data, must be speciaily mentioned, as in previous
years.

One of the principal labours of the year was a thorough
revision of the Hope Library and comparison of the volumes
with the (very incomplete) slip catalogue. The thousands of
works which require cataloguing were rendered easily recog-
nizable, while the numerous duplicate volumes and pamphlets
were identified and separated. At this point the work was
postponed owing to the needs of the Collection, but the com-
pletion of the catalogue at an early date is absolutely necessary
for the assured safety and proper and convenient use of this
valuable library. It is much to be hoped that some means
may be found whereby this work can be done without en-
croaching further upon the amount of time and work devoted
to the Collections. Mr. W. Holland commenced this revision
in 1897, and it occupied almost the whole of his time for the
first half of 1898. Mr. R. McLachlan, F.R.S., kindly assisted
in the classification of some of the more obscure pamphlets,
and Miss Cora B. Sanders, of Lady Margaret Hall, in writing
a number of slips.

In the latter part of the summer Mr. Holland arranged the
large group of Danaine butterflies confined to tropical America
and now separated as a distinct sub-family, the /thomiinae.
The immense growth of the Collection is well seen in this
group. The whole of the specimens in the Department in
1893 were crowded into two drawers, and might have been
expanded into five or six after the species had been worked
out. Now that the Godman-Salvin and other accessions of
less importance have been incorporated, with a fair allowance
for future additions, the sub-family occupies sixty drawers.

2

During the remainder of the year Mr. Holland began to
arrange the first part of the collection of Phytophaga (Coleo-
ptera), which has been examined and named by Mr. M. Jacoby.
A considerable part of this work has now been done, and early
in 1899 the arrangement of the general collection of butterflies
was resumed, that of the Satyrznae being already complete
in 1co drawers, while the Elymniinae, Amathusiinae, and
Morphiinae are worked out ready for the final arrangement.
Making some allowance of space—viz. 40 drawers—for the
Pierinae now being arranged by Dr. Dixey, the above-men-
tioned groups will, with those already arranged, occupy the
whole of the 400 drawers recently ordered and delivered in
November 1897 and March 1898. It is of the utmost impor-
tance that a further consignment, if possible of equal size,
should be ordered forthwith, and during the six months which
must elapse before the order is complete the final arrange-
ment of the butterflies must be postponed and that of some
other group undertaken.

Mr. A. H. Hamm has been chiefly occupied in setting and
in printing labels for the additions to the Collection, especially
the large number of specimens (over 3,000) brought by Prof.
Poulton from Canada and the United States in September
1897. These specimens were captured, generally a few at
a time, in so many different localities that the labour involved
in labelling them has been very severe. He has also reset
the butterflies, which have been arranged by Dr. Dixey and
Mr. Holland in the general Collection, so far as they required
it. The work upon the Pzerdvae arranged by Dr. Dixey has
been especially troublesome, because of the flimsy condition
of many of the older specimens.

Towards the latter part of the year Mr. Hamm arranged
the Collection of British Hymenoptera Aculeata (ants, bees,
wasps, &c.), which had been kindly named by Mr. Edward
Saunders. A large amount of preliminary work upon these
insects had been done by Miss Cecil Vernon Harcourt in the
previous years. The Collection occupies 30 drawers, and is
now in a most efficient state.

In addition to his work for the Department the Delegates
employ one-fourth of Mr. Hamm’s time in printing for the

3

Museum, and the Hope Collections have greatly benefited by
the share which has been devoted to them. At these times
labels in large type have been printed on the Museum press,
and are now placed in the card-holders on the outside of
cabinets and of drawers, thus facilitating reference to the
Collections. This only applies to the parts of the Collection
which have been finally arranged.

The immense additions made in 1897 have occupied so
much time in setting, printing, labelling, cataloguing and incor-
porating, that those of 1898 and 1899 have necessarily been
kept back. Even at the present date (June 1899) the 1897
additions are not all finished. In the meantime those parts of
the additions of 1898 and 1899, which were specially needed
in relation to the work which has been going on, have been
labelled and incorporated. In spite of the great strain thrown
upon our small staff by the 1897 accessions, I hope that we shall
nearly have overtaken our arrears by the end of this year.

The Department has been greatly helped as in previous
years by the kindness of friends. Col. Swinhoe has continued
his work upon the Eastern moths. Dr. Dixey has continued
the arrangement of the Pzerzzac, and has worked out many
questions of the deepest interest, especially as regards the
seasonal differences in many species, and the transitional forms
which he has shown to occur connecting species which were
believed to be distinct. Mr. Arthur Sidgwick and Mr. Pogson
Smith have now arranged the British Lepidoptera as far as
the middle of the Tortrices. Mr. M. Jacoby has continued
his valuable examination of our collection of Phytophaga.
Mr. Malcolm Burr (New College) has worked from time to
time in the Department, and has often rendered kind assistance
in the Orthoptera. Col. J. W. Yerbury visited the Department
and rapidly surveyed the general collection of Diptera, upon
which he wrote a very useful report, pointing out the groups
in which the Hope Collections are especially strong, and giving
a list of many of the important types which we possess.

Several distinguished entomologists visited the Collections
in order to examine specimens in the groups in which they
are authorities. Such work always does great service to the
University in the better understanding of its Collections.

4

Dr. W. Horn of Berlin came to work at the Crczndelidae,
Dr. Roeschke of Berlin at the Caraéidae. Professor Roland H.
Thaxter of Harvard University remained in Oxford for several
days in order to work at the Ladoulbeniaceae, a group of
minute fungi which are parasitic upon certain beetles, &c.
Mr. W. J. Lucas examined the British Odonata (Dragon-flies).

Many other eminent visitors paid a brief visit to the
Department. Among these were Prof. Mitsukuri of Tokyo,
Dr. Karl Jordan, H. Frithstorfer, Prof. Haswell, René Oberthiir,
Albert Fauvel, Aug. Lameere, and Rev. O. Pickard-Cam-
bridge, F.R:S:

The annual visit of the Council of the Entomological
Society took place on July 2-4. The President, Mr. Roland
Trimen, Hon. M.A., F.R.S., the Treasurer, Mr. R. McLachlan,
F.R.S., the Secretary, Mr. F. Merrifield, Mr. C. O. Waterhouse,
Mr. M. Jacoby, Mr. J. W. Tutt, and Mr. A. H. Jones were
present, and, as in the past, the Collections greatly benefited
from the experience of our visitors. The members of the
Council were entertained by Hope Curators and others
interested in the Department. By the death of Mr. Osbert
Salvin, F.R.S., who was with us in 1896 and 1897, and had
been expected in 1898, the Hope Department has sustained
a grievous loss. Mr. Salvin was not only a very generous
donor to the Hope Department; we relied continually upon
his deep knowledge, and immense experience in dealing with
Natural History Collections.

I must again thankfully acknowledge the invariable kindness
received in the Insect Department of the Natural History
Museum and the Godman-Salvin Collection.

A considerable amount of scientific research has been
published in the course of the year, describing work done
in the Department or in some cases elsewhere upon material
belonging to the Department.

The insects collected by Mr. E. N. Bennett, M.A., in Socotra

(December 17, 1896, to February 11, 1897), were worked out :
the Lepidoptera by Dr. F. A. Dixey, who described 2 new
species, Papilio bennetti and Lyblia boydi, the Orthoptera by
Mr. Malcolm Burr, who described ft new species, the
Arachnida by Rev. O. Pickard-Cambridge, who described

SS Sa

5

4 new species, The paper appeared in Proc. Zool. Soc.,
1898, p. 172.

Mr. Herbert Druce described 23 new species of South
American moths (Syztomidae) in the Hope Collections (Ann.
and Mag. of Nat. Hist., 1898, p. 401).

Dr. F. A. Dixey wrote on recent experiments on the
hybridization of insects (Science Progress, April 1898), and
on seasonal dimorphism in South American Pierine butterflies
(Proc. Ent. Soc., December 7, 1898).

Professor Poulton published a paper on natural selection
as the cause of Mimicry (Journal Linnean Society, 1898, p. 558),
and in conjunction with Miss Cora B. Sanders attempted to
obtain experimental evidence of the struggle for existence in
certain insects (British Association Report, 1898, p. 906).
The former paper describes many examples in the Hope
Collections, the latter enquiry, aided by the Government
Grant Committee of the Royal Society, was in part carried
on in the Department.

The memoirs published in 1898 and 1899, together with
others which appeared in the latter part of 1897, nearly suffice
for a second volume of “ Hope Reports,” and it is hoped that
the issue will take place before the close of the present year.

ADDITIONS TO THE COLLECTION IN 1897
(tmperfectly recorded in the Report of that year).

It has been explained that the additions made in 1897
were labelled and catalogued only in small part when the
Report of that year was issued in June 1898. Only about
1,600 specimens out of a total of about 12,000 had been
catalogued and could be adequately acknowledged. The
present Report begins where that of 1897 becomes brief
and incomplete.

Rey. G. Dexter Allen presented 16 Lepidoptera Rhopalocera
from the Alps and 10 Coleoptera from the Alps (with the
exception of one from the Pyrenees).

Dr. E. Crompton and his brother, F. Crompton, Esq.,
presented a fine set of insects captured (1894-7) in the

6

neighbourhood of Victoria, Vancouver Island, comprising
74 Lepidoptera Rhopalocera, 121 Lepidoptera Heterocera,
6 Hymenoptera, 6 Diptera, and 3 Neuroptera. Dr. Crompton
also presented the insects which he captured in September,
1897, during Professor ’Poulton’s visit to Victoria, amounting
to 48 specimens of many Orders, including one Arachnid.

Many other naturalists also presented specimens of insects
captured at the same time in the same locality. Professor
Meldola, F.R.S., presented 49 specimens of many Orders, Pro-
fessor G. F, Fitzgerald, F.R.S., 23 specimens, Dr. Kirker, R.N.,
4 specimens, Professor Sherrington, F.R.S., 3 specimens, E. A.
Carew-Gibson, Esq., Ig specimens, and Professor Poulton 176
specimens. Eight specimens captured at an earlier date were
presented by E. Baynes Reed, Esq.

Thus the Hope Collections have been enriched by over 540
specimens with excellent data from this interesting locality.

Professor Meldola presented 18 insects of various Orders,
Professor Poulton 46 insects, and C, A. Ellis, Esq., 2 rare
Coleoptera (Carabidae) from the neighbourhood of Vancouver.

An Acridian from Vernon, B.C., was presented by E. M.
Walker, Esq., of Toronto.

Small numbers of insects of many Orders captured at
various places on the Canadian Pacific Railway between
Vancouver and Glacier House were also presented, viz.
31 specimens by Professor Poulton, 8 by Professor Meldola,
and 1 by Miss Cecil Vernon Harcourt. The localities repre-
sented are Mission Junction, North Bend, Revelstoke, Albert
Canyon, and Illecilliwaet.

Insects captured at or near Glacier House, on the summit
of the Canadian Pacific in the Selkirk Range, at a height of
from 4,000 to 6,000 feet were also presented—137 specimens
by Professor Poulton, 23 by Professor Meldola, 11 by Miss Cecil
Vernon Harcourt, and 1 by Professor W, Ramsay, F.R.S.

Thirty specimens from various localities between Glacier
House and Lake Louise were presented by Professor Poulton,
the places represented being Roger’s Pass, Six-Mile-Creek,
Beaver Mouth, Donald, and Palliser.

Twenty-three specimens from Lake Louise near Laggan, —

on the eastern slope of the Rockies at a height of from 5,550

a

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to 5.700 fect, were presented by Professor Poulton, and 16 by
Professor Meldola.

Five hundred and fifty-three specimens, captured in small
sets at a very large number of localities on the Canadian
Pacific and Grand Trunk Railways between Banff and Toronto,
were also presented. The principal sets came from the
following places, proceeding eastward from Lake Louise—
Banff, Medicine Hat, Waldeck, Rush Lake, Parkbeg, Moose-
jaw, Regina, Brandon, Carberry, Rennie, Rat Portage and
Nepigon. Of this number of specimens 8 were captured and
presented by Miss Cecil Vernon Harcourt, 10 by Miss E. C.
Ramsay, 14 by Mr. W. G. Ramsay, 9 by Professor Meldola,
3 by Professor G. F. Fitzgerald, 2 by Dr. W: Saunders of
Ottawa, 4 by E. H. Chapman, Esq., of Magdalen College,
24 by E. M. Walker, Esq., of Toronto, and the remainder by
Professor Poulton.

A very fine consignment of 633 insects of many Orders
from Russell (W. Manitoba) was presented by Mr. H. R.
Smith, late assistant in the Hope Department. They include
over 400 specimens of Lepidoptera, several of which will be of
much value for the series illustrating the subject of Mimicry.

One hundred and twenty-two insects from Virden (W. Mani-
toba) were captured and presented by Miss Mary G. Holmes.
The specimens are beautifully collected and accompanied by
excellent data.

Concerning the specimens from Southern Canada enumerated
above, it may be well to point out that even the common species
are very rare in European Collections. Thus Mr. H. R. Smith’s
valuable present included many specimens of two species of
Lrebia (£. epipsodea and £. discoidalis), which had never
before been obtained further East than Calgary. Such a large
number of specimens with excellent data from this part of the
world forms an accession of the utmost value to the University
Collections, and has also enabled us to enrich the British
Museum with many of the duplicates.

Professor Poulton presented 234, and Professor Meldola 24
‘specimens of insects captured, in August and September, in
the neighbourhood of Toronto. Professor J. Mavor, of Toronto,
and his son presented 12 insects from the same locality.

8

Professor Poulton presented 42 insects from the river St.
Lawrence between Quebec and Montreal, 129 from Quebec,
and 40 from Montreal. Miss Cecil Vernon Harcourt presented
I specimen from the latter locality.

A very interesting set of 41 insects of many Orders, captured
July 1886 in Hudson Strait (Lat. 62° 35’ N., Long. 77° 22’ W.),
was presented by F. F. Payne, Esq., of Toronto.

Miss Mary G. Holmes presented 69 insects from Ottawa.

Professor Poulton presented 26 insects from various localities
in Ontario, and 150, and Professor Meldola 2, from Niagara
Falls.

When not otherwise stated, all the captures in Canada were
in the months July, August, or September 1897.

Professor Poulton presented 53 insects from Detroit, 274 from
the neighbourhood of Chicago, and 287 from the neighbour-
hood of Hartland (Wis.); and in association with Professor
W. M. Wheeler of Chicago University 182 insects, and with
Professors Wheeler and S. Watase 14 insects from the neigh-
bourhood of Chicago. Professor Wheeler also presented 32
insects from the same locality, and 12 insects from various
localities in the United States. The latter will be especially
valuable for the Mimicry Collection.

Dr. G. W. Peckham presented 179 insects from the neigh-
bourhood of Hartland (Wis.), and in association with Professor
Poulton 12 from the same locality.

Twelve insects from Montana, U.S.A., were captured and
presented by Mrs. W. Ramsay and her son and daughter.

With the exception of some of Dr. Peckham’s (1896) and
Professor Wheeler's (1895 and 1896) captures, all these speci-
mens from the United States were taken in the same months
of 1897 as those from Canada.

The interesting Collection of 102 insects from Socotra,
captured 1896-7 by E. N. Bennett, Esq. M.A., has been
already alluded to. Mr. Bennett also presented 1 specimen
(Papilio) from Crete. One specimen (Hemipteron) from Tripoli
was presented by A. J. Evans, Esq., M.A.

Professor Poulton presented 8 Lepidoptera captured in
North America by his father in 1874-5, and a rare moth
from the Naga Hills, Assam (188g).

9

“Mrs. Bazett of Reading presented a useful set of 101 Lepi-
doptera, captured in 1893-4 in Switzerland (Vaud), and 28
from the Spanish Pyrenees (1892).

Mr. H. R. Smith presented 12 Lepidoptera from W. China,
the Khasia Hills, Assam, and Europe.

Mr. A. H. Hamm presented 14 Lepidoptera from a variety
of localities. These were especially useful in providing
specimens which were much wanted from certain localities
unrepresented in the Hope Collection, or of great interest in
themselves.

Colonel Swinhoe presented a fine set of 248 Heterocera
from the Khasia Hills, Assam (1893).

Mr. W. Holland presented 34 specimens (chiefly Lepido-
ptera) captured at various localities in Britain, to be added to
the general collections.

Mr. A. H. Hamm similarly presented 30 specimens.

Rev. O. Pickard-Cambridge, F.R.S., presented a useful set of
68 insects, chiefly Lepidoptera, received from Rev. J. T. Roberts
Rea from the Potaro River, Central British Guiana (1896).

Dr. F. A. Dixey presented 8 very interesting and valuable
hybrid Saturnian moths, obtained by Dr. M. Standfuss, of
Zurich.

Twenty-seven Lepidoptera, chiefly intended for the Mimicry

Collection, were purchased from Messrs.Watkins and Doncaster,

The British Collections also received a very large number of
accessions—by far the most important of recent years.

Rev. H. A. Harvey presented a specimen of a species of
Ichneumonidae captured in Oxford.

A small but valuable set of 42 Hymenoptera Aculeata was
presented by Edward Saunders, Esq., F.L.S., filling up some
of the gaps in our fine British Collection of this Order.

Miss E. E. Thoyts presented a valuable set of 339 insects
of many Orders captured by her chiefly in Sulhampstead Park,
Berkshire. Two hundred and eighty-eight specimens of
Hymenoptera were from the latter locality. All the specimens
are accompanied by careful and excellent notes.

Mr. W. Holland presented a splendid set of 1,630 specimens,
chiefly of Lepidoptera, from many localities, principally in the

10

neighbourhood of Reading. The data accompanying this
great accession are excellent.

Mr. H. R. Smith presented, on his leaving England for
Canada, a very fine set of 996 well-dated and localized
specimens, also chiefly Lepidoptera from the neighbourhood
of Reading.

Professor E. B. Poulton presented 1 specimen of Tadanus
bovinus, another Dipteron, and 3 moths captured by him at
Oxford in 1896 and 1897, and 136 specimens of Lepidoptera
and 2 of Diptera captured, with only one exception, by
him at Lulworth or Freshwater in July 1883; and his son,
Edward P. Poulton, presented 2 rare moths captured by him
(August 1897) at St. Helen’s, Isle of Wight.

A specimen of Szvex gigas, captured in the Oxford Market
in 1896, was brought to the Museum.

Mr. A. H. Hamm presented 167 specimens, mostly Lepi-
doptera, from the neighbourhood of Oxford, accompanied by
excellent data.

Eighteen moths, captured in 1895 at Freshwater, were pre-
sented by W. M. Geldart, Esq., M.A., St. John’s College.

One dragon-fly, captured at Oxford, was presented by
Mr. Burden; and a wasp by Mr. H. Trim.

Colonel Swinhoe presented 1 bee, captured (1896) at Har-
penden; Professor Poulton, 6 Hymenoptera Aculeata, captured
at Great Malvern (1897); and Dr. Dixey, 1 beetle, taken in
Oxford (1897). .

The insects presented in 1897 have not been catalogued
further than this point (about 9,700).

There still remain the following collections:—A valuable
collection of many hundreds of insects brought by C. V. A.
Peel, Esq., from: Somaliland. These have received their
printed labels, but cannot be catalogued until they have been
worked out. Several new species and some new genera are now
being studied by specialists in the various groups, and it
is expected the memoir will be finished by the end of the
summer.

A large and varied collection of butterflies and a few moths,
chiefly from India and Burmah, presented by Colonel Swinhoe.

via

il

Many of these will be of great value in the Mimicry Col-
lection.

A large collection, presented by Herbert Druce, Esq., F.L.S.,
consisting of butterflies from Africa and South America, and
moths from Central America. These will be of great value
to the general collections.

Dr. Dixey’s important present of British Insects from the
neighbourhood of Morthoe, N. Devon, is also at present
uncatalogued.

ADDITIONS TO THE COLLECTION IN 1898.

The great strain thrown upon the Department by the acces-
sions in 1897 has only permitted a small proportion of the
specimens presented in 1898 to be as yet catalogued.

One of the most interesting additions made in recent years
to the Department and the Museum is dte to the kindness of
Abbott H. Thayer, Esq., of Scarborough, N.Y., U.S.A., who
prepared and painted models of two birds, illustrating the
reason for the gradation of the colours of animals in nature
—from dark on the back, through lighter shades, to white
on the belly. These models show the difference between
a uniform tint and colours so graded. In the latter case
the increasing lightness is seen exactly to neutralize the
increasing shade as the sides and under-surface of the animal
are more and more shielded from the light of the sky in
passing from the back to the belly. Furthermore the cold
white light of the sky combined with the brown of the
animal's back produces a colour-effect similar to that of the
brown reflection from the earth combined with the cold white of
its under-surface. The uniformly coloured model stands out in
startling contrast beside the graded one which is inconspicuous
and almost ghost-like, as so many animals are in nature.

The nest of a Marsh Warbler (Acrocephalus palustris) from
Kingham, containing a cuckoo’s egg buried under the lining,
was presented by W. Warde- Fowler, Esq., M.A. (Lincoln
College). A nest of the Long-tailed Tit (Parus caudatus)
was presented by G. Holding, Esq., M.A. (All Souls College).
A nest of the Great Reed Warbler (Acrocephalus turdoides),

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taken on the banks of the Guadalquivir by H. F. Witherby,
Esq., was presented by him.

A valuable set of 51 insects of many Orders, chiefly Lepi-
doptera, captured in Mexico almost exclusively in 1897, and
71 captured in 1898, were presented by Osbert H. Howarth, Esq.
Many of the specimens are of great value, and the data accom-
panying them most full and precise.

A valuable set of 175 insects, almost exclusively Lepidoptera,
from Queensland (1897), Himalayas (1897), India (1897), Cape
of Good Hope (1897), and Manila (1896), was presented by
G. C. Griffiths, Esq.

Ninety-eight Lepidoptera from the Upper Engadine and
the Upper Rhone Valley (1869) were presented by Rev. A. G.
Butler, M.A. (Oriel College). |

One hundred and two Lepidoptera from the South of
France (1898) were presented by Colonel J. W. Yerbury.
The data sent with the specimens are excellent.

Two Diptera from Arizona, captured (1897) with the 2
Hymenoptera they mimic, were presented by R. C. L. Perkins,
Esq., B.A. (Jesus College).

Twenty-two insects from various localities in Palestine and
Syria (1898) were presented by E. N. Bennett, Esq. M.A.
(Hertford College). .

Twelve valuable specimens of Orthoptera of various groups
from Java, Celebes, and Ceylon (1889-96) were presented by
Malcolm Burr, Esq. (New College).

The eggs of a Locustid from France (1897) were presented
by W. J. Lucas) Esq.

A valuable set of 194 insects of many Orders, Arachnida
and Myriapoda, from Manchuria (1898), was presented by
R. T. Turley, Esq.; and’ Mrs. Turley:

Two rare Coleoptera from Patagonia (1898) were presented
by M. Jacoby, Esq.

Twenty-two Lepidoptera, with excellent data, from Mexico
and Trinidad (189%) were presented by L. Richardson, Esq.

A pair of a rare Danaine butterfly from Madagascar (1891)
was obtained by purchase from Messrs. Watkins and Doncaster.

The British Collections also received many accessions,

13

a relatively few of which have been catalogued and _ in-
corporated.,

Seventy-two insects, captured in 1898 in various localities —
Oxford, Berkshire (Sulhampstead Park), &c.—were presented
by Professor Poulton.

An interesting variety of the male of Pzer?s brassicae from
N. Devon (1897) was presented by Selwyn Image, Esq., M.A.
(New College).

A fine specimen of Bembex from Oxford (1898) was pre-
sented by Miss Butler,

A series of 7 Zygaena trifolit, including some interesting
varieties from Sussex, together with a pair of Vyssza lapponaria
from Scotland, were presented by W. M. Christy, Esq.

Ten specimens, comprising dragon-flies and a few specimens
of Orthoptera from various localities, all much needed by the
Collection, were presented by W. J, Lucas, Esq., B.A.

In addition to the catalogued and incorporated specimens
large numbers have been received in 1898, the majority of
which have been set, while many have received their printed
locality labels. These uncatalogued additions will how be
briefly mentioned, and will be acknowledged in detail in next
year’s Report.

Very large and valuable additions, chiefly from tropical
America, to the collection of moths have been made by
Herbert Druce, Esq., F.L.S., and Col. Swinhoe has presented
many Oriental moths from his own collection in the progress
of his work in the Department: also a few butterflies from
various localities.

The most important of all additions made in the year is, as
has been already inferred, the captures made and presented
by Guy A. K. Marshall, Esq., in S. Africa. These consist of
a fine series of butterflies and Orthoptera, and a few insects
of other Orders, from Salisbury, Mashonaland, and a few rare
butterflies from Cape Town, and moths from Delagoa Bay.
In addition to these great accessions to the general Collection,
Mr. Marshall has made special collections to add to the series
illustrating various zoological principles. Thus he has sent
mimetic groups caught in one place on one day, and groups

14

of butterflies showing the attacks of birds or other enemies,
such as the species of Charaxes with the “ tails” bitten off the
hind wings. But perhaps the most interesting and valuable
gift of this kind is a set of specimens proving that Precis
octavia-natalensis, a red and black butterfly with under side
very similar to the upper side of the wings, occurring in the
damp summer season, is only a seasonal form of the blue,
black, and red Precis sesamus,a butterfly occurring in the dry
winter, with an under side entirely unlike the upper, and with
a wing of a different size and shape to that of P. octavia-
natalensis. Mr. Marshall has sent two examples of the
parent red form which laid the egg, and the blue offspring
which he succeeded in rearing from it.

In the present year (1899) Mr. Marshall has further helped
us by presenting a fine series of Coleoptera from Salisbury,
and many other groups for the mimicry and other series,
including one of unique interest from the extraordinary
number and diversity of the forms which fall into it. All
the specimens received from Mr. Marshall are set, and will
soon receive their printed labels.

The most important addition that has yet been made to
the rather poor set of European species contained in our
Collections was due to a visit to Switzerland in July, 1898,
made by Professor and Mrs. Poulton and Miss Cora B.
Sanders. Many hundreds of specimens with very full data
have been set, and will receive printed labels as soon as time
will permit.

Very fine sets of Lepidoptera from Mexico and Queensland
have been presented by G. C. Griffiths, Esq., in addition to
those which have been described as already catalogued. We
owe to Mr. Griffiths’s kindness further sets from the same
localities presented in the present year.

An interesting series of insects of many Orders, unfortunately
in very bad condition, was brought from the Omdurman
campaign, and kindly presented by E. N. Bennett, Esq., M.A.
A brief account of the captures is printed in Mr. Bennett’s
recently published book on the campaign.

A small but valuable set of insects from Eastern Europe
was presented by Malcolm Burr, Esq.

15

Dr. Dixey presented a large number of butterflies from
Singapore.

Professor Poulton presented a few moths which emerged in
Oxford during the course of the year 1898 from pupae obtained
from larvae found by him in N. America the year before.

Miss Merivale presented a moth from Barbadoes, and 2 beetles
from the River Niger were presented by B. Tomlin, Esq.

The British Collections have also been enriched by a large
number of specimens which have not as yet been catalogued.
Kind help is here briefly acknowledged from the following :—
fewinim, VW: Holland, L. Clifford Brown, Dr. Bo A] Dicey,
A. H. Hamm, E. B. Poulton, R. W. Poulton, E. W. Blagg,
Rev. E. C. Dobree Fox, W. J. Bruce, F. Bolton, F. Swanton,
A.W. Daish, — Bartlett, A.G.S.Symonds, E. Wheal, J. Taylor,
Mrs. Pelham, Miss Thoyts, Miss C. V. Butler, Miss E. M.
Ramsbotham, Miss K. D. Mossman, and Miss C. B. Sanders.
Full acknowledgment will be made in the Report of next year
when the specimens will have been catalogued and incorporated.

Special mention must however be made here of the fine set
of British Lepidoptera presented by H. Goss, Esq., F.L.S.

AOMOTELONS TO. THE HOPE, LIBRARY EN» 1898:

The Smithsonian Institution (United States National
Museum, Washington) presented the publications which deal
with the subjects of the Department.

The Boston Society of Natural History and the Bombay
Natural History Society presented their publications for the
year.

_ The University of the State of New York presented its
Reports.

The Linacre Professor of Comparative Anatomy, Oxford,
presented “ The Linacre Reports,” vol. iii.

‘The Radcliffe Librarian, Oxford, presented the Catalogue
of Books added during 1897. .

The Transactions of the Entomological Society, and the
Transactions and Journal of the Linnean Society for the year
1898, were presented by Professor Poulton.

Canon W. W. Fowler presented three more parts of his

16

monograph on the Rhynchota Homoptera, vol: ii, in the
Biologia Centrali-Americana.

The Agent-General for Queensland presented the “ Annals
of the Queensland Museum,” No. I.

The National Museum of Rio de Janeiro presented its
Journal, vol. i.

The Superintendent of the Museum of Zoology, Cam-
bridge, presented the Thirty-second Annual Report (for 1897).

The Radley College Natural History Society presented its
list of the Fauna and Flora of the district.

Sir John Evans presented the Memoirs of the Academy of
Science, Bologna, series v, vol. vi, 1896-7.

Copies of original papers were presented by the.following
authors:—P. P. Calvert, Esq., of Philadelphia, Dr. A. G.
Butler, Abbott H. Thayer, Esq., of New York, Dr. H. J.
Hansen and William Sorensen, Frank Finn, Esq., Herbert
Druce, Esq., Guy A. K. Marshall, Esq., Roland Trimen, Esq.,
Hont M.A. F.R.S;,.and)\ Dr: Karl Jordan.

Miss E. A. Ormerod very kindly completed the set of her
valuable ‘‘ Reports of Observations of Injurious Insects” in
the Hope Library, presenting the 16th to the 21st for the
years 1893 to 1898 both inclusive, and also the “ Report” for
1884 which was missing from the earlier set. Miss Ormerod
also presented the following volumes of which she is the
authoress, ‘ Handbook of Insects Injurious to Orchard and
Bush Fruits, with Means of Prevention and Remedy,” London,
1898, ““ A Text-book of Agricultural Entomology,” London,
1892, and “A Manual of Injurious Insects,’ London, 18go.
This kind present will be of great value to the important
section of the Hope Library which deals with Economic
Entomology.

“ British Orthoptera,” Huddersfield, 1897, was presented by
the author, Malcolm Burr, Esq. \ \

A few second-hand books were purchased together with
the Zoological Record, the parts of Barrett’s “ British
Lepidoptera,” the Ray Society volume, and the parts of
Rippon’s ‘‘ [cones Ornithopterarum” for the year.

EDWARD, B:. POUL TOR

Report of the Hope Professor of Zoology.

The additions to the Hope Department during 1899 are
very large and important, greatly surpassing those of 1898.
Nearly 7,000 specimens have been already catalogued and
incorporated, and large numbers can only be provisionally
acknowledged on the present occasion. Mr. Guy A. K.
Marshall’s specimens from Mashonaland are, as on previous
occasions, the most perfectly collected and valuable specimens
received by the Department. Many of them illustrate
problems of natural history of the widest interest, and form
the evidence upon which the interpretation of several
extremely difficult questions is to be sought. Other accessions
are also of the highest importance, especially the valuable
collection of moths collected by Cecil Barker, Esq., in Natal,
presented by Roland Trimen, Esq., Hon. M.A., F.R.S:; of
butterflies from Karwar, on the Malabar Coast, collected and
presented by G. Keatinge, Esq. ; and of butterflies from Sanda-
kan, N. Borneo, collected by A. L. Cook, and presented by
Herbert Druce, Esq., F.L.S. These, with the fine additions to
the collections of British insects, mark the year 1899 as a very
important one in the history of the Hope Department.

The final arrangement of the general collection of butterflies
was continued by Mr. Holland, the Satyrinae, Elymniinae,
Amathusiinac, and part of the A7orphinae, being completed in
150 drawers. At this point the available cabinet space was
exhausted, 40 drawers being kept in reserve for the Pzerinae,
which Dr. Dixey was arranging.

Towards a fund for the provision of new cabinets the
Delegates of the Museum granted 450 to be paid in 1899,
and £50 in 1900, and similar amounts were voted by Deeree.
A further consignment of 200 drawers was therefore ordered
of Messrs. O. E. Janson & Son. Ten cabinets of 20 drawers
each were completed and delivered in March of the present
year, and the final arrangement resumed, the J7orphinae and
Lrassolinae being completed, and considerable progress made
with the immense group of the Mymphatinae.

B

2

Dr. Dixey continued the arrangement of the Pierinac. His
chief work consisted in the completion of the arrangement of
the extremely difficult genera Colias and Terias. The species
are arranged in a most instructive manner to show the geo-
graphical and seasonal variations.

The arrangement of the Phytophaga (Coleoptera) so kindly
worked out for the Hope Department by Mr. M. Jacoby, was
next continued by Mr. Holland.

The classification is now complete up to the point reached
by Mr. Jacoby, and the collection occupies 41 drawers in
a cabinet of an old type with shallow drawers, not unsuitable
however for these small beetles. A large amount of labour
has been devoted to this collection, which is now in a very
satisfactory condition. :

Mr. A. H. Hamm has been chiefly occupied in setting and
printing labels for the large number of accessions, and in reset-
ing a large portion of the Lepidoptera which have been finally
arranged. Many of these are very old specimens, in poor
condition, and require the greatest care. During the Long
Vacation he named and arranged the Syntomid Moths of the
Department, following Sir George Hampson’s recently pub-
lished monograph.

Colonel Swinhoe completed the arrangement of the
Oriental moths, thus finishing an arduous piece of work which
has occupied him intermittently for many years. The working
out and naming of this high proportion of our large collection
of moths will be of the greatest value in the labour of bringing
together the whole into one comprehensive series.

The arrangement of the British Lepidoptera has been
continued by Mr. Sidgwick and Mr. Pogson Smith. The
Tortrices are now complete, and the 7zezza about half finished.
Mr. Edward Saunders and Mr. R. C. L. Perkins visited the
Department in the course of the year, and afforded valuable
help in naming the most obscure among the recent additions
to the British Hymenoptera. . Dr. Frederick Moore kindly
eave information as to the history of specimens formerly in
the collection of the East India Company. Mr. Gilbert J.
Arrow studied the section of Coleoptera he was at work upon

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6 —"?

3

in the British Museum, and named many of our specimens.
Mr. Roland Trimen kindly helped in the identification of
many African butterflies. Mr. E. E. Austen looked through
the general collection of Diptera, and made notes on some of
the specimens. Mr. M. Jacoby continued his kind assistance
with the Phytophaga.

The visit of the Council of the Entomological Society took
place on July 1-3. Mr. Roland Trimen, Hon. M.A., F.R.S.,
Vice-President, Mr. R. McLachlan, F.R.S., Treasurer, Mr, J. J.
Walker, R.N., and Mr. F. Merrifield, Secretaries, Mr. J. W.
Tutt, and Mr. H. St. J. Donisthorpe were present.

Professor Meldola, F.R.S., Mr. Herbert Druce, F.L.S., Mr.
Martin Jacoby, were also present, together with the Oxford
students of the Insecta. The Hope Collections were inspected,
and many interesting and useful suggestions made by the
distinguished specialists who were present. ‘The visitors were
entertained by the Hope Curators and their friends. Mr.
Arthur Evans kindly invited the party to Youlbury on the
afternoon of July 2.

-The collections have also been visited by many other
eminent naturalists in the course of the year, including
Meee. Godman, F.R-S:, Hon. DC... Professor Enc.
Mion Reo. Mr W. Le Sclater, Mr. C. W.. Dale, MrG: C:
Griffiths, Mr. F. A. Heron, and Mr. W. J. Lucas.

I have much pleasure in acknowledging the great kindness
of the members of the staff of the Insect Department of the
British Museum of Natural History. It is a satisfaction that
the Hope Department has been able to render some assistance
by sending types for comparison with the specimens in the
national collection.

As regards research upon the specimens in the Department,
Mr. C. V. A. Peel’s Somaliland insects were worked at by
many eminent authorities in the various groups, and the paper
was read before the Zoological Society in the beginning of the
present year. The issue of “ Hope Reports,” Vol. II, is being
deferred until this memoir can be included.

Professor Poulton, in association with Mr. F. Merrifield,
brought out a paper in the “ Transactions of the Entomological

Bae

4

Society,” on the relation of the colours of pupae to the
surroundings of the larvae, &c. Much of the recorded work
was done in the Hope Department in 1898 and 1899, by
Professor Poulton, assisted by Miss Cora B. Sanders, Miss
Mabel E. Notley, and Miss Florence A. Wright. Observations
of Mr. W. Holland and Mr. A. H. Hamm are also described
in the paper.

The experimental inquiry into the struggle for existence in
insects was continued by Professor Poulton, Miss Notley, and
Miss Wright, aided by the Government Grant Committee of
the Royal Society.

A large amount of work was involved in studying and
making out the species contained in the collections illustrating
mimicry and allied subjects made by Mr. R. Shelford, chiefly
in Borneo, and by Mr. Guy A. K. Marshall, chiefly in
Mashonaland. In this work Professor Poulton has received
much kind help from the staff of the British Museum of
Natural History, from Mr. R. McLachlan, F.R.S., and
Mr. M. Jacoby.

Dr. F. A. Dixey, M.A., Wadham College, has undertaken
a prolonged investigation into the secondary sexual characters
of the Prerinae.

Mr. T. N. Annandale, B.A., Balliol College, investigated the
insects and other forms brought by him from Siam, and
commenced a paper upon various aspects of their natural
history.

ADDITIONS TO THE COLLECTION IN 1896.

A valuable collection of 357 insects, chiefly Rhynchota,
Hemiptera and Homoptera, from Sydney, Brisbane, New
Zealand, and Java (1891-1893), was presented by Julius
Langhans, Esq., Hertford College. In the press of other
work, the setting, printing, and cataloguing was deferred until
189g. The specimens are now incorporated, and are a very
welcome addition to the important collections by which these
two Sub-Orders are represented in the Hope Museum.

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5

ADDITIONS TO THE COLLECTION IN 1897.
(Catalogued and incorporated since the issue of the tast Report.)

About 9,700 specimens presented in 1897 were recorded in
the Reports for that year and for 1898.

The following additional donations have now been cata-
logued :—

Colonel C. Swinhoe, Hon. M.A., Wadham College, pre-
sented 250 Lepidoptera from a great variety of Oriental
localities (various dates) and a few from America. Many
specimens will be of value in the series illustrating Mimicry.

Mr. W. Holland presented a specimen of 72ramala limniace
from Sikkim.

A valuable series of 150 butterflies from Sierra Leone
(about 1890) was presented by Herbert Druce, Esq., F.L.S.

A small set of 27 butterflies from various localities was
presented by C..V A. Peel, Bsq., F-Z:S:

The British Collections were also enriched by 321 specimens
collected in 1897, with great care, by Dr. F. A. Dixey, M.A.,
Wadham College. Nearly the whole were captured near
Morthoe, N. Devon, and all are accompanied by full and
excellent data.

In addition to the above, Mr. Peel’s large and valuable
collection made in Somaliland in 1895 and 1897 is still waiting
for the appearance of the memoir which has been written
upon it, in order that references may be placed with the
specimens. The paper is now in the Press, and will appear
in the forthcoming Part I of the “ Proceedings of the Zoological
Society” for 1g00, so that the incorporation of this fine
collection cannot be delayed much longer.

The valuable collection of South American butterflies and
Central American moths presented by Mr. Druce is also as
yet uncatalogued.

ADDITIONS TO THE COLLECTION IN 1898.
(Catalogued and incorporated since the issue of the last Report.)

Only 835 specimens, presented in 1898, were recorded in
the Report for that year. Since then much progress has

6

been made, and comparatively few specimens remain un-
catalogued.

A very valuable set of 346 insects was presented by Guy A. K.
Marshall, Esq., of Salisbury, Mashonaland. They consist of
266 Lepidoptera (263 butterflies and 3 moths), chiefly from
Salisbury and other localities in Mashonaland (1895, 1897,
and 1898), and a few from Delagoa Bay (1897), and Cape
Town (1897); 3 Coleoptera, 4 Hymenoptera, 4 Hemiptera,
and 69 Orthoptera, all from Salisbury (1898). The majority
of the specimens are a most welcome addition to the general
collection, but among them are many groups of even greater
value, inasmuch as they illustrate various aspects of the
natural history of insects. Thus there are many specimens
showing the apparent attacks of birds or other enemies: butter-
flies with notched wings or with the tail-like prolongations of
the hind wing (in the genus Charaxes) wanting and probably
bitten off. Also groups of mimetic insects of many kinds
captured on the same day, and specimens showing the
parasitic attacks to which insects are subject. The specimens,
like all those which we owe to Mr. Marshall’s generosity, are
beautifully collected,andare accompanied by extremely full data.

Mr. A. H. Hamm presented 36 Hymenoptera and Diptera
from the neighbourhood of Oxford (1898), and a specimen
of Limunas chrysippus from N. Borneo (1892).

A fine set of 114 Lepidoptera from Brisbane (1897, 1898)
was presented by G. C. Griffiths, Esq. The specimens are in
beautiful condition, many of them being bred specimens.
In some cases Mr. Griffiths presented the pupae from which
the butterflies emerged in the Museum. Mr. Griffiths also
presented 39 Lepidoptera from Jalapa, Mexico (1893).

Seven bred specimens of Lepidoptera, together with 3
pupa-cases, and 14 Hymenoptera and Diptera from the
neighbourhood of Oxford (1898), were presented by Mr. W.
Holland (for the General Collection).

A set of 47 insects of many Orders from the Upper Nile
near Khartum (1898), was presented by E. N. Bennett, Esq.,
M.A., Hertford College. Many of the specimens will be of
considerable value on account of the locality.

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7

A small collection of 35 insects, chiefly Lepidoptera, from
South-Eastern Europe (1898), was presented by Malcolm
Burr, Esq., New College. The localities render the specimens
of special value to the Hope Collection.

Two Coleoptera from Lokoja on the River Niger (1896)
were presented by Brockton Tomlin, Esq.

Colonel C. Swinhoe, Hon. M.A., Wadham College, presented
6 butterflies from various localities.

A hawk-moth .from Barbados (1896) was presented by
Mrs. Merivale.

A moth from the Adirondack Mountains (larva found 1891)
was presented by Henry Balfour, M.A., Trinity College.

Professor Poulton presented 2 specimens of Catocala elocata
bred in 1889 from European eggs, 2 specimens of a Smerin-
thine hawk-moth (with the pupa-cases) bred from larvae
found in Vancouver Island (1897), a moth similarly bred from
a larva found in Toronto, and 2 moths bred from larvae from
Hartland, Wisconsin, in the same year.

Twenty-two Lepidoptera from South-East Mexico and
Trinidad were presented by Lawrence Richardson, Esq.

Five British Lepidoptera for the General Collection were
presented by L. Clifford Brown, Esq.

A very interesting collection of 99 specimens captured in
1898 in the neighbourhood of Urumiah (North-West Persia),
and 2 moths from Kivorak (Transcaucasia), were presented by
R. W. T. Giinther, Esq., M.A., Magdalen College. The species
have been described by experts in the respective groups in
Mr. Giinther’s paper in the “ Linnean Society’s Journal,” vol.
XXvii, p. 345, and references to the appropriate page now
accompany every specimen. The locality and exact data also
render the accession of great value to the Hope Collection.
Some of the specimens will be of special value for the series
illustrating biological problems.

Mr. Giinther also presented a collection of Lepidoptera
made in the neighbourhood of Urumiah by the Rev. S. J.
Daltry. These specimens have not yet been catalogued, in-
asmuch as it is hoped that further data may become available.

8

Professor Poulton presented 1,069 specimens and Mrs.
Poulton 8 specimens of insects of many Orders captured in
Switzerland (Valais and Bernese Oberland) in 1898. Many
specimens are of value for the Biological Series.

Miss Cora B. Sanders, Lady Margaret Hall, presented 404

specimens captured in Switzerland at the same places and
times as those last mentioned. Three specimens of Pafzlio
podalirius, bred in Oxford from larvae found near Visp, were
presented, together with their pupa-cases, by Miss Sanders
and Professor Poulton.

This addition of nearly 1,500 Swiss specimens, mostly in
excellent condition and with full data, is an important
accession to the General Collection.

In addition to the above there remain uncatalogued a large
and valuable collection of moths, chiefly from tropical America,
presented by Herbert Druce, Esq., F.L.S.; a fine set of butterflies
from Singapore, presented by Dr. F. A. Dixey, M.A., Wadham
College ; the latter are waiting in the hope of further data.

The British Collections have also received*the following
accessions :—Miss E. E. Thoyts presented 25 specimens of
Lepidoptera, Hymenoptera and Diptera, from Sulhamstead
Park, Berkshire (1898).

Four insects from Oxford (1898), 3 having been captured in
the University Museum or its grounds, were presented by
Mr. H. Trim.

A specimen of Sphinx convolvuli, captured in High Street,
Oxford (1898), was presented by Mr. W. T. Bruce; another
specimen, captured in Wellington Street, by Mr. F. Bolton;
a third, captured at Youlbury, Boar’s Hill, by Prof. Poulton;
and a fourth from St. Helen’s, Isle of Wight, was purchased
from a boy.

Professor Poulton also presented 26 specimens of various
Orders from Marlston (Newbury), Birdlip (1891), and
St. Helen’s (Isle of Wight), and Miss Cora B. Sanders 12
Diptera from the latter locality. Two Lepidoptera from
St. Helen’s were presented by Ronald W. Poulton, and 6 fine
specimens of Pyralis costalis by Miss Sanders. All undated
captures were made in 1898.

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A fine specimen of Acridium aegyptium was captured alive
in 1898 at St. Helen’s, and presented by Mr. A. W. Daish.
This specimen is of interest as bearing upon the introduction
of insects into new countries.

Mr. W. Holland, of the Hope Department, presented
175 insects of various Orders from St. Helen’s and the
neighbourhood of Oxford (1898). They include a fine set of
Hymenoptera, with the Diptera which mimic them, for the
Biological Series.

A fine series of 12 Zygaena minos, captured in Wales (1896)
by E. W. H. Blagg, was presented by Dr. F. A. Dixey, M.A.,
Wadham College, who also presented 66 insects of many
Orders, from the neighbourhood of Morthoe, N. Devon, 10
from Lundy Island, 12 from near,London (Coldfall Wood,
East Finchley), and 8 from near Oxford (all 1898 except the
latter, which are from 1893 to 1897). The specimens are
rendered specially valuable by full and precise data. Two
specimens of Hemiptera from Morthoe (1898) were presented
by Giles Dixey. Dr. Dixey also presented 6 British butter-
flies from various localities (1876-97), showing the apparent
attacks of birds.

Thirteen butterflies from near London (Wandsworth) and
the Isle of Wight were presented by L. Clifford Brown, Esq.
Mr. A. H. Hamm, of the Hope Department, presented 337
insects of various Orders from the neighbourhood of Oxford,
S. Devon, and Wellington College, all captured in 1898, and
one captured at Basingstoke about 1888. Many Hymenoptera
and mimetic Diptera captured on the same day will be
valuable for the Biological Series, together with specimens set
in the attitude of rest, and others showing the apparent attacks
of enemies.

Four specimens of Caradrina ambigua from S. Devon (1897)
were presented by the Rev. E. C. Dobree Fox, together with
a specimen of Lycaena aegon from Westmorland (1897).

A female specimen of Gonepteryx rhamnt, found hybernating
on. Dec. 30, 1898, was presented by Mrs. H. F. Pelham,
Trinity College.

10

Mr. Joseph Walker, of the Radcliffe Library, presented
a queen wasp found hybernating in the University Museum
on Dec. 13, 1898.

A fine set of 327 specimens of Lepidoptera from various
British localities was presented by Herbert Goss, Esq., F.L.S.
The specimens are in very fine condition, and were captured or
bred at various dates by Mr. Goss himself. They form a very
welcome accession to the collection of British Lepidoptera.

Miss E. M. Ramsbotham and Miss K. Mossman presented
6 cast skins of a species of Per/a from Yorkshire (1898).

A species of Cassida from Enslow Bridge (1898) was
presented by A. M. Bell, Esq., M.A., Balliol College.

The cocoon and pupa-case of Procris geryon, from Birdlip

(1898), was presented by Miss C. V. Butler.

A fine specimen of Arctia caja from Oxford (1898) was
presented by Mr. J. Taylor.

Five specimens of Crabro interrupta were bred (1898) from
some rotten wood containing their pupae. Presented by
Miss Ward. From Oxford.

Miss Cora B. Sanders and Prof. Poulton presented the
pupa of Vanessa urticae, with the parasitic Zachina which
emerged from it, and the puparium of the latter.

Miss Sanders presented 12 bred specimens (1898) of the
third brood of Pieris brassicae. Four of these are for the
General Collection.

A pair of Exdromis versicolor, captured in Kent about
1868, was presented by the Rev. A. G. Butler, M.A., Oriel
College.

ADDITIONS TO THE COLLECTIONS IN 1899.

Very large accessions have been made in the year 1899, and
although nearly seven thousand specimens have already been
catalogued, large numbers are still waiting, and can only be
provisionally acknowledged on the present occasion.

A pair of Colias hecla from Greenland, and gg specimens of
the genus /redza from various localities, representing nearly
the whole of the species, were presented by H. J. Elwes, Esq.,

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F.R.S. These latter were of great value in working out the
species of this important group of mountain butterflies, upon
which Mr. Elwes is the recognized authority.

An invaluable collection of 495 insects from Mashonaland,
nearly all from the neighbourhood of Salisbury, and 2 from
near Durban, Natal, were presented by Guy A. K. Marshall,
Esq. The General Collection has been enriched by an
important addition of specimens, in beautiful order, and with
more complete data than almost any others in our possession.
Furthermore a large number of specimens represent the
material which Mr. Marshall has specially collected, in
order to work out and illustrate various biological problems
of the utmost interest.

Thirteen examples of Precis sesamus were in reality
presented in 1898, and are briefly referred to in the Report
of that year. They were however inadvertently omitted from
the catalogue of Mr. Marshall’s donations in 1898, and are
therefore included in those of 1899. These specimens are
historic, for they constitute the evidence of the most wonderful
known example of seasonal dimorphism, proving that the red
and black Preczs octavia-natalensis is the summer form of the
blue, black, and red P. sesamus, a butterfly which flies in the
comparatively dry winter, and differs from the former in
habits, pattern, size, shape of wing, and in the relation of the
colouring of the under side to that of the upper side of the
wings. The series contains specimens of the former captured
at the end of summer, and of the latter at the beginning of
winter, 1898 (a fourteenth specimen, of octavia, captured in 1899
carries the sequence on for another season), as well as two
female specimens of P. octavia-natalensis, and butterflies of the
other form obtained from the eggs which they laid. A beautiful
photograph of the whole group, taken by Mr. Alfred Robinson
of the University Museum, is now at the Paris Exhibition.

Mr. Marshall also presented many groups illustrating
Mimicry and Common Warning Colours among insects.

A group of 18 specimens of 5 very similar species of Acraca,
captured on one day, illustrates the tendency of the insects of
specially protected groups to assume a similar appearance.

12

Another smaller group of 4 specimens showed the same
tendency to be strongly marked in two larger species of
Acraca (A. anemosa and A. natalica), also captured on the
same day.

A specimen of Teracolus phlegyas (female) subjected to
damp heat in the pupal condition.

Twenty butterflies showing injuries probably caused by the
attacks of enemies.

Five beetles taken almost uninjured from the crop of a
guinea fowl (Wumida coronata).

A large convergent group of 45 insects of various Orders,
all red-brown in colour with a black patch on the posterior
part of the body or wings. The appearance is evidently mimetic
of Lycid beetles, of which several species are included. The
group comprises many beetles in addition to the Lycidae,
many //ymenoptera, a fly, two moths, and a bug.

Other interesting groups are—g small beetles with a very
similar appearance (type of pattern that of the included
Phytophaga); 24 large beetles with the pattern most charac-
teristic of the Cantharidae; also small groups exhibiting
resemblance between Coccinellid beetles and a bug (4 speci-
mens); an Ichneumon, a beetle, and a bug (3 specimens) ;
wasps and an Ichneumon (3 specimens); 4 pairs, each con-
sisting of a different species of stinging Hymenopterous insect
and a mimetic fly; 13 specimens of J/utellidae, spiders
and beetles with a similar colouring; 2 large species of
Carabidae (3 specimens), with very similar apparently warning
markings produced in quite different ways; 16 beetles of
various groups with a similar type of colouring.

Many of these examples are new species hitherto undescribed.
In addition to the above, Mr. Marshall has sent a magnificent
new fly of the genus //yferechia, mimetic of an African
xylocopid bee.

The 2 specimens from Natal (1897) are bugs of entirely
different groups, and yet with an almost identical pattern and
colouring.

I have described this especially valuable material at length,
but we also owe to Mr. Marshall’s generosity important

13

additions to the General Collections, including 3 Hymenoptera,
2 Hemiptera, and a very fine collection of 295 Coleoptera, all
from the neighbourhood of Salisbury (1898, 1899), and a further
fine set of Lepidoptera which arrived later, and have not as
yet been catalogued.

A fine set of 205 specimens of Limunas chrysippus and some
allied forms, from various localities in Afghanistan, Baluchistan,
and India (1879-83), was presented by Mr. W. Holland. The
specimens are from the Collection of Colonel Swinhoe, Hon.
M.A., Wadham College, and are a valuable addition to that
important department of the Hope Collection which deals
with variation of species in relation to geographical dis-
tribution.

Five specimens of Z. chrysippus, and an allied form, from
Greece and various localities in the eastern part of its range
(1877-92), were presented by J. J. Walker, Esq., R.N.

A fine set of 234 Lepidoptera, in beautiful condition, from
the neighbourhood of Brisbane (1897, 1898)—170 from Jalapa,
Mexico (1893); and 15 from various localities in tropical
S. America (1896, &c.)—was presented by G. C. Griffiths, Esq.

A specimen of Avxosia plexippus and its mimic Lementtis
misippus, captured on the same day (July 26, 1899) at Lake
Simcoe, Ontario, were presented by E. M. Walker, Esq., of
Toronto.

A set of 180 butterflies from Ashanti (1896) was presented
by Captain the Hon. Grosvenor Hood. The suggestion was
kindly made to Captain Hood by the authorities of the British
Museum (Natural History). The specimens will be of great
value to the Hope Collection on account of the locality—Cape
Coast Castle to Kumassi.

Ninety Lepidoptera, Hymenoptera, and Coleoptera, from
Virden, Manitoba (1898), were presented by Miss Mary G.
Holmes. The collection includes specimens for the Mimicry
Series, and all the insects are accompanied by the fullest data.

A very valuable collection of 700 butterflies from Sandakan,
Borneo (1895-96), was presented by Herbert Druce, Esq.,
F.L.S. This fine series is a very welcome addition, the
majority of specimens being in beautiful condition ; many

14

specimens will be of value for the Mimicry Series. Mr. Druce
also presented 5 specimens from other localities in Borneo
(1895-96).

A fine collection of 772 insects of many Orders from the
Towranna Plains, W. Australia (1898), was purchased from
E. Clement, Esq., Ph.D. Dr. Clement kindly permitted a
selection of the specimens required by the Department for
a very moderate remuneration. The collection is especially
rich in moths, but also includes a fine set of Australian wasps
for the Mimicry Series.

Nineteen moths from St. Helen’s, Isle of Wight (1899), were
presented to the General Collection by Mr. W. Holland and
Mr. A. H. Hamm, and 81 insects of various Orders from the
same locality by Mr. Hamm.

A valuable collection of 162 insects from Nairobi, and
between Nairobi and Mount Kenia, British East Africa (1899),
was presented by H. J. Mackinder, Esq., M.A., Christ Church,
and C. B. Hausburg, Esq. The specimens were excellently
collected and with full data. They include some species
which are new to science, and all are of great value to the
Hope Collection, on account of their rarity or the fact that
they are unrepresented from this interesting locality.

One of the most important accessions of recent years is the
generous gift of a fine collection of butterflies, containing 805
specimens, from Karwar, Bombay Presidency (1894-97), by
G. Keatinge, Esq. Large numbers of the specimens were
bred from the larvae, and almost all are in the most beautiful
condition, with data far superior to all but a few of our Indian
insects.

Lady Denton presented a scorpion and a Cicada from Ilaro,
near Lagos Island, W. Africa (1899).

The Rev. G. B. Simeon presented a specimen of Axosia
plexippus from Hawaii (1885).

Sixty butterflies from Switzerland, Valais (1899), were
presented by H. M. Wallis, Esq.

Thirty-two insects of various Orders from Switzerland,
Valais (1897), were presented by Mrs. E. C. Bazett.

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15

Two moths from the Khasia Hills, Assam (18g0- 9°), were
presented by the Rev. G. Dexter Allen, B.A.

Three butterflies from the Oriental Region (1890-96) were
presented by Mr. W. Holland.

Seven butterflies and a moth from various localities were
presented by Col. Swinhoe, Hon. M.A., Wadham College.

A fine collection of 293 butterflies, moths, and other insects,
from Ecuador (1896-97), was purchased from Mr. W. F.
Rosenberg. Many of these will be of great value for the
Mimicry Collection.

The collection of Phytophaga has been enriched by 1.475
specimens selected from the duplicates of the British Museum
(Natural History). Nearly the whole of these have been
_named and placed in the part of the collection which is now
completely arranged.

In addition to the accessions recorded above, a large number
still remain uncatalogued, and are now provisionally acknow-
ledged.

The Curators of the Indian Institute presented the
collection of insects formerly in the Museum of the East
Indian Company, together with the cabinets in which they
were contained. Although the condition of the majority of
the specimens is poor, many of them will be of value to the
Hope Collection.

The British Museum of Natural History has presented a
very large number of butterflies selected from among their
duplicates.

The very fine collection of S. African moths made by Cecil
Barker, Esq., of Malvern, Durban, Natal, was presented by
Roland Trimen, Esq., Hon. M.A., F.R.S.; also butterflies
from S. Europe.

E. S. Goodrich, Esq., M.A., Merton College, presented the
insects captured by him in Ceylon (1899), accompanied by
excellent data.

Herbert Druce, Esq., F.L.S., presented a fine collection of
moths chiefly from Central America.

16

Dr. T. A. Chapman presented a valuable set of Lepido-
pteras, chiefly Erebias, from Switzerland and Norway.

A collection of butterflies from Banff, Canada, was pre-
sented by H. J. Elwes, Esq., F.R.S.

Five butterflies captured at sea and Lepidoptera from various
European localities were presented by J. W. Tutt, Esq.

A very fine series of Lepidoptera was collected in Siam by
Richard Evans,:\ Esq., -B.A.,” jesus, Collese” Mir vans
proposes to work them out in the Hope Department, and
will present many of the specimens to the Hope Collection.

A large number of insects of various Orders were presented
by G. C. Griffiths, Esq., from Japan, Sumatra, the Moluccas,
the Khasia Hills and other Indian localities, Zomba,
N. Queensland, Mexico, Switzerland.

Colonel J. W. Yerbury presented a fine collection of
Diptera (Syrphidae) made by him at Aden. This valuable
collection includes many types. He also presented specimens
of Diptera for the British Collections, and examples of Diptera
resembling Hymenoptera for the Mimicry Series.

A fine series of insects, chiefly from British North Borneo,
illustrating mimicry, and allied subjects, was presented by
R. Shelford, Esq., B.A. (Cantab.), of the Sarawak Museum,
British N. Borneo. Mr. Shelford also presented a very fine
series of insects from the same district for the General
Collection.

Small collections of insects from the localities indicated were
also presented by the following donors:—From Uganda, by
Mrs. Bazett; from Switzerland, by Hugh Richardson, Esq. ;
from Norway, by E. N. Bennett, Esq., M.A., Hertford College ;
from various localities, especially W. Africa, by Mark L.
Sykes, Esq.; from the United States, by Abbott H. Thayer,
Esq.; from Fiji and Vancouver, B.C., by Professor Gilson ;
from Darjeeling, by Henry Ward, Esq.; from Europe, by
D. Chaplin, Esq., M.A., University College; from Sylhet, by
W. Hatchett Jackson, Esq., M.A., Keble College; from
Mexico, by Osbert H. Howarth, Esq.; from Brittany, by
Professor Poulton, M.A., Jesus College; from Vancouver

ne

17

Island, by Mr. A. H. Hamm; from California, by Ronald W.
Poulton; from Europe, by F. Merrifield, Esq.; and from
Europe, by Professor R. Meldola, F.R.S.

The additions to the collections of British Insects in 1899
have been very numerous and valuable, due especially to
a week’s visit of the staff of the Department to St. Helen’s,
Isle of Wight, at the beginning of July, 1899. As a result
of untiring work the following specimens have been incor-
porated. Mr. W. Holland presented 243 insects, chiefly
Coleoptera, many of them of great rarity, exclusive of a large
number which he will add at a later period. Mr. A. H. Hamm
presented 587 insects, chiefly Hymenoptera, many being very
rare and interesting species. Together they presented 68
moths. Professor Poulton also presented 19 insects, while
9 others were given by all three naturalists. All the specimens
have very full data, and area most important accession. Many
specimens will be added to the series illustrating the natural
history of insects.

A wasps’ nest found at Headington Hill (1898) was pre-
sented by Hugh F. Hall, Esq.

An interesting specimen of a large locust (Acridium
aegyptium) was presented by Mr. J. Gee, having been intro-
duced into Oxford from Italy in broccoli. The insect was
alive when brought to the Department (Feb. 8, 1899).

A pair of very rare Hymenopterous insects (A stata stigma),
from Brandon (1898), was presented by R. C. L. Perkins,
Esq., B.A., Jesus College.

Thirty-nine insects were presented by J. J. Walker, Esq.,
R.N., including rare Coleoptera from Kent and S. Devon
(1898-9), 3 Colias Hyale from Sheppey (1899), and 25
Lepidoptera from the neighbourhood of Rannoch (1899).

Eighty Lepidoptera, captured by Mr. William Reid in
N. Shetland and various localities in Scotland (1895-98),
were presented by G. C. Griffiths, Esq., together with a pair
of the rare Platypteryx sicula (1899), confined in this country
to the Leigh Woods, Bristol.

A dragon-fly from the neighbourhood of Oxford was pre-
sented by Dr. F. A. Dixey, M.A., Wadham College.

18

Two wasps and a mimetic fly captured at Dorking, on the
same flower-heads, were presented by Edward Saunders, Esq.,
F.L.S.: for the Mimicry Series: ~:

Many further accessions to the British Collections have not
yet been catalogued. °

A set of moths from N. Cornwall were presented by A. G.
Cardew, Esq., M.A., Queen’s College. A valuable set of rare
British Hymenoptera, filling up many gaps in the Hope
Collection, was presented by Edward Saunders, Esq. British
Coleoptera, with excellent data, and many of them very rare
species, were presented by Horace St. J. Donisthorpe, Esq.

Small numbers of British insects were also presented by the
following donors:—Mrs. A. G. Butler, Miss Ruth Butler,
Miss C. V. Butler, Miss Janet Poulton, Miss C. B. Sanders,
Miss Mabel E. Notley, Miss F. A. Wright, Professor E. A.
Minchin, Professor E. B. Poulton, Major R. B. Robertson,
Mr. Henry Higgs, Mr. W. Holland, Mr. C. J. Bayzand, Mr. A.
Overington, Mr. A. H. Hamm, Mr. Edward Webb, Mr. A. Drew,
Mr. A. Robinson, Mr. H. Trim, and Ronald W. Poulton.

ADDITIONS TO THE HOPE LIBRARY? EN aégee

The Trustees of the British Museum presented the “ Cata-
logue of the Lepidoptera Phalaenae,” vol. i., by Sir George F.
Hampson.

The Smithsonian Institution (United States National
Museum, Washington) presented the publications which deal
with the subjects of the Department.

The Boston Society of Natural History and the Bombay
Natural History Society presented their publications for the
year.

The University of the State of New York presented its
Reports.

The. Linacre Reports, vol. iv, were presented by Prof. E.
Ray Lankester, F.R.S.

The Radcliffe Librarian, Oxford, presented the Catalogue
of Books added during 1898.

19

The Transactions of the Entomological Society, and the
Transactions and Journal of the Linnean Society for the year
1899, were presented by Professor Poulton.

“Fauna Hawaiiensis,” vol. i, part 1, and vol. ii, parts 1 and 2,
were presented by the author, R. C. L. Perkins, Esq., B.A.,
Jesus College.

**Colouration in Animals and Plants,” and “On the Growth
Ol ureeswcce, by the late Alfred Tylor, Esq. F:G:S., were
presented by Mrs. Alfred Tylor.

Miss E. A. Ormerod presented her valuable “ Report of
Observations of Injurious Insects, &c.,” for 1898 (22nd Report),
and for 1899 (23rd Report).

Copies of original papers were presented by the following
authors :—Herbert Druce, Esq., F.L.S.; W. F. Kirby, Esgq.,
Pele. j. Elwes, sq. Y.R:S.3; Dr. A. -G. Butler; Rev:
spe Stepping, M.A; F.RiS., Worcester College; S: H:
Scudder, Esq.; R. W. T. Giinther, Esq., M.A., Magdalen Col-
lege; Horace St. J. Donisthorpe, Esq., F.E.S.; G. W. Kirkaldy,
Pegnerebase: G. C. Bionell, Esq. P.E.S.3. Professor Roland
Thaxter, Harvard Univ.; and Sir George F. Hampson.

A large and valuable set of original papers has been pre-
sented by Lionel de Nicéville, Esq., F.E.S.

The Committee of the Bristol Museum presented their
Report for 1896-98. ©

A paper on the Collembola, &c., of Devon, by Edward
Parfitt, was presented by F. J. Haverfield, Esq., M.A., Christ
Church.

A few second-hand books and papers were purchased,
together with the Zoological Record, the parts of Barrett’s
‘British Lepidoptera,” the Ray Society volume, and the parts
of Rippon’s.“ Icones Ornithopterarum”’ for the year.

EDWARD BoPOULTON,

Si Sav egos SMITHSONIAN INSTITUTIO

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