> en ip, a — -_ San ta eae RI See ; aay Ae Puate IV. Eaas or THESTORIDS AND POLYOMMATIDS. Natural History of British Butterflies, January, 1906. Puate IV. (To be bound facing Plate IV.) Haes or RURALIDS. Fig. 1.—CALLOPHRYS RUBI. Fic. 4.—PoLYoMMATUS ICARUS. Fic. 2.—CALLOPHRYS RUBI. Fig. 5.—AGRIADES BELLARGUS. Fic. 3.—ARICIA var. ARTAXERXES. Fic. 6.—AGRIADES CORYDON. All x20 diameters. t 7 — P — pen wa 1S ay ally EE v.83 Gad oy ae A NATURAL HISTORY OF THE BRITISH LEPIDOPTERA ne nr BOOK FOR STUDENTS AND COLLECTORS BY qme? {Lam Nine ole ele es ES. Author of ‘‘ The British Noctuz and their Varieties,’’ ‘‘ Monograph of the British Pterophorina,’’ ‘‘ British Butterflies,’ ‘‘ British Moths,”’ ‘Migration and Dispersal of Insects,’’ ‘‘ Melanism and Melanochroism in Lepidoptera,’’ ‘‘ Practical Hints for the Field Lepidopterist,’’ etc. ZEM\THSONIARS VoL. VILA OCT 95 VI BERLIN: © FRIEDLANDER & SOHN, 11, Carlstrasse, N.W. NOVEMBER, 1905—DECEMBER, 1906. PREFACE. In presenting this volume to my brother lepidopterists, some little explanation is necessary. Although itis essentially vol. vii of The Natural History of the British Lepidoptera, it has been appearing for a considerable time in parts as A’ Natural History of the British Butterflies. The real reason for publishing this volume out of its proper order was threefold—(1) The large amount of material on the group that had been slowly amassing during the last twenty years, and the increasing difficulty of dealing with it; (2) the fact that a really good scientific work on British butterflies was an undoubted desideratum among advanced workers; and (3) the long time which must necessarily elapse before the material to be dealt with in the two intermediate volumes. vi and vii, can possibly be worked out and prepared for publication; volume v is being cleared up for publication contemporaneously with this. It is trusted that these reasons will be considered sufficient to excuse my action. It is exceedingly difficult to foretell the extent of detailed and exhaustive treatises of this character. When one is working at a eroup, one accumulates material from all possible sources, and such detail can only be finally estimated when the printer has set it in type, and one sees the actual printed matter before one. That I should only be able to treat of ten species in a large volume would, two years ago, have appeared to me absurd, and that an account of Rumicia phlaeas would extend to 84 pages, or 10 pages more than our account of Manduca. atropos in vol. iv, would have been considered impossible, but, as one pieces all the facts together, one finds the accounts of some well-known species growing beyond all previously calculated dimensions, whilst, of others, one is astonished at the necessity of working out, almost de novo, the whole of the life-history, and the trouble attached to this is not to be despised. The practical completion of the life-histories of Chrysophanus dispar, Thymelicus acteon, Urbicola comma, ete., are the result of inuch painstaking and careful work on the part of my collaborators, and the detailed account of the world- wide variation of such species as Rumicia phlaeas, Urbicola comma, and Cyclopides palaemon, should bring home a recognition of the importance and bearings of a knowledge of species outside the narrow boundaries of our own islands. If, in all the preceding volumes, I have happily had to acknowledge much generous help, in this my obligations have been increased tenfold. As usual, to Dr. T. A. Chapman and Mr. A. W. Bacot, first thanks are due, but in no smaller degree also to Mr. A. Sich, Mr. M. Gillmer, and Mr. S. Edwards; whilst no less am I indebted to Mr. F. Noad Clark, Mr. A. Tonge, and Mr. Hugh Main, to whose kindness I owe almost all the beautiful photographs by means of which the volume is illustrated. Mr. H. Rowland-Brown, too, has done yeoman service with regard to our distribution lists, and there are many more whom one ought certainly to mention— ill. Professor Blachier, Mr. A. W. Kappel, Mr. W. F. Kirby, Paymaster-in- Chief G. F. Mathew, Rev. G. H. Raynor, Mr. Raleigh S. Smallman, etc. Jt is impossible to mention all those to whom we are further indebted for incidental observations, notes, and lists of localities. I can only say that without their kind help the volume, as it now stands, could never have appeared. 1 may add that I have had no hesitation in writing for information to any lepidopterist, at home or abroad, who, [thought, might help me to clear up any point concerning any insect of which he might have special knowledge. In every case my queries have been most courteously answered. I trust that such will not mind if in the near future I have to worry them again. For the index we are largely indebted to the Rev. G. H. Raynor, who kindly responded to our request for help in this direction. I sincerely trust that the result of the work expended on this volume will commend itself alike to field naturalists and scientific lepidopterists, and that it will meet with their approval. Information on the British species of Theclids and Lycznids is greatly desiderated, and any such will be most gratefully received. Our knowledge of some of the commonest species of ‘‘ blues” is of the most scrappy character, and material to complete in detail our knowledge of their life-histories and habits is urgently needed. We shall be most thank- ful for any help in this direction. CONTENTS. Part I. CHAP. PAGE. I. GENERAL OBSERVATIONS ON BUTTERFLIES ... pee me aS oa 1 IJ. EGGLAYING OF BUTTERFLIES ap beat : ee oA. ws 2 III. EGGS OF BUTTERFLIES bts ee ae Ae sf ee = 5 IV. PHOTOGRAPHING BUTTERFLY EGGS ... oe ie a a “de 8 V. OBTAINING EGGS OF BUTTERFLIES ... = be i ae cat 12 VI. BUTTERFLY LARVE AND THEIR MOULTINGS.. a er: io ae VII. EXTERNAL STRUCTURE OF THE BUTTERFLY LARVA... an ae ow eS VIII. INTERNAL STRUCTURE OF THE BUTTERFLY LARVA... .. ... nee cay eee IX. THE ASSOCIATION OF ANTS WITH BUTTERFLY LARVE ee ee a. ew X. CARNIVOROUS HABITS OF BUTTERFLY LARVE pe oe a Mee XI. COLLECTING BUTTERFLY LARVZ ae yee Bi ee Joe eee XII. THE SILK-SPINNING HABIT OF BUTTERFLY LARVAE ae ee a ae 50 XIII. THE COLORATION OF BUTTERFLY LARVE ... a ee 61 XIV.. THE RESTING-HABITS OF BUTTERFLY LARVE ae bee Ae, _12- 78 Part II. URBICOLIDES oe oe ee bo oe: ee tie 4 eee URBICOLIDE . e wee ee. a tat a es THYMELICINE ‘AND THYMELIOIDI oie aa Me bulk ae i Se ADOPHA LINEOLA ... a te Bre ae ae rae i ee ADOPHA FLAVA nee ae ‘gal ees ae nh ie Sas THYMELIOUS . es a ee Ges f wa aE ce MS THYMELICUS ACTEON ae ae SE pei ae ne a jy sleet, URBICOLINA AND URBICOLIDI ae i ae ae Ps oo Be SKUGIADES © 42, : a Sa ae att = 2 ee AUGIADES SYLVANUS.. na ie ike eas ae Wee ul Shee. URBICOLA: ... he za ews ae oe a ae Ras irs URBICOLA COMMA ... af ae Ae se Re. yo nae OYCLOPIDID® Wek ie sie ae ae of ee bt) CYCLOPIDINE AND CYOLOPIDIDI ath ae ue ae oe 4 @on CYCLOPIDES ... ; Me ae a ae eee fet ee kee OYCLOPIDES PALEMON He ce as 4s & _ ac} Age HESPERIIDA ... es ay ~~ i my cae a. Ku a HESPERIINAE ... a he he re oh ae re see HESPERIIDI ... ay eae an vs Be ne se ao the HESPERIA... ies a a Hes ee fee me a oe HESPERIA MALVE ... a, ae er ae Bee Hip aoe NISONIADIDL ... ne a a ee ae a xg vas’ eee NISONIADES ... oe cae tig A tee Res. ak ak oe NISONIADES TAGES.. : aa mae ba A a gene CATALOGUE OF PALEAROTIO URBICOLIDES Sch ue a stg Were 5 RURALIDES ... a ett si ie ae ca aS 4 ee RURALIDE ... or oh i fe a PO CHRYSOPHANIN A AND CHRYSOPHANIDI ve ne a As ce) ke RUMICIA Be et ae si ob ae a a ee RUMIOIA PHLHAS ... oe =n a ek a an ad OU CHRYSOPHANUS ae ae rat ss a a ay se ee OHRYSOPHANUS DISPAR xe si ae ee 2 ve eal: ee, GENERAL INDEX TO PARTS I. AND II. ir Shs rn =F > ee SPECIAL INDEX TO PART II. oe a a i me .. 468 REFERENCES TO PLATES ie: EP bi ioe i a . ate Seetrtst) BUTTERFLIES. CHAPTER LI. GENERAL OBSERVATIONS ON BUTTERFLIES. Butterflies and moths together make up the order of insects known as Lepidoptera (scale-winged insects), so called because their wings are covered with delicate scales which are very easily removed. The popular idea is that there are profound differences between butterflies and moths, but this is not so, the butterflies comprising simply two of the superfamilies of the “‘upright-egged”’ stirps of the lepidoptera, and finding probably their closest relatives in the Castniids, Noto- dontids, Noctuids, etc. There is, indeed, no real line of demarcation between butterflies and moths, those superfamilies of moths just mentioned being much nearer to the butterflies than to many other superfamilies of moths. The general characters that are supposed to distinguish butterflies are (1) The knobbed antenne. (2) Resting with their wings upright, ¢.e., with their wings raised vertically face to face over their bodies. (8) The thorax and abdomen being separated by a waist, etc. Some moths, however, have knobbed antenne, others rest with their wings upright over their backs, and yet others have the waist between the thorax and abdomen more marked than in many butterflies. The fact is that all lepidoptera have been evolved from the same original stock, the various superfamilies having been produced by modifications and changes taking place through the course of ages, yet all, different as they now are, bearing the stamp of a common origin. The differences have no doubt been brought about by changes in environment, the individuals having to adapt themselves to the great physical changes that we know the earth and its atmosphere have undergone in past time. Geological remains of insects are scarce, due no doubt to the fragile nature of the organisms themselves. Yet there is much fragmentary testimony of the history of insects written in those remains that have been found. Butterflies and moths are among the more recently evolved insects, and appear to have been contemporaneous with flowering plants, the newer rocks of the Tertiary period providing most fossil examples. Although above 30000 fossil insects have been found in the tertiary beds of Europe and America, only about 20 examples of lepidoptera have been found. Scudder notes that over 15000 fossil 2, BRITISH BUTTERFLIES. insects have been found in the neighbourhood of the small lake of Florissant, high up in the Colorado Parks, yet he has only seen eight butterflies among them, each belonging to different genera. He further notes that this is also the case with the fossil butterflies found at Radoboj, at Aix, and at Rott in the European tertiaries, and with two exceptions, Hugonia and Pontia, no representative of genera found today has been discovered. The fossil species are all extinct. It is largely a matter of speculation, from which of the main stirpes or root-stocks of the lepidoptera the butterflies have sprung, but they have, as we have already noticed, upright eggs in common with Castniids, Noctuids, etc., and the arrangement of the tubercles and the presence of the remarkable chinglands of their larve also suggest that these are among their nearest allies at the present time. The matter is just here of less importance than the knowledge that, since the butterflies branched off on their own account, they have developed, in various directions, strongly marked characters of their own, and that now, each of the two, usually-accepted, large superfamily groups, Urbi- colides or Skippers and Papilionides or Papilios, 1s subdivided into large groups or families, some containing vast numbers of species exhibiting very varying degrees of specialisation. Some butterflies are very generalised in their structure, t.e., they show a number of very simple characters which are commonly observed among a large number of other lepidopterous insects, others again are exceedingly specialised and show some well-marked peculiarity very strongly. Of our two large groups the Urbicolides are assumed to be more generalised than the Papilionides, although, compared with many other superfamilies, the Urbicolides are very highly specialised. Whilst, therefore, there are marked differences between the super- families and families into which butterflies are subdivided, it will be readily understood that they are, in reality, very closely allied to the moths. No hard and fast lines of distinction can be shown to exist ; variation is the basis of all progress in evolution, and the student, therefore, must look for differences as well as similarities, and try to fathom the meaning of both when he observes them. Similarities in some one particular do not always denote close relationship between species, they may be mere analogies and not homologies, whilst marked differences do usually denote considerable separation and more distant relationship. CHAPTER. If. EGGLAYING OF BUTTERFLIES. The egelaying habits of butterflies are very varied, yet almost absolutely constant for the same species. So much is this so that, in many instances, the butterfly selects almost exactly the same portion of the plant on which to deposit her eggs, the upperside of a leaf, the underside of a leaf, the pedicel of a flower, the sheathing part of a erass blade, ete. Thus it has been noted (Ent. Rec., iv., p. 225) that, whenever the ?s of the early brood of Cyaniris argiolus choose holly on which to lay their eges, they almost always lay them on the calyx, EGGLAYING OF BUTTERFLIES. 3 and it is further observed how important this is to the insect, for should they lay them a fraction of an inch higher up, they would almost certainly be destroyed, as, when the buds once open, the petals are very easily blown away by the wind. Similarly it is noted (op. cit., xii., p. 269) that, when the ?s of the late brood of this species lay their eggs on the ivy-umbels, the base of the calyx is also almost always chosen. It is remarkable that, if the twigs are chosen instead of the flowers, the eggs are laid much more indiscriminately. In the same manner both the 9s of the early brood of this species, and of Callophrys rubi, frequently choose Rhamnus frangula, and, when this is so, the base of the calyx appears again to be almost invariably chosen. Similarly, the eges of Nemeobius lucina are laid on the undersides of primrose leaves, generally one on each leaf, but two, three, four or even five are sometimes to be found on the underside of the same leaf, possibly laid by different ? s, but it is the underside and not the upper- side of the leaf that is practically always chosen. As we propose giving a separate paragraph on the mode of egglaying of each of our British species in the systematic part of this work, there is no need to give a large number of examples here. One of the most remarkable facts connected with the egglaying of butterflies is what may be termed the botanical instinct. Species whose larve are usually confined to one or two plants will sometimes, under the stress of necessity, select an introduced plant, and.it is remarkable that, in most cases, this will be a plant closely allied botanically to its natural foods. The ¢ Huchloé cardamines chooses the base of the footstalk of a somewhat passé flower of Cardamine pratensis on which to lay its yellow egg. Failing this it will accept Alliaria officinalis, and other common Cruciferae, and, if it enters a kitchen- garden will utilise the flowers of horse-radish. Never by any chance does it select any plant unless it belongs to this natural order. Similarly the 9s of Gonepteryx rhamni choose the underside of the leaf of a Rhamnus bush, laying the egg on one of the veins, and passing by all other plants in the hedge or woodside, however similar they may appear. This selection appears to be made by the sense of smell, and the accuracy with which the selection is made is sometimes very remarkable. Fritz Muller cites some curious instances in which butterflies appear to have recognised the affinities of certain plants before they had been discovered by botanists, ¢.y., he says (Nature, xxx., p. 240): “ The caterpillars of Mechanitis, Dircenna, Ceratinia and Ithomia, feed on different species of solanaceous plants (Solanum, Cyphomandra, Bassovia, Cestrum), those of the allied genus Thyridia on Brunfelsia. Now this latter genus of plants had been placed unanimously by botanists among the Scrophulariaceae, until, quite recently, it was transferred by Bentham and Hooker to the Solanaceae. It thus appears that butterflies recognised the true affinity of Brunfelsia long before botanists did so. Another and more curious instance of butterflies confirming the arrangement of plants in the Genera Plantarum is exemplified by Ageronia and Didonis, which were formerly widely separated by lepidopterists, being even considered as belonging to distinct families, but now placed near each other in the Nymphalids, the larve leaving no doubt of their close affinity. The larve of Ageronia feed on Dalechampia, those of Didonis on Tragia. These two euphorbiaceous genera were widely separated by Endlicher, who placed 4 BRITISH BUTTERFLIES. the former among the Huphorbiaceae, the latter among the Acalyphaceae, Bentham and Hooker, on the contrary, place them close together in the Plukenetieae, so that their close affinity which had been duly appreciated by butterflies has finally been recognised by botanists also.”’ Another remarkable fact connected with this subject is the narrow range of plants from which some species have to select and the wide range of others. Aglais urticae is confined to stinging-nettle, Limenitis sibylla to honeysuckle, yet Polygonia c-album has foodplants as widely different as stinging-nettle, black-currant and hop, and Pyrameis cardui as stinging-nettle, thistle, mallow and_ bugloss. Still, on the whole, our butterflies have a very small range of foodplants compared with that of many moths, some of which appear to be more or less polyphagous, e.g., Saturnia pavonia (Brit. Lep., l., p. 838), Manduca atropos (op. cit., iv., pp. 482-3). In America, however, Scudder notes that the ‘‘swallow-tail,’ Jasoniades glaucus, feeds upon plants belonging to no fewer than 15 different families. Not only is the range of plants which a species chooses as food for its larvee very restricted, but there is usually a very close alliance between the plants selected by the ¢?s of all the species of the same family, e.y., all the European Urbicolid skippers choose grasses and rushes of various kinds, the Hesperiid skippers on the other hand have a much wider range among Malvaceae, Rosaceae, Lequminoseae, ete. ; the Chrysophanids prefer plants belonging to Polygonaceae, the Lycenids and Cohads choose Leguminoseae, the Argynnids use Violaceae, the Pierids Cruciferae, the Coenonymphids, Satyrids and Erebiids, erasses,and so on. It will be known to most lepidopterists that these general statements are largely true of butterflies belonging to these eroups throughout the whole of the Palearctic and Nearctic regions. We have already noted that the egglaying habit of butterflies is usually constant and fixed even to the extent of considerable detail. Almost all butterflies lay their eggs definitely on their foodplant, the only British species which appears not to do so being Melanargia galathea, which drops its eggs loosely among the roots of the grasses on which the larve live. Occasionally a 2 of a species with a fairly constant egelaying habit appears to blunder and will choose a dead stem of grass or other object in the immediate neighbourhood of the foodplant, but the occasions on which this happens are rare, and the choice of the foodplant and the selection of an exact position in so many cases are instinctive processes which have been brought to a high stage of perfection in the development of the race. Chrysophanus virgaureae chooses a dead or dying stem on which to lay its hybernating ego. The choice of the foodplant, as we have already noted, is almost certainly effected by means of scent. The antennal sensory organs (among which those of scent are included) have been worked out in detail (Hnt. Record, viii., pp. 225, 261). We know that the sense of vision is exceedingly limited in butterflies, and experiment is against the assumption that they use their power of sight for any such delicate discrimination as is required when they are selecting a special species of foodplant on which to lay their eggs. On the other hand, their movements, when on this important business, are such that it would seem obvious that they are seeking by scent the plants on which to lay. A @ will often settle on a dozen different plants similar and dissimilar to that wanted before she finds the right one on which to EGGS OF BUTTERFLIES. 5 lay her eggs. On the other hand, some will fly rapidly from plant to plant when on egglaying intent without much delay, and especially is this the case where the species is local and the foodplant abundant, e.g., Polyommatus bellargus, etc. CHAPTER III. EGGS OF BUTTERFLIES. Lepidoptera are holometabolic insects, 7.e., they are insects that have a complete metamorphosis, assuming in turn, the egg, larval, pupal and imaginal stages. The eggs of butterflies are naturally of small size, and photographing them has added much charm to the description of these beautiful objects. Under an ordinary lens much of their remarkable beauty may be discovered, but for a proper appreci- ation of their lovely forms, delicate sculpturing and detailed structure, a good microscope is necessary. The egg itself is a cell, consisting of an outside shell, enclosing the living protoplasm which is, at first, homogeneous; the shell is a thin, elastic pellicle, often transparent, and, when opaque, usually made so by the ribs that longitudinally and transversely cross its surface, although, in some cases, aS in some of the Urbicolids, etc., there is a general thickening of the wall. The butterfly egg is of the ‘“‘upright’’ type, 7.e., it has its micropylar axis perpendicular to the surface on which it is laid, whilst its transverse section (at right angles to the micropylar axis) almost always forms a circle (the only exceptions known being those of the Thymelicids), and its base is almost always flattened at the point of attachment. At the apex or point opposite the base, a number of microscopic canals lead into the interior of the egg; these canals are surrounded by a rosette of tiny cells, the whole structure being termed the micropyle. Through these little canals the spermatozoa pass to fertilise the egg. In all the species of any given family, the eges are almost always very similar, although the details vary in each species. The Thymelicid eggs are “flat”? in outline, with three axes of different lengths, whilst all other butterfly eggs have a horizontal section that is circular ; the Urbicolid (sens. rest.) skipper eggs are somewhat more than hemispherical, those of the Lycenids are shallow, flattened, tiarate or echinoid in shape, those of the Papilionids and some Satyrids almost globular, of the Pierids, long, slender and spindle-shaped, of the Nymphalids somewhat barrel-shaped, with projecting ribs reaching from apex to base, of most of the Satyrids cylindrical or spheroidal, etc. The barrel-shaped egg is sometimes more or less conical, and it is usually rather broader at the base than at the summit. The surface of the egg may be practically smooth, reticu- lated with a surface sculpture, or ribbed longitudinally from base to apex, and often with finer transverse ribs runniny round and crossing the longitudinal ones. These various modes of sculpture run into one another, and are often not sharply defined, and in the Lycenid and Limenitid eggs the sculpture forms patterns and traceries of exquisite beauty. The ribbing generally fades off into the micropylar area, 6 BRITISH BUTTERFLIES. usually a slight depression surrounding the micropyle proper, of which mention has already been made. It may be well here to remark that whilst the Urbicolid (Pamphilid) group of skippers tends to smooth eggs, forming rather more than a hemisphere, those of the Hesperiid group are hemispherical, with well-defined, clearly-cut, strongly projecting longitudinal ribs, which give them an appearance much more resembling that of the eggs of Argynnids than the other skippers. We have already noted that, in the same group, the Thymelicids have flat eggs, super- ficially like those of many moths, except that the micropylar axis is upright. The number of ribs running from the base to the apex varies considerably, in the Vanessids they may be reduced to eight, and rarely have more than twelve, in the Coliads there may be as many as thirty or forty. The eggs of butterflies are usually of a pale yellow or aan ereenish colour when first laid, but those with transparent shells change their colour very rapidly, the tint becoming quickly that which will best preserve each in the position in which it is laid. It is usually placed on a leaf, and then hatches in a few days, but if the eggstage lasts for any length of time, the position chosen is on a twig or other permanent part of the plant. The ege of Huchloé cardamines is pale yellow when laid, becomes deep orange in about twenty-four hours; and remains of this tint till just before the larva hatches, when the embryo can be seen coiled up within the transparent eggshell. Other eggs change from yellow to pink, or brown, or pale salmon, and then to greyish in different species. As the young larve mature within the egg, there isa considerable change observable inside in those of many species, due to the development of the embryo within. These changes can, in some of the more transparent-shelled species, be readily traced under a microscope, - e.g., Pararge megaera, Nemeobius lucina, etc. Those British butterflies whose eges go through the winter are few in number, viz., Argynnis adippe, Thecla w-album, T. pruni, Zephyrus betulae, Z. quercus, Urbicola comma, Adopaea lineola and Thymelicus acteon. The hybernating stage of Lampides boetica is very uncertain. The old myth of the eggs of Aporia crataegi (teste Rennie and other authors) going over three winters and then hatching, is absurd. The eges of butterflies are usually laid singly, rarely more than two or three on a leaf, but a few British species, e.g., Aporia crataeyi, Pieris brassicae, Melitaea cinaia, M. athalia, M. aurinia, Vanessa io, Aylais urticae, Hugonia polychloros, and Euvanessa antiopa, and many of their Continental and exotic allies, lay their eggs in clusters, either as regular rows or irregular heaps, on a twig or the upper- or under- side of a leaf. Some of “the Polygonias lay their eggs upon each other in a string, as it were, from three to ten eges in a ‘single file. Eggs are subject to many dangers after being laid. A particular group of Hymenoptera, called Chalcids, lay their eggs inside the eggs of lepidopterous insects, and find within them sufficient nutriment to come to perfection ; in fact, a dozen or more perfect Proctotrupids some- times emerge from a single moderate-sized egg, having passed their entire existence within, and obtained the whole of their nutriment from the contents of, this tiny receptacle. It has already been noted that the egg consists of an outside shell and its protoplasmic contents. After fertilisation, the time varying much with the species, the contents begin to thicken visibly and to EGGS OF BUTTERFLIES. ih undergo development, which can, in some species, be readily observed through a microscope. The first cells formed unite together, and present some parts darker than others, and after a time the fluid mass - breaks away from the eggshell, and a tubular structure, which soon becomes deeply segmented, forms within. The rings become distinct, and those which form the head (apparently four in number) are at this time much larger than the others. As development proceeds these get welded together, and the hard mouthparts become visible. At the same time the three segments forming the thorax have little cells developed on their outside. These are the rudiments of the legs. Then a great change takes place in the position of the embryo whose _ development has thus far been traced. Up to this point, the embryo has been lying in a somewhat circular form with the legs outside, ¢.e., towards the eggshell, but now it gradually changes its position until the body is somewhat S-shaped, the movement continuing until a complete reversal has taken place and the embryo has returned once more to a circular position, but with the legs now pointing towards the centre of the egg. At this time, distinct patches appear on the cheeks, and gradually six black spots develop on each dark patch. These are the ocelli, five of which are arranged in lunular form, the sixth being isolated and at some little distance from the concave side of the others. The ocelli are simple lenses, and very different from the complicated structure of the com- pound eye of the imago. During the time that this has been going on, structural changes have been taking place inside the embryo. A hollow sac forms along the back, and after a time this keeps up a regular pulsation. This is the dorsal vessel, and the centre of the circulatory (blood) system. The alimentary canal also becomes traceable, and possibly, whilst you are watching, you may observe tiny silvery threads suddenly come into view, which look as if they traverse the larva in all directions; this appearance is due to the sudden expansion of the air-vessels which, as we can now see, start from the little spiracles along the sides of the body, this respiratory (air) system having been invisible hitherto because of their transparency. After this, little lines gradually appear, crossing the embryo in different directions, and usually traceable to little chitinous buttons from which they arise; these are the hairs or primary sete. Certain marks on the embryo also become distinct, and other peculiarities may possibly be noticed. At last the embryo is not noticed to undergo any further change; its jaws are seen to move steadily to and fro against a particular part of the eggshell, usually at or very near the micropyle, until a little hole is made in it, the edge of which the contained larva continues to nibble until it can squeeze its body through the aperture. It is well to remember that in some of the hybernating eggs the larva is formed in the early autumn, and remains all the winter in the egg, hatching only with the spring, e.y., the egg of Argynnis adippe, laid in July, has its embryo fully developed in less than a month, but does not hatch until the following March, the egg of Adopaea lineola, and possibly others, is in similar case. 8 BRITISH BUTTERFLIES. CHAPTER IV. PHOTOGRAPHING BUTTERFLY EGGS.* My first attempts at photographing ova were made with an ordinary 4-plate stand camera, focussing from the back, attached to a student’s microscope, and with this I was able to obtain results which surprised me, but was handicapped by the short extension obtainable with the camera (about 15in.), as this necessitated using the eyepiece of the microscope to bring the image to a focus at a point nearer the objec- tive than would otherwise be necessary. This caused some loss in definition, and I also had considerable difficulty in getting an objective of long enough focus to cover the whole of a small batch of ova and at the same time admit of the object being focussed with the eyepiece I possessed. Having obtained a 3in. objective which filled these neces- sary requirements, the next step was to construct a base board on which to fix the camera and microscope in proper alignment. For this purpose I used a smooth deal board about 3ft. long, 6in. wide, and lin. thick, and ruled a straight line down the centre from end to end as a guide to obtaining a proper alignment of the completed apparatus. On placing the camera and microscope upon it, with the body tube of the latter lowered to a horizontal position, I found that the line of projection was considerably below the lens aperture in the camera front, so the next step was to construct a block for the micro- scope to stand upon, which would raise it sufficiently to bring the eyepiece exactly in the centre of the lens aperture, and permit of an imaginary line being drawn horizontally from the centre of the objective to the centre of the focussing screen of the camera, and pass centrally through the intervening portions of the microscope body and the camera. This block I screwed down firmly in the required position, and fixed small wooden stops at the sides and ends to prevent the microscope from slipping when placed on it. I then drilled a hole for the camera screw in the centre of the base board, in such a position that, when the camera was screwed down to it, the lens front just touched the eyepiece of the microscope, when the latter was racked out to its fullest extent. It was now necessary to devise some means of connecting the microscope with the camera in such a way as to exclude all light except that passing through the former, and yet to admit of focussing being done as usual. To do this I obtained from an instrument maker a short brass tube about lin. in length, with a screw thread cut on one end to fit the lens flange of the camera. This I screwed in, in place of a lens, and then made a sleeve of black velvet to slip over it, long enough to be drawn over the microscope body tube, and held in place at either end with an elastic band. This answered admirably, and I found that after focussing an object on the stage of the microscope and then con- necting the microscope with the camera, I was able to get a sharp image on the ground glass of the latter with the extension at my disposal, and to reduce or enlarge this by altering the amount of camera extension, and then to refocus with the microscope by stretching * This chapter is by Mr. A. E. Tonge, who is doing the photographs by which this book is illustrated. “GOGT ‘taquIsAON ‘saufiazng ysumwg fo huojysreT poungon “SAITHUALLAG dO SONY AHL ONIHGVUYNOLOHG YOL SALVAVddY ‘TI gLvIg PHOTOGRAPHING BUTTERFLY EGGS. 9 out my arm along the side of the camera while I kept my eyes fixed on the focussing screen. : This somewhat primitive apparatus answered well so far as it went, but I soon determined to work without the microscopic eyepiece, and set to work, therefore, to build a very long extension camera. I made it 4-plate size, so that the accessories I already possessed could be used, and gave it a bellows 42in. long in three sections to obviate sagging. I found this was a great improvement, but had, of course, to get a much larger base-board, and used for this an oak plank 5ft. x 8in.x lin. The focussing of the microscope could not now be done by hand, as the distance from the ground glass was too great for “my arm, so I carried a long brass rod through wooden blocks under the camera, and fitted a small grooved pulley-wheel on it just under the fine adjustment screw of the microscope, and put a milled screw on the other end under the focussing screen of the camera. A fine elastic band passed round the pulley-wheel, and the fine adjustment screw then enabled me to focus comfortably, with my head under the black cloth, by turning the milled screw already . mentioned. As the microscopic objective had to be brought very considerably closer to the object in order to throw a sharp image on the focussing screen than when focussed with the eyepiece in the microscope alone, I noted from experiment the amount of this variation by measurement, and screwed down the coarse adjust- ment of the microscope the necessary amount before connecting it with the camera, and, in this way, left only the final focussing to be done with the apparatus described above. This answered much better and gave me many very excellent results, but I was still not satisfied, as I found that, unless the ova I wanted to take were small or had a comparatively flat upper surface, only a portion of the resulting picture came out sharp, and all the rest was out of focus, owing to the lack of penetration in the microscopic objective. ‘To improve this, I adapted a rapid rectilinear photographic lens of 5in. focus to fit the body tube of the microscope in the place of the usual objective, and, as this rendered the stage useless for holding the object to be photographed, and the distance obtainable between it and the lens was much too short for a lens of so long focus, I took advantage of the circular opening in the stage to work through, and fixed up a movable slide carrier in proper alignment working behind the supporting block on which the microscope stood. This was a great advance, but I was not, of course, able to get so large a magnification as with the microscope objective, as, even with the aid of a special extension which I fixed up between the camera lens front and the microscope body tube, which enabled me to obtain a maximum extension of 5ft. between the focussing screen and the lens, I could only get a magnification of about ten diameters, while the exposure required was nearly double. On the other hand, I had the advantage of an iris diaphragm in the lens, so that I could focus with this open to the fullest extent and admitting the maximum amount of light, and then stop down to f. 16 or f. 22 for the exposure, and in this way get the whole of the largest butterfly ova including even the background. A very powerful illuminant is an absolute necessity for this kind of work. I started with a paraffin lamp, but soon found this useless, not only on account of the amount of heat it gave out, but also owing 10 BRITISH BUTTERFLIES. to the lack of contrast obtainable in the illumination, and particularly to the very great difficulty in focussing accurately when the ground glass image was so faintly hghted. Fortunately I was able to avail myself of an electric light installation, and found a -25 Amp. Nernz lamp admirable in every way, as it gave a 25 to 30 candle-power hight, and this, with a single condenser on one side of the object and a mirror reflector on the other to reduce the blackness of the shadows it made, appeared as nearly ideal as I could imagine. I have not tried incandescent gas myself, but I understand from friends who use it that it gives equally good results. Still 1 should imagine that the heat trouble would obtrude itself, especially if the light were placed near to the object, whereas, with the Nernz lamp, the light can be approached to within 3in. of the ovum being operated on, without any untoward results. The great points to bear in mind are to get a brilliant illumination of the object from one side, and a somewhat less powerful lighting on the other, so as to show up the structure and rotundity of the ovum by contrast. The final critical focussing is best accomplished with the aid of a focussing magnifier held against the eround glass focussing screen, and fixing the attention on some brightly illuminated spot on the surface of one of the ova to be photographed. After focussing and placing the sensitive plate in position, it is only necessary to cover the lens with a piece of black card while drawing out the flap of the dark slide, as any small amount of indirect side light entering the lens is quite negligible, and the card is more conveniently manipulated than a lens cap. The exposure necessary will naturally vary with the brilliancy of the illumination and the colour of the ova, but there is a large amount of latitude permissible with most of the dry plates on the market. J use the most rapid isochromatic plates I can obtain, always backed to minimise halation, and find that, with the 3in. microscopic lens and a magnification of twenty diameters, an exposure of two and a half to four minutes, according to the colour of the ova, is about right. With the din. Rapid Rectilinear, stop f. 16, magnification ten diameters, and similar illumination, I should give from four to seven minutes, but these exposures might be doubled without detriment to the resulting negative by a slight addition of bromide of potassium to the developing solution. For holding ova in position while being photographed I use ordinary glass slips 8in. x lin., such as are used for mounting micro- scopic slides. Ova in sit on bark, portion of leaves, paper, ete., are easily fixed with a small dab of adhesive material, and, where needed, a background of suitably toned paper can be gummed on the glass slip first, and then the support for the ova fixed upon it. When the ova are loose, and particularly if it be desired to retain them uninjured for subsequent hatching, I find a most suitable method of mounting to be as follows :—Cut out a small square of gummed paper, say 8in. each way, and punch out a circular hole in the centre }in. in diameter. An ordinary cork boring drill does this admirably. Then cut another smaller square of paper, tinted to suit the ova in question, and gummed on one side. ‘This should be a little larger than the hole alluded to above, say ;°;in. square. Place the small square, gummed side upwards, in the centre of the glass slip, moisten the gum PHOTOGRAPHING BUTTERFLY EGGS. 11 of the larger square, and press it down upon the smaller, so that the hole is entirely filled up, and all is held firmly to the glass shp. Now moisten a fine hair pencil between the lips and pick up a single ovum on the point of it. Breathe upon the prepared gum surface within the tin. hole, and place the ovum lightly upon it. Continue the process with as many ova as you wish to photograph at once, and they will all be found to adhere quite as firmly as is necessary for the purpose in hand, while they can be easily brushed off afterwards, and will be none the worse for the treatment they have undergone. At least one ovum of each species should be mounted upon its side, and it will then be available afterwards to afford measurements otherwise unobtainable from the resulting photograph. It is a very good plan to line the box, in which living ?s are put for the purpose of obtaining ova, with paper, as then the ova will be laid in most cases on the paper, and are easily accessible, whereas, if they are laid directly on the sides of the box, these must either be cut up or a thin shave taken from them to obtain the ova uninjured. The very worst material I know for photographic purposes on which ova can be laid is cotton wool. Hach individual ovum must be separated from every strand of the wool before it can be mounted with any hope of getting it to he flat, or of placing the entire batch as nearly as possible in one plane (a most important point to remember, or your photograph cannot be in focus all over), while every strand of the cotton which does get mounted with the ova, and there are sure to be some, comes out like a piece of rope, and quite spoils the picture. Ii is only necessary to try to manipulate ova so laid once, and you will thereafter be very careful to ask all your friends, when sending you any, to avoid getting them laid on cotton wool as they would avoid the plague. It is unnecessary to go into any detail as to methods of developing the negative after it has been taken, as these will vary with the make of plate and the particular developer used, and are easily obtained from the manufacturers; but it is helpful to keep a full note with each exposure of at least the name of species, magnification, hghting, make of plate, exposure given, developer used, and colour and appear- ance of the ovum. I write these particulars upon the outside of a paper envelope, into which I afterwards slip the finished negative, before storing it away, and I always pencil the name of the species upon the corner of the plate before development, so that if it gets separated from its proper envelope at any future time it can be identified with certainty. The envelopes alluded to above should each bear a consecutive number, and, if an alphabetical index to these is compiled as they are made, any particular species required in future can be found without loss of time. The exact amount of magnification is important, and once a con- venient size has been fixed on, that, or multiples of it, should be strictly adhered to, as otherwise any comparison will be difficult. An easy method of ascertaining the magnification is to photograph a finely ruled scale in place of the ova, and then compare your negative with the original scale. 12 BRITISH BUTTERFLIES. CHAPTER Y. OBTAINING EGGS OF BUTTERFLIES. The student of the eggs of butterflies, be he biologist, field-naturalist, or photographer, will undoubtedly attempt to collect or otherwise obtain the eges required for study. The best modes of obtaining these in confinement, and the best means of surmounting the difficulties that occur, are, perhaps, rather out of place in a work of this kind, and have already been dealt with fully elsewhere (Practical Hints for the Field Lepidopterist). A few hints, however, as to the best methods of obtaining the eggs of some of the species will, perhaps, not be so out of place, especially as the systematic portion of this work will take some considerable time in publication. The following hints are arranged under the headings of the various months in a manner likely to be followed by the field-naturalist, and in the order requiring attention. January-Aprit.—Hges of Zephyrus querctis on oak, Thecla w-album on elm, 7’. pruni and Ruralis betulae on sloe, are all placed as a rule near leaf-buds on twigs, and hatch towards the end of April. Eggs of Plebetus aegon (argus) pass the winter in this stage; towards the end of February a sharp eye should be kept on them, as they always hatch in the last few days of February or the commencement of March. Females of Polygonia c-album captured in late March and early April will lay freely on hop, currant and nettle, if carefully sleeved on plants that obtain a fair amount of sunshine. Hybernated Hugonia polychloros taken in the spring should be ~ retained for eggs; both sexes should be sleeved together for the purpose, as the earliest caught 9° s after hybernation are rarely fertilised. Sallow is one of the best plants for the purpose of sleeving, and if the captives be fed with syrup, soaked into pieces of bark, they will lay freely. When a 2 Gonepteryx rhamni is observed flying by a hedgeside or on the outskirts of a wood, watch it until it selects a Rhamnus bush for egelaying; collect carefully the leaves and shoots afterwards; you will readily find the spindle-shaped eges on the leaves and petioles. The underside of a leaf, the twig itself, or a terminal bud, is usually chosen, although the upperside of a leaf is not despised. At the end of April and in early May the eggs of Cyaniris argiolus are laid singly on the underside of the calyx of holly-buds, so that, when the flowers open, the sepals fold over the ege, hiding it altogether from sight; the eggstage lasts about ten days. The eggs are also laid on the footstalks of flowers of Rhamnus frangula, and also on the young leaves of ivy. May ann June.—The pale greenish eggs of Nisoniades tages are laid on the leaflets of Lotus corniculatus from the end of May to the middle of June, the eggstage lasting about a fortnight. Females of Augiades sylvanus will lay their eggs in confinement in late June, on cock’s foot grass, if they be enclosed under a leno cover and placed in the sun. Similarly, females of Cyclopides palaemon may be enclosed over a plant of Brachypodiwm sylvaticum. The circular, flattened, greenish-drab eggs of Polyommatus astrarche OBTAINING EGGS OF BUTTERFLIES. 13 are laid in June, in little groups of two, three, or more, on the under- side of the leaves of Helianthemum vulgare. In June, by following up a female Polyommatus bellargus when on egglaying intent, one can obtain eggs quite freely by picking the leaves one after the other as she quits the plants on which she has been engaged. The egg of Lycaena arion is deposited in June among the flowers of Thymus serpyllum, being circular in outline, and covered all over, except a central depressed spot on top, with fine raised irregular reticulation, which, in profile, stands out strongly. Pick the flower-heads of Anthyllis vulneraria in June for the eggs of Cupido minima; although they are placed low down on the calyces of the Anthyllis flowers, and thus hidden from casual observation, they may be easily detected on careful search. Females enclosed over a plant of Anthyllis vulneraria, and allowed plenty of sun, will lay eggs freely among the flowers. The eggs of Nemeobiuslucina are readily found on the underside of cow- slip leaves in late May and June, not more than four or five on a leaf ; also to be found similarly on primrose leaves. They may also be readily obtained by enclosing caught ¢s on potted plants of cowslip or primrose. The eggs of Melitaea aurinia can be obtained freely by enclosing caught females in a leno sleeve over a plant of Scabiosa succisa, the egos being laid in heaps on the surface of the leaves. In nature, the egg-batches may be found by careful searching and turning over the scabious leaves in their haunts. Captured @s of Brenthis euphrosyne, taken in late May or early June, lay their eggs freely on theleaves of Viola canina. So also do those of B. selene, choosing indiscriminately the upper- and underside of leaves and the stems. The imagines of Melampias epiphron will lay their eggs in confine- ment, if placed in a suitable receptacle with a supply of grasses on which the larve will feed, e.g., Nardus stricta, Aira flexuosa, etc. In late June (and during July) the beautiful eggs of Limenitis sibylla, covered with deeply-set hexagonal basins, and sharp prominent spiny points, giving rise to fine gossamer-like hairs, are laid on the edge of the underside of a honeysuckle leaf. Tigges of Colias edusa may be obtained in June by placing a under a bell-glass with a sod of white clover; they are laid on the upper surface of the leaves; the eggs in a batch often hatch irregularly, even when the whole is deposited within a few hours. The ?s should be supphed with a little honey and water for food, and will lay their eggs pretty freely so long as the weather is bright and sunny; during dull weather the butterflies will not lay. Immigrant females of Pontia daplidice lay their eggs occasionally in June, on Reseda luteola. In May, the flowering stems of Cardamine pratensis and Alliaria officinalis should be collected for the orange-coloured (yellow when first laid) eggs of Huchloé cardamines, which are usually laid (one on each flower-head) on the pedicel of a flower nearly over. The long spindle-shaped eggs of Leptidia sinapis are to be found readily in May on Vicia cracca and Lathyrus tuberosus. The globular, greenish-yellow or greenish-white (when newly-laid) 14 BRITISH BUTTERFLIES. eges of Papilio machaon are to be found, in early June and on through the month (often, indeed, until August), laid usually on Peucedanum palustre, in its local haunts in Cambridge and Norfolk. JULY-SEPTEMBER.—The eggs of many species of butterflies may be obtained in July, e¢.g., that of Limenitis sibylla (a most beautiful micro- scopic object) on honeysuckle; Apatura iris on sallow, etc. (We have seen a dozen of the latter collected in a single morning by watching a ? , in a place where larve could never be beaten owing to the density of the vegetation). The eggs of Urbicola comma, laid in August, on grass, do not hatch until the following March. Similarly, those of Adopaea lineola and ? Thymelicus acteon, laid in July. As most of the ‘“‘skippers”’ hybernate as larvee, care must be taken to look after the eggs of these species, and not throw them away with the idea that they are infertile. The eggs of Plebeius aegon are laid in July on Ornithopus perpusillus, etc., but do not hatch until the early part of the following March. The circular, flattened, greenish-drab eggs of Polyommatus astrarche are laid in August and September, in little groups of two, three, or more, on the underside of the leaves of Helianthemum vulgare. Kegs of Cyaniris argiolus are laid in August, sometimes beneath the flower-heads of the umbels of ivy; the young larve then feed on tender ivy-leaves and flowers. They are at other times laid on the flower-stalks of holly, then the young larve burrow in the unexpanded buds. Worn ¢sof Thecla w-album will oviposit freely if sleeved out in the sun on elm, in the early part of July. In nature, they are to be found above or directly below an aborted leaf-bud, and harmonise so exactly with the colour of the bark of the elm-twig on which they are placed that they are only to be detected with the utmost difficulty. The yellowish milk-white eggs of Zephyrus querciis, covered with a rough raised reticulation, are laid upon oak-twigs, where they may be found during the winter months. In spite of its colour, the egg is not at all easy to see, looking like a small, inconspicuous fungoid growth ; after the maturation of the embryo it grows somewhat darker. The females of Coenonympha tiphon will lay their eggs in confine- ment, if placed in a suitable receptacle, and exposed to the sun, with a supply of their foodplant, the beaked rush (Rhyncospora alba); this should be potted, and the young larve will feed thereon until their hybernating stage with little trouble. Females of Hrebia aethiops will lay their eggs quite freely in August, if supplied with grasses in a suitable receptacle, and placed in the light and sun; the eges are glued to the culms of Aira praecox, A. caespitosa, etc., and are large and conspicuous. ‘The young larve appear in about three weeks. The females of H'pinephele tithonus will lay their eggs fairly freely in confinement on Poa annua, Dactylis glomerata, and other common erasses; they hatch in about three weeks, and the young larve hyber- nate when exceedingly small. The females of Melanargia galathea give their eggs perhaps more freely than any other butterfly; they are unattached, hatch in August, the young larvee feeding well on common grasses. The eges of Aryynnis adippe are laid in July and August on the leaves of Viola canina, generally on the underside or on the stems. BUTTERFLY LARVZ AND THEIR MOULTINGS. 15 They change colour very rapidly as the embryos mature, but the larve do not appear till late February or early March the following year. The eggs of Dryas paphia are laid in July and August; the egg- stage, however, only lasts about a fortnight, although the larve feed very little (or not at all) on Viola canina before hybernation. Autumnal females of Colias edusa, enclosed on a growing plant of Trifolium repens, Lotus corniculatus, etc., placed in the sun, and supplied with a little honey and water for food, will lay their eggs pretty freely so long as the weather is bright and sunny. Captured females of Colias hyale will lay their eggs on Trifolium repens, Medicago lupulina, M. sativa, etc.,in August. They hatch in about a fortnight, and the young larve feed up slowly to hybernation. 99 These are only a few “hints” extracted from our work Practical Hints for the Field Lepidopterist, and are simply inserted as illustrations of the details to which the attention of the seeker for eggs of lepidoptera must be directed. CHAPTER VI. BUTTERFLY LARVHZ AND THEIR MOULTINGS. The newly-hatched larve of many families of butterflies are very similar to each other, much more so in many cases than are the newly-hatched and adult larve of the same species. The term “embryonic”’ has been applied to newly-hatched larve in their first instar or plumage, 7.e., before their first moult, and the term is a happy one, because, structurally, the larva until this moult retains all the characters that the embryonic larva possesses just before it leaves the egs. This stage rarely lasts more than a few days (although Dryas paphia and Argynnis aglaia are both reputed to have larve that leave the eggs in August, and, without feeding, remain in this state till the following March). The marked peculiarity of many newly-hatched lepidopterous larvee is the similarity in the position and arrangement of certain little chitinous buttons or knobs, each bearing a conspicuous hair or seta; these buttons, with their sete, are known as the primary tubercles. This similarity the young butterfly larva shares, with scarcely any modification, with the larve of most other lepidopterous superfamilies, and, as we know that the embryonic stages often recapitulate the past history of the development of the species, we speak of this general or common form of butterfly larve as a generalised type, and consider it as exhibiting, more nearly than the adult larve, a primitive or ancestral form of the butterfly caterpillar. With the first (or, at latest, the second) moult a very considerable change takes place, and the larva becomes more specialised. The larve of our butterflies live more or less exposed on their food- plants; their enemies are numerous, and they are sought eagerly by various animals as food. Their colours, hairs, etc., are so modified as to make them difficult of detection on their foodplant in their normal position of rest, or by making them unpalatable if they be detected. Hence the larva of each species is so far modified or specialised in the direction of its colour and markings, or in its armature—spines, hairs, 16 BRITISH BUTTERFLIES. etc.—or in both, as will best protect it from the enemies which would otherwise prey on it. Similar, therefore, as the newly-hatched larve may be, they present, usually, at their first moult, a marked change in their appearance, and this frequently becomes more pronounced at each successive moult until the larva is fullzrown. The difference between the newly-hatched larve of the Vanessids, Argynnids, etc., and their adult forms is very great, e.g., compare the newly-hatched and adult larve of Dryas paphia, Aglais urticae, etc. If we study the habits of these larvee we shall find that these changes culminate in producing just that form which is best protected by the particular environment which surrounds it. The preservation of the more suitable, and the weeding out of the less suitable, individuals by natural causes, is known as ‘‘natural selection.”” Butterfly cater- pillars have, therefore, been brought to a high state of fitness to their surroundings by natural selection. Their independent mode of life makes the larval specialisations run in entirely different directions from those which are most effective in the preservation of the pupa, or imago, where the conditions of its environment are so entirely different. i All newly-emerged larve, however, do not conform to the generalised type just noted, but hatch from the eggs already in a _ highly specialised condition, e.g., the larva of Papilio machaon is well provided with spinous processes when it leaves the egg, and others have undergone even more development before hatching. We assume that, as these larve hatch in a more than usually specialised condition, they go through a generalised stage of development in the egg before reaching this more specialised one in which they hatch, and that these earlier stages have at some distant time taken place outside the egg, and that the necessities of a changed environment have forced these later stages into the egg, so that the larva is more specialised and more able to respond to its present environment when hatching takes place. We have, however, no real evidence that this is the case. The larva undergoes a certain number of moults or changes of skin before it becomes adult. The period between one moult and another is called a stage or stadium. The appearance of the larva at any particular moult is known as its instar. Thus a larva that moults four times has five stadia, and five different appearances of its plumage or instars corresponding with the stadia. It may be noted that the head, being chitinous, is of fixed size throughout a given stadium, and that this is of great importance in determining the stage in which a larva is, apart from the size of the body. It is also to be noted that, at a larval moult, not only does the larva cast off its old skin, but the linings of the mouth, gullet, and even of the large air-passages are shed as well, and, if the cast-off skin be examined, you may observe the latter as fine thread-like processes curled up, and starting from the spiracles down the sides of the caterpillar’s body. Previous to moulting, the larva spins a silken pad on which it rests for two or three days, inserts into this the hooks of its prolegs, and here it remains motionless whilst the new skin is being matured beneath the old one. During this time a surface fluid collects between the two skins, the old skin splits, and the larva in its new skin frees itself from the old one, which remains attached to the pad by means of the proleg hooks, which cling tightly to it. BUTTERFLY LARVE AND THEIR MOULTINGS. 7 An excellent account of the exuviation of the larval skin in Dryas paphia is given by Buckler, who writes: ‘‘I observed the larva in _ preparation for its last moult, fixed belly upwards to a leaf, on May 20th. It remained quite still until noon of the 25th, when I noticed it moving its anterior legs a little free from the leaf, a circumstance which claimed my whole attention; it was but a slight movement, and was repeated at intervals of about half-an-hour until between 2 and 3 o'clock in the afternoon, when it began to stretch its 1st segments downwards from the leaf, making the forepart of the back concave, and then presently gently reversing the movement. It continued thus at short intervals to increase the stretching curve of the body so much that, by 10 minutes past 3, its hold on the leaf was retained only by the fourth pair of ventral prolegs and the anal pair, when, suddenly, the skin snapped asunder close to the head, with quite a shock to the larva, which instantly returned its ventral prolegs to the leaf, whilst the elastic skin, relieved of the tension, was itself, from the impetus of the rupture, gliding backwards. The anterior legs were held back until divested and then returned forwards to their natural position one after the other, but kept just free from the leaf, each pair being elevated in unison for a moment, and let fall as though to test their complete freedom; otherwise the larva remained passive, the skin only con- tinuing to move backwards, and, whilst passing the ventral prolegs, each foot was lifted up in turn out of it and then replaced on the same spot of the leaf, and, when the old skin had shrivelled up at the end of the body, the larva, with all the ventral prolegs, took two steps forward and drew forth the anal pair free. At the first breaking of the skin the head became exposed, with the old headpiece adhering to the parts around the mouth, but now, at last, the larva gave its head a sudden twist or two, and the old piece fell off. From the rupture of the skin to this final riddance the operation occupied nearly ten minutes; the spines were all uncovered in a remarkably small, wet and flaccid condition, the front pair even smaller than the others, but now this pair began gradually to grow and in fifteen minutes were far longer than ever, and in another half-hour all the other spines had grown considerably both in length and rigidity. After this the larva remained still for 24 hours longer.”’ With each moult there is usually some change in the appearance, markings, or structure of the larva. Thus, after the first moult, the larva of Apatura iris develops its long horns, the primary tubercles of the Vanessids appear to be replaced by long spines, etc., whilst, on the other hand, in some of the Pierids and Satyrids, they get smaller, and become almost obsolete. Larvee in which the gradual obsolescence in these tubercles may be well observed are Huchloé cardamines and Pararge meyaera, although almost any members of these families will do. The larva of every species of butterfly will, however, show some modifica- tion or other, and all are exceedingly interesting. 18 BRITISH BUTTERFLIES. CHAPTER VII. EXTERNAL STRUCTURE OF THE BUTTERFLY LARVA. The butterfly larva is composed of a head, thorax and abdomen. The last two form a cylindrical tube slightly drawn in by a series of constrictions following one after the other, and dividing it into segments, separated by the drawn-in parts or incisions. The head of the larva appears to consist of four segments, in most cases, closely welded together, but in some instances traces of division are here and there noticeable. The newly-hatched larva of Pararge megaera also shows a somewhat peculiar development, the last head-segment bearing four typical trapezoidal tubercles, arranged as a trapezoid (as on the body-segments), each carrying the usual hairs. The marks on the other head-segments seem to have the same significance, and it affords strong suggestion that the head-segments were originally ordinary tubercle- and hair-bearing segments like those of the body. It is also interesting to note that the newly-hatched larve of Limenittis sibylla and Eugonia polychloros have fleshy spikes on the head, which are probably of the same origin and significance as the spines of the thoracic and abdominal segments. The three segments following the head are the thoracic segments, and the ten following these comprise the abdomen. The Ist of the thoracic segments is known as the prothorax, the 2nd as the meso- thorax, and the 3rd the metathorax. Of the ten abdominal segments the hindmost is known as the anal segment. There is often considerable difference between the armature or clothing of the thoracic and abdominal segments both as to the structure and arrangement. The prothorax usually differs more from the meso- and metathorax than do these from the abdominal segments; the 8th, 9th and 10th abdominal seoments are also generally considerably modified. The number of subsegments, into which the segments are subdivided transversely, ordinarily differs in the thoracic and abdominal segments, the prothorax being usually greatly specialised and differentiated in this as well as in other details. The peculiar structure of the neck-like prothorax of the Urbicolids is very remarkable and exactly opposite to that of the swollen prothorax of the Ruralids or Lycenids, into which the head is freely retractile. There is also a remarkable swollen vesicle, which is placed (usually in a transverse slit) below the Ist thoracic segment, just in advance of the first pair of legs. It is known as the chin-gland. This structure, butterfly larve have in common with those of the Notodontides, Noctuides, and a few closely allied superfamilies. Again, the spines or _hair-bristles arising from the skin are usually arranged in _ longitudinal rows, which have a regular position on each abdominal seg- ment; yet, whatever the arrangement on the abdomen, it will usually be found that this either stops altogether, or changes in direction, as soon as the thoracic segments are reached. The thoracic segments bear the true legs, and the meso- and metathorax, under which the structures that will form the future wings begin to develop in the earliest larval life, have no spiracles, although the prothorax carries one on either side, The abdominal segments that are probably EXTERNAL STRUCTURE OF THE BUTTERFLY LARVA. 19 the most modified are the 8th and 10th; in the Satyrids the latter is modified to form a pair of pointed projections extending out behind the larva. The 8rd, 4th, 5th, 6th and 10th abdominal segments each bear a pair of prolegs, the last pair often known as the anal claspers. The spiracles are tiny holes for the admission of air into the tracheze or breathing-tubes ; they are placed in pairs (one on each side) of the prothorax and the first eight abdominal segments. Hach spiracle leads, by means of a little tube, into a larger longitudinal one so that all the trachez connected with the spiracles are brought into connec- tion ; injury to one spiracle, therefore, throws the work on others and does not kill the larva. It will be observed that the spiracles on the 1st thoracic and 8th abdominal segments are invariably larger than the others, which are usually equal among themselves. The reason for this is obvious, for each has to supply air to a much larger area, the head and thoracic segments being dependent on the prothoracic, and the 8th, 9th and 10th abdominal segments on the 8th abdominal, spiracle. -It is to be observed also that these have usually a different position from the others, being generally placed much higher than the other spiracles. The caterpillar has, as we have noted, three pairs of true legs, one pair situated on each of the thoracic segments. These are usually small, jointed, horny, and provided with terminal hooks, but often so ill- developed as to be comparatively useless for walking purposes. For this purpose, and to support the long cylindrical abdomen, the skin on the underside of the 3rd, 4th, 5th, 6th, and 10th (anal) segments of the abdomen is prolonged downwards to form the prolegs and anal claspers (or false legs), and, to make them more effective for walking purposes, they are composed of joints which are partly retractile one into the other ; the end joint of these prolegs is provided with minute hooks, the arrangement of which is most important as giving clues to the line of evolution of the various groups of butterflies. At the same time they enable the larva to cling tenaciously to its foodplant. In the most highly developed butterfly larvee these hooks exist only along the inner margin of what is now a flange, but what was once a circular pad, and the larve of the skippers still have an almost complete circle of hooks on the prolegs, hence the conclusion that the butterflies have been derived from lepidopterous ancestors of a low or generalised type such as the Hepialids, etc., which possess somewhat similar prolegs, and not from the higher or more specialised moths which have the same type as the other butterflies, 7.e., excluding the skippers. We have already referred to the primary tubercles and the hairs carried by them in the newly-hatched larva. It may be well now to note them a little more particularly. The simplest form consists of a little chitinous knob bearing a single hair or seta. Sometimes these are modified so that a slightly raised base bears several hairs, in addition to the primary seta, when we get a tubercular wart; again they may be modified into spines, fascicles of hair, etc. Examination of the dorsum (or back) of a larva will show, at least on the abdominal segments 1-8, two rows of chitinous-based hairs, or spines, running down the length of the body, two on either side of each segment, those in front, rather nearer the middle line of the back than those behind ; the front ones are known as tubercles i, or the anterior trapezoidals, the hinder ones 20 BRITISH BUTTERFLIES. as tubercles ii, or the posterior trapezoidals. Another row is found along the side just above each spiracle, these are known as tubercles ill, or supraspiracular tubercles, another row is placed just behind each spiracle, called tubercles iv, or postspiracular (in many superfamilies of the lepidoptera tubercle iv is subspiracular like v), whilst a row placed directly under the spiracles are called tubercles v, or subspiracular. At the tops of the outside of the prolegs is another series known as tubercles vii, or marginal tubercles, whilst between v and vii is a row of secondary, rather than primary, tubercles, known as tubercles vi. These should be thoroughly worked out by every lepidopterist who wishes to study the structure of butterfly larve, and particularly to describe them. We have already stated that the position of those on the thoracic segments is often much modified, and their homologies are not always easily to be determined. The differences in character and position between the corresponding tubercles on the different seoments are of the greatest possible importance, particularly is this so in the case of a comparison between the positions of those on the thoracic and abdominal segments. _ Besides the hairs or sete carried by the tubercles, which have fairly fixed positions in all butterfly larvee, the skin of most butterfly larve has, scattered more or less regularly over the body, little elevations, resembling somewhat a fine pile or covering of minute hairs. This pile is a very common feature in butterfly larve, is supported by very minute papille, and is generally distributed with considerable regularity, usually in a transverse, though sometimes in a longitudinal, direction. It is, however, occasionally scattered irregularly over the body, and when it is arranged transversely, it:is usually somewhat closely related to the subsegmental divisions into which the segments are subdivided. It is not at all confined to butterfly larve, but is found very generally in those of many other superfamilies. As an illustration of the vagaries connected with its appearance one may note that in the larva of Aglais urticae it is strong in the early stages, whilst in that of Zephyrus querets it is strong when, and not until, the larva is fullfed. Scudder thinks that ‘‘the clothing prevents the too rapid evaporation of the heat from the surface of the body, for, although larve are cold-blooded animals, they, nevertheless, have an internal heat above that of the surrounding atmosphere, which originates from the activities of the organs and the respiratory functions, and which they would lose more rapidly but for this investing pile.”’ Many larvee are provided with what may fairly be termed glandular hairs. They are more especially abundant in young butterfly larvee and occur in all the larval stages among the Satyridsand Pierids. In the Pierids, they form an open basin, fringed with cilia, supported on an exceedingly slender hollow pedicel. In the basin a drop of transparent fluid may be secreted when the hairs look as if tipped with dew. The purpose of the secretion is possibly protective, as the volume of the fluid is visibly increased when the larvee are excited. Scudder writes: ‘‘ They are generally arranged in longitudinal rows, and their use is wholly unknown, but they probably have a protective function, for this fluid is odoriferous, the secretion increasing when the larve are disturbed.’’ He describes them as ‘* papilla-mounted bristles, each furnished with a trumpet-mouthed tip,’ and adds that they “are the ducts leading from glands at their bases, secreting a transparent fluid, which, after secretion, is borne in a EXTERNAL STRUCTURE OF THE BUTTERFLY LARVA. 21 little globule in the mouth of the trumpet, and sometimes kept in its place by a few microscopic bristles which surround its rim.” These glandular setze are really the hairs of the primary tubercles. Scudder, speaking of the change occurring in butterfly larve (Butts. of New England, 11., p. 805), at the various skin-moultings, says: The mature Satyrine larve have a rough skin the result of a multitude of minute tubercles, each bearing a simple hair scarcely visible to the naked eye; in the young larve of the Satyrids, the skin, instead of being supplied with an almost innumerable number of microscopic hairs, is furnished, in some instances, with an exceedingly scanty -number of little club-shaped bristles, proportionally many times longer than the hairs of the adult . . . . arranged in definite longitudinal series; in others, with compressed, ribbon-like hairs as long as the body, serrated on one edge and bent in the middle; on the abdominal segments, these hairs point backward, and on the thoracic, forward. In the Nymphalids, the segments of the young larve are equal in size, and have regular series of stellate warts; in the mature larva, the body is grotesquely hunched, while the warts have changed to very variable tubercles, etc. In Anosia archippus the fullgrown larva is naked, but adorned with a pair of long thread-like fleshy flexible tentacles at either extremity of the body; in the young larva, these tentacles or filaments are absent, but their future position is marked by little conical black points, while the body is covered with minute black bristles, arising from still more minute warts, and arranged six on the back of each segment (?1, 11, 111), and three on either side of the body (?iv, v and vii). In the Vanessids, the larval spines are com- pound in the adult, and arranged in certain definite rows; in their earliest life, these same larve are furnished with long tapering hairs, also arranged in definite series, but not occupying the same positions as the spines of the mature larva. In Agraulis vanillae and Apostrophia charithonia, two Heliconians, the appearance of the larva after the 1st moult is entirely different from that preceding it. In the 1st stage, the head is unarmed, and the body supports longitudinal rows of very large papille, each bearing a long slender naked hair with a delicate ovate apical club. After the 1st moult, the head is armed above with a pair of stout spines nearly as long as itself, bristling with distant thorns; and, in the place of the primary hairs, are long tapering spines as high as the body, with a very slight basal enlargement, and furnished along their whole length with minute papille supporting little needles, the position of these spines is quite different from that of the papille of the 1st stage, and, as if to mark this more distinctly, there are but three longitudinal series above the prolegs; these differences become intensified in every subsequent ecdysis. The adult Ruralid (liyczenid) larvee appear to be quite smooth, although covered with microscopic hairs, whilst the newly-hatched larve of this group are provided with long spiculate primary hairs that sweep backwards behind their bodies. In the Urbicolids, the primary sete of the newly- | hatched larvee are always shaped lke little clubbed mushrooms. The adult Papilionid larva is always nearly naked; a few scattered hairs may be found with a lens, with a few minute tubercles or smooth and shining lenticles; in some, the front part of the body is swollen, and furnished with striking eye-spots; at birth, however, the body is always cylindrical and supplied with several prominent series of bristle- 29, BRITISH BUTTERFLIES. bearing tubercles, one tubercle to a segment in each row, and one row often more conspicuous than the others; sometimes the entire body bristles with these appendages. As hasalready been noticed, there is, in certain of our British butterfly larvee—Urbicolid, Pierid, Satyrid, etc.—a tendency to the obsolescence of the primary sete. In other cases, however, the bases of the tubercles are developed into long fleshy processes, carrying aborted sete, e.g., in Vanessid, Argynnid, Meliteid larve, etc. That these complex processes, often bearing long sharp spines, are modifications of the tubercular structure, and are dermal appendages, appears certain if we examine the newly-cast skin of a Vanessid or Argynnid larva. The structure of the spines of the larva of Dryas paphia, the movable prothoracic horns of Apatura iris, with moving tubercular bosses, and the anterior flexible filaments constantly in motion backwards and forwards (especially when eating or alarmed), are all important from the point of view of the development of special external structures for protective purposes. The larvee of butterflies have then undergone special development along various lines for protective purposes. The resemblance of some, especially grass-feeding larvee, to their foodplants, makes them readily overlooked; those of the Melitzeids closely resemble the long bloom-heads of plantain and allied plants to which some species are attached, whilst those of the Apaturids and others are most difficult to detect owing to their resemblance to the leaves, etc., of their foodplant. Others, again, are protected by the sharp, prickly spines into which the tubercles are modified, e.g., the Vanessids, whilst others again have bright warning colours, or are protected by nutant spines, etc., evaginable osmateria, e.g., the larve of the Papilionids. Butterfly larve, therefore, show considerable variety in their means of defence and consequent ability to escape their vertebrate enemies. They are, however, subjected to the serious attention of a vast army of smaller foes, especially diptera and hymenoptera, which lay their eggs in them, the caterpillars from these eggs devouring the internal organs of the larve, and, after maturing thereon and killing their host, pupating either in the body or directly after leaving it. The destructive powers of some of the smaller species are very ereat. One minute species lays its eggs in tha newly-hatched larvee of Melitaea aurinia, in June, and, according to Wolfe, after the larva stops feeding preparatory to hybernation, the parasite forms its cocoon inside the web spun by the larva, in which to hybernate, and the latter of course dies. ‘The imagines of these ichneumons emerge in spring, sting fresh M.aurinia caterpillars, and, so rapidly are their own metamorphoses completed, that even a third brood of the parasites will attack the same katch of larve before the latter are full-grown. The destruction caused by such parasites can readily be understood. Many peculiar structures are to be observed in butterfly larve. We have already referred to the chin-glands—eversible bladder-like olands hidden in a slit on the ventral surface of the prothorax, which are everted when the larva is disturbed, but which appear to have no power of emitting any fluid, although it is possible that some scent may render them of service as a means of protection to the larve possessing them, and which appear to be general among Nymphalid larve. But the most striking of the eversible glands in the butterfly larvee are the well-known osmateria of the larve of Papilionids, Parnassiids and EXTERNAL STRUCTURE OF THE BUTTERFLY LARVA. 23 Thaids, which are placed in a transverse slit on the upper part of the prothorax. When irritated, the larve thrust therefrom a large orange- yellow, Y-shaped, fleshy, tubular process (the osmeterium), from which is diffused a very appreciable odour, varying in its nature according to the species, usually more or less objectionable, and, in some cases, exceedingly so; this is frequently accompanied by a drop of fluid which Packard says is acid, and turns litmus paper red. The mechanism has been described and figured by Klemensiewicz. Packard notes that, when at rest or retracted, the osmaterium lies in the upper part of the body in the three thoracic segments, and is crossed obliquely by several muscular bundles attached to the walls of the body, and that by the action of these muscles, the evagination of the osmaterium is strongly promoted. After eversion, the tubes are slowly retracted by two slender muscles inserted at the end of each fork or tube, and arising from the sides of the metathorax, crossing each other in the median line. Secretion takes place in an oval mass of glandular cells at the base of the forks; in the glandular mass is a furrow-like depression about which the secretory calls are grouped. The secretion collects in very fine drops on the side of each furrow opposite the glandular cells. Its particular structure in Papilio machaon and the details of secretion and method of movement have already been dealt with (Nat. Hist. Brit. Lep., 1., p. 95). We need only add here that the osmateria are probably protruded by the muscular contractions of the walls of the body, forcing the contained fluid into the tube, and thus pressing out the reversed osmaterium. On the dorsum of the 7th abdominal segment of most (probably all) Ruralid (Lyczenid) larve is a narrow transverse slit. In this slit, in many species, is a very minute eversible sac, whose function appears to be directly opposite to that of the osmateria already described, for the sac exudes a sweet fluid very attractive to ants, which may be diffused more widely by the delicate spinulose bristles crossing the summit. It is assumed that, in return for a supply of the sweet fluid, the larvee are protected by the ants from predaceous enemies. Scudder says that all Lycenid larve have the slit, though all do not possess the gland; in those that do possess it, it is found to be a vesicle of somewhat tubular shape that can be thrust through the transverse slit, which, when closed, looks exactly hke a transverse line running across the dorsum of the segment. The connection between these larve and ants has already been noticed (Nat. Hist. Brit. Lep.,1., pp. 97-98). In addition, Edwards states that, in several Lycznid species, there is, besides the gland on the dorsum of the 7th abdominal segment, a pair of minute dorsal evaginable tubercles. The larva of Pirochala isocrates, the well-known Pomegranate butterfly, is said by Pargiter to have two white spots near the anal end of the body, in each of which is a small hornlike process, which the larva continually protrudes and retracts. Nicéville gives (Butterflies of India, vol. 111) an excellent account of two tubercles with protruding flagella, found one on each side of the 8th abdominal segment of Curetis thetis. These are de- scribed as two diverging cylindrical rigid pillars, arising from the sub- dorsal region, and of a pale green colour. When the insect is touched or alarmed, a deep maroon tentacle, as long as the right pillar, bearing on its end long parti-coloured hairs (the basal third black, and the upper two-thirds white),is everted. The maroon tentacle, with its long 94 BRITISH BUTTERFLIES. hairs spread out like a circular fan or rosette, is whirled round with ereat rapidity ina plane parallel to its body, its use being, almost certainly, to frighten away its enemies. Similar eversible glands are described by Hagen as occurring in the larve of Plebeius argqus and Polyommatus corydon. He writes: ‘“ You find on the penultimate seg- ment, outside and behind the stigmata, two large white spots, each of which evaginates a white membranous tube, just like the finger of a glove, the top of which is not entirely drawn out.” Exactly what measure of protection is afforded by these flagella is not known, nor the manner in which they afford it. Many authors, e.g., Scudder and Dimmock, incline to the opinion that they are of the nature of osma- teria and diffuse odours, but so far the odours do not appear to have been detected. The glands are by no means present in all Lycenid larvee, closely allied species differing in this respect. In many lepidopterous larve (Nat. Hist. Brit. Lep., 1., p. 40) there are present two bristles, each standing out backwards from a papilla, placed directly under the anal flap, sometimes looking as if they pro- jected from the base of the anal prolegs, and used by the larve to throw the pellets of frass to some distance from where they are feeding. These are called ‘‘paranal forks’’ or ‘‘paranal tubercles.”” In butterfly larvee, their place seems to be taken by the “‘ anal comb,” which Chapman considers may possibly be homologous with the paranal forks. It is present in Urbicolid larve. Scudder also figures the structure in Colias (Hurymus) philodice. It will be dealt with at length in some of our detailed larval descriptions. One of the most remarkable external features of the Urbicolid larvze is the development of pecuhar glandular structures on the venter of the 7th and 8th abdominal segments in their last instars. We have described them somewhat at length in our descriptions of the Urbicolid larvee (in the systematic portion of this volume), and there is no need to redescribe them here. They appear, in the case of the Palearctic species, to be active only in the final larval stage, when the puparium is thickly sprinkled with the asbestos-looking material secreted by these glands, apparently for the purpose of keeping it watertight, but, in some of the exotic Urbicolids, it would appear that the larval shelter itself may be covered with the secretion for the same purpose, e.g., Nicéville notes (Butterflies of Sumatra, p. 588) that the larva of Erinota thraw is covered with a white waxy powder, and that it lives in a shelter made of a portion of one of the enormous leaves of Musa. He adds that the pupa is also covered with the same white powder, which is of the greatest service to the animal, as, in consequence of the heavy showers of rain in the tropics, much water often collects in the rolled-up leaf, and the pupa, if not so protected, would soon be drowned and rot ; as it is, the powder keeps the pupa dry until the water has drained away or dried up. The downy larva of the allied Gangara thyrsis is similarly covered with a white waxy powder. Little is known of the remarkable structures, detailed at length in our accounts of the larve of the various Urbicolid species, and there called lenticles. Scudder describes them as crateriform chitinous annuli, which are ranged in longitudinal rows along the abdominal (and sometimes the thoracic) segments. They are found only in certain groups, appear to be an universal characteristic of the earliest stage of the Lycwnids, having the appearance of spiracles, only they INTERNAL STRUCTURE OF THE BUTTERFLY LARVA. 25 are usually quite circular, whilst spiracles are ordinarily oval, and they present no opening in the centre, but only a simple pit of more delicate structure than the chitinous annulus itself. They are also found in some Urbicolid larve in their first stage and sometimes also throughout life, but for the tenuous structure of the pit in the centre, they would have all the appearance of suppressed spines, and, indeed, the central pit seems sometimes to be wanting, and we have simply a shining lenticle, similar to those which are common in the Papilionids, but whether they should be looked upon as structures on their way to some use, or as effete structures, degenerated spines, so to say, we have no evidence at present to show, and an explanation of their purpose has still to be sought. CHAPTER VIII. INTERNAL STRUCTURE OF THE BUTTERFLY LARVA. The internal anatomy of the butterfly larva is exceedingly complex. The external features of the butterfly larva are comparatively easily de- scribed, and the position of the structures located owing to the segmented form of the body, and the fact that special organs and appendages are re- stricted to certain segments. The dealing with the internal organs and structures, is, however, a much more difficult matter, as most of them are not restricted to certain segments, but run longitudinally through the body, sometimes extending from the thorax, forward into the head, or backward into the abdomen. It is, therefore, necessary to consider each separately, both as regards position and function. The movements of the body are of the first importance, and the larve have, in various species, undergone great modifications to enable them to vary their movements according to their needs. Movements are dependent on the muscular system, and the changes that take place in the appearance and configuration of the larva, when movement occurs, are due to the muscles. The nutrition of the various parts is maintained by food, and, to comprehend this, the digestive or alimentary system must be studied. The absorption of the digested food into the blood and its carriage to the different parts of the body, necessitate a circulatory system, whilst the oxygenation of the blood leads up to a consideration of the respiratory system. This latter is so intimately connected with the excretion of waste, that one is forced to consider the excretory system, whilst the organs, by which the whole of these various systems is governed, comprise what is known as the nervous system, and this has to be considered, both in its relation to volition and sensation. These various systems comprise, then, the different organs (and their functions) by means of which the life of an insect is carried on, and their external results, as exemplified by their movements, etc., are the outward signs of their vitality. The reproductive system, which is not, however, matured, nor very largely developed, in the larval stage, must take the highest place in relation to the continued life of the species. Closely related, too, with the digestive, is the cellular system by means of which the caterpillar is able to store up large quantities of surplus material for use in the later stages of its metamorphoses. We 26 BRITISH BUTTERFLIES. have already dealt with the internal anatomy of a lepidopterous larva at length (Nat. Hist. Brit. Lep., pp. 54 et seq.), and, since that of butterflies does not differ, in any of its essential features, from that of other lepidoptera, we shall not do more here than repeat a few of the salient facts, which may be considered under the following heads :— (1) The muscular system: The voluntary muscular system of the larva, is that by means of which it is enabled to move about in order to obtain its food. The muscular fibres are usually arranged as flat ribbons or conical bundles, the latter making up almost the whole structure of the head, and are attached to the headwalls, stretching to the mandibles, labium, labrum, etc. A series of contiguous muscular cords, or bands of longitudinal muscular fibres, run from one end of the body to the other on each side, just under the skin, between the spiracular line and the venter of the body ; other longitudinal muscular bands run above the spiracles; a transverse muscular belt encircles the body at the front of each segment, whilst oblique transverse muscular bands run from the front of each segment, and are attached to the medioventral line further back in the segment. Besides these, complicated muscular systems bring about the movements of the legs and prolegs. The involuntary muscular system is principally connected with the digestive and the circulatory organs. The esophagus is provided with fine longitudinal, and with less well-developed transverse encircling, bands of muscular fibre. The inner coat of the stomach is enclosed in delicate strips of muscular fibre crossing each other diagonally, whilst longitudinal muscles run throughout its length, and the well-developed encircling muscles are similar to those of the cesophagus. The arrangement of the muscular tissue in the intestines, is very similar to that of the rest of the alimentary canal, only the longitudinal bands are often thick and glistening, whilst near where the small intestine joins the stomach, the walls are plentifully supplied with short longitudinal muscles; the diagonal bands and encircling muscles found in the stomach also have their representatives here. The alimentary canal is held in its place by a series of muscular bands attached to the body wall, one set passing round that portion of the intestine where it is connected with the stomach, another set being attached to, and supporting, the posterior end of the small intestine, these muscles stretching horizontally from the middle of one side of the 8th abdominal segment to the opposite side. (2) The alimentary system: The mouth opens into a short gullet, and this in turn expands into a crop and gizzard, before it extends into a somewhat wider sac or stomach, which in its turn narrows into the intestine and ends at the anus. The nutritious parts of the food when dissolved are absorbed almost directly into the blood. A number of long tubules pass into the csophagus and appear to represent the salivary glands of the higher animals, a fluid being discharged which is swallowed with the food; the fluid dissolves certain parts of the food, fitting it to soak through the walls of the alimentary canal so that it can enter into the system. ‘The crop is a sort of food-receptacle, from which the food is passed on to the gizzard, provided with somewhat hard plates for the grinding up of the food, the latter being then passed into the stomach, the walls of which secrete another fluid that renders still more of the food soluble, this part being then readily absorbed by the walls of the stomach and INTERNAL STRUCTURE OF THE BUTTERFLY LARVA. 27 intestine. Near the union of the stomach and the small intestine a number of tubular glands, supposed to represent the liver of the higher animals, open. The intestine ends in a chamber called the cloaca, in which the indigestible and waste portions of the food are collected before being expelled from the body as feces. (8) The circulatory system: The blood circulates, not through actual blood-vessels but, through lacune or hollow channels in the tissues. The blood is collected into a longitudinal membranous sac which is placed just beneath the skin, in the middle of the dorsum or back, and is known as the “ dorsal vessel,” and its rythmic contraction, when it drives the blood into the tissues, can be detected in some of the more thin-skinned lepidopterous larve. In a larva of Brotolomia meticulosa it was observed to beat 44 timesina minute. The functions of the dorsal vessel are analogous with those of the heart in the higher animals, but it consists of only one chamber, although the latter is divided into a num- ber of sacs. The muscular tissue of which it is formed contracts from its hinder part forwards, 7.e., towards the head, and, by its contraction, forces the fluid in it out in front into a number of little vessels which soon come to an end in the little hollow passages or lacune in the tissues. These lacunze are very abundant around the tiny air-tubes which branch off from the trachez, and it is here, after the blood has been over the system, that it is aérated. From here it is carried to the dorsal vessel again to be once more distributed over the system to be again aérated and to be returned again to the dorsal vessel or heart. In vertebrates, the nervous system is placed dorsally, and the circulatory and respiratory systems ventrally,in relation to the alimen- tary canal. These positions are exactly reversed in insects, the nervous system being placed ventrally, the circulatory and respiratory systems dorsally, the alimentary canal being placed between them. It has, however, been shown that this difference is more apparent than real, the dorsum of the insect being really analogous with the venter of the vertebrate, but with the position of the limbs reversed. The dorsal vessel, although consisting of only one chamber, is divided into 8 or 9 sacs, the latter with openings along the sides called ostia. It is composed chiefly of muscular tissue, and is connected with the roof of the body by short stout muscles, which keep it in position. In its passage through the tissues, the nutritious parts of the food, which soak through the walls of the stomach and intestine, enter the blood in the lacune found near these organs. The blood of insects is so ditferent from that of vertebrates, that one feels that it is a great mistake to call them by the same name. Its function is to carry the nutritious matters to the tissues, and to feed, as it were, the tissues it bathes. It is frequently filled with somewhat crude fatty matters, and Graber calls it a refined or distilled chyle. Beneath the dorsal vessel, a fine membrane is stretched in such a manner as to separate the dorsal vessel from the surrounding organs and, at the same time, leave a cavity around the dorsal vessel itself. This cavity is called the peri- cardial cavity or sinus. The membrane itself is incomplete, and, when certain muscles contract so as to pull it down tightly upon the tissues below, the movement at once increases the size of the sinus. The tissues thus pressed upon are full of chyle and blood, and the fluid is squeezed from these structures through the incomplete membrane, 28 BRITISH BUTTERFLIES. into the pericardial cavity, and from thence re-enters the dorsal vessel again. (4) The respiratory system: The air is conveyed into all parts of the body by means of the trachez, elastic tubes, held open by an inner chitinous layer, which are all intimately connected. Large tubes connect the spiracles longitudinally, others pass from one side of the body to the other, whilst a set of trachez in the lower part of the body, is connected with another set in the upper part by ascending tubes. These main branches give out small branches, which fork in all directions, and by them the body is supplied plentifully with air. The tubes have a white glistening appearance, and hence can be readily detected in a freshly killed insect without difficulty. The finest tracheal tubes are supposed to penetrate cells, but it is not known whether they terminate with open or closed extremities. (5) The fat-body: The fat-body is a very prominent part of the structure of lepidopterous larve. It consists of fat masses of various sizes and colours, loosely connected together, and enveloping most of the organs. It varies in colour and appearance in almost every species, and appears to consist essentially of a reservoir, as it were, of reserve material, which increases in the larval stage, when feeding is going on rapidly, and upon which the insect can draw in the future, when it is unable for a long period to take food, ¢.g., at the exuviation of each larval skin, and the more exhausting periods of metamorphosis. It must also be looked upon as material which the insect can utilise, during the period of histogenesis in the pupal stage, in the formation of the imaginal structures. Bessels notes that in Pzeris brassicae, the fat-body is white. Jackson, however, observes that, in LP. brassicae, the fresh fat- body posteriorly to the 6th segment is greenish or olive-yellow, | anteriorly to it, opaque yellow or green on the dorsal aspect, but on the ventral aspect, white. He also says that the fat-body of the larva of Vanessa io is yellow, and that it becomes orange in the pupa. (6) The nervous system: The nervous system of the caterpillar is, in its broad outline, not very dissimilar from that of the butterfly and is very interesting, and its structure helps to explain why it is that, when the thorax of the butterfly is crushed and the insect is, to all appearances, dead, the abdomen, head and antenne continue to twitch and moye, and give to the kindhearted, but ignorant, observer, the notion that one’s cruelty is unbounded in pinning an insect alive to suffer tortures through being spit on a pin when in a moribund condition. The fact is, the central nervous system of an insect is apparently very different from that of vertebrate animals, and is situated in the ventral or belly part of the body, not, as in the latter, in the dorsal. In each of the abdominal and thoracic segments there are two ganglia (little masses of nervous tissue) placed one on either side of the central line. In the head, which appears to be composed of four segments, the eight ganglia are massed together around the esophagus. Hach ganglion is united to its fellow in the same segment by minute transverse nerve fibres, whilst other fibres pass from it in a longitudinal direction to the ganglia of the same side, next in front of and behind it. In addition, the ganglia of the thorax and abdomen give origin to numerous nerves which are distributed to the organs of alimentation and circulation, and to the muscles, those from the thoracic ganglia being chiefly distributed to the muscles that move the wings (when these organs are present). INTERNAL STRUCTURE OF THE BUTTERFLY LARVA. 29 From the ganglionic mass in the head arise the nerves which supply the eyes and antenne, so that this is evidently the nerve-centre for _ such special senses as insects may possess. If now we turn to the vertebrates, we shall find that their central nervous system consists of two distinct parts: (1) The brain and spinal cord, placed in the skull and vertebral column. (2) The sympathetic nerve system, which is composed of a double chain of ganglia running through the neck thorax and abdomen in front of the spine. The nerves which arise from this double chain of ganglia supply chiefly the digestive canal and the walls of the blood-vessels, but some of them join the nerves which spring from the brain and spinal cord, and thus connect the two systems together. The nerves from the brain and spinal cord are principally distributed to the skin and muscles of the body and convey impressions from them to the centre, giving rise to sensation, and from the centre to the muscles, giving rise to movement; those from the spinal cord are provided near their origin with independent ganglia by means of which certain automatic movements, and others called ‘reflex,’ can be carried on without the intervention of the brain. The ganglia and the nerves in the thorax and abdomen of an insect are similarly automatic in their action. When the thorax is pinched the nervous tissue in it is crushed and its functions abolished, so that no pain is felt when a pin is thrust through it. The insect, however, still moves the abdominal segments and the antenna, because the muscles, which effect this movement, derive their supply of nervous force from the ganglia situated respectively in those parts which have not been included in the pinch, and which act independ- ently of each other. It must not be supposed that the ganglia in the head are, to any considerable extent, comparable with the brain of the higher animals; there is no evidence that they are in any special degree the centres for sensation, other than sight or smell, and it is highly improbable that insects feel pain even in the slightest degree. This hurried sketch of the nervous system of insects will help us to understand why a wasp, whose abdomen has been severed from its thorax, will go on sucking up juices, even those exuding from its wounded body, for a long time after the mutilation has occurred. Although, as noted above, apparently so different, the development of the nervous system in the embryo is analogous with that of vertebrates, and, although the nervous system of insects is apparently ventral, whilst that of vertebrates is dorsal, the ventral part of an insect corresponds with the dorsal part of a vertebrate, 7.e., in reality, opposite parts of the body are placed ventrally in insects and vertebrates respectively, owing to the limbs being turned in opposite directions in the two cases. (7) The reproductive system: Herold, as long ago as 1815, figured the changes that he observed the essential reproductive glands to undergo in the larva and succeeding stages of Pieris brassicae, but, up to the present time, there appear to have been no external openings, in connection with the sexual organs, discovered in any lepidopterous larva. The internal glands, however, are not difficult to observe in some larve, and can usually be obtained by a little careful dissection. The testes and ovaries are placed just beneath the skin of the 5th abdominal segment. They exist in pairs, one on either side of the dorsal vessel, just above the position of the alimentary canal. The 80 BRITISH BUTTERFLIES. testes form two lobes of a not very distinctly reniform shape, wh.lst the ovaries, which are only to be seen with a lens, and then in comparatively few species, are much smaller and consist of tubes. The testes are generally much more readily observed than the ovaries, being usually yellow or brown, and may be seen distinctly in the larve of those species of lepidoptera that feed internally, or that have fairly transparent skins. Jackson says that the larval ovaries are situated in the 5th abdominal segment and close to the dorsal middleline. Their proximal or attached extremities are approximated, and they diverge from one another posteriorly. The colour gets deeper during the quiescent period preceding pupation. Four opaque white lines, the future ovarioles, traverse the larval ovaries lengthwise, and converge towards their hinder extremities, from which the larval oviducts sprmg. The latter are very delicate filaments, and difficult to make out. _Bessels notes of Pieris brassicae that the ovary is yellow, the testes violet, the fat-body white. CHAPTER IX. THE ASSOCIATION OF ANTS WITH BUTTERFLY LARV#. A matter, of which little is known, has here to be considered, viz., the character of the association of ants and certain Ruralid larve. On the middle of the dorsum of the 7th abdominal segment in these larve is an evaginable gland, which, in some species, secretes tiny drops of sweet fluid, greedily lapped up by ants, that stroke the larve with their antenne until the desired liquid is obtained. Other allied larve are said to possess the gland, without, however, its having any secretive powers, and it is recorded that these larve are not accompanied by ants. Until, however, many more observa- tions have been made, this latter conclusion must be accepted with great caution. It is assumed that, whilst the ants benefit by obtaining the saccharine fluid desired, their presence tends to warn off ichneumons and other enemies that prey on the larve. Seudder observes that ‘“‘it is a curious thing that, among the Lycaenidi, the glands are found in some species, whilst not in others closely allied; their presence in many members of the other two tribes of Lycaenidae, together with the impossibility of their independent origin in different genera, render it probable that these glands first arose as long ago as before the differentiation of the three Lycenid tribes; the brotherhood of the ants and caterpillars may, therefore, be of great antiquity.’ Seudder says that Esper was the first to notice the relation of the larva and the attendant ants, and that Guenée observed the gland in the larva of Lampides boeticus, describing and figuring it, whilst Freyer figures the gland as two white dots in the larva of Plebeius argus, but does not describeit. Scudder observes that the gland is present in the larve of many hair-streaks, and in that of T'hestor ballus, although the association of ants with these larve has escaped notice. Edwards gives (Can. Fnt., x., pp. 181-186) a detailed account of the connection between the larve of Cyaniris pseudargiolus and their attendant ants, He noted that ants frequented the same THE ASSOCIATION OF ANTS WITH BUTTERFLY LARVE. 31 flower-spikes as the larve, and thought they were attracted by the nectaries of the flowers, until he observed an ant running up and down _ the back of one of the larve, drumming and gesticulating with its antenne, the feeding larva not at all disturbed by the treatment. Three kinds of ants were observed, and, on one occasion, six examples of a small ant were seen to be busy with one larva at the same time, but the movements of all the species were similar. They run over the body, caressing the larva incessantly with the antenne, and undoubtedly with the object of persuading it to emit the fluid. Much of the caressing is done about the anterior segments, and, while the ants are absent from the posterior segments, the tubes (on the 8th abdominal) are almost constantly exposed to their full extent, and so remain, without contracting, until the ants come tumbling along in great excitement, and put either foot or antenna directly upon, or close by, the tubes, when these are instantly withdrawn. The ants pay no heed to these tubes, so far as touching them with intention, but at once turn to the median gland, caress the back of the 7th abdominal segment, put their mouths to the orifice, and show every sign of eager expectancy. With a lens, a movement will speedily be apparent, a dark green mammilloid membrane will protrude, from the top of which exudes a tiny drop of clear green fluid. This the ants drink greedily, two or three of them perhaps standing guard over it. The demonstra- tions of the ants are of the most gentle nature, caressing, entreating ; and, as the little creatures drink the fluid, they lift their heads as if to prolong the swallowing. There is a manifest satisfaction and delecta- tion that is amusing to see; they lick away the last trace and stroke the back of the segment, and wait to see if their coaxing avails anything ; if not, they run about, but presently all return and the caressings go on as before. The intervals between the appearance of the globule vary with the conditions of the larva; if exhausted, by yielding to the frequent solicitations, some minutes may elapse, and the tubes, meanwhile, will remain concealed; but a fresh larva requires little urging, and the mere intimation of the presence of an ant in the vicinity is enough to cause the tubes to play rapidly, and one globule to follow another, sometimes without a retracting of the membrane and before the near approach of the ants ; six emissions were once counted in 75 seconds. The tubes are usually expanded when the ants are away from the posterior segments, and are retracted when they come near; counting the length of these periods of complete and quiet expansion, they were found to be 10, 20, 50, up to 80, seconds, the period always ending with the approach of the ants. HKixperiment, by placing larve upon stems of the growing plants where the ants had access to them, showed that, as soon as the ants discovered one of them, there was an immense excitement and a rush for the hinder larval segments; the larva forthwith relieved itself by the excretion of the fluid, and the tubes stood out with tops expanded between the periods. A larva, placed on a stem on which there were no ants, showed no excitement, no appearance of the tubes, and no movement of the median glands; if ants were transferred to the stem, the larva at once changed its behaviour. Scudder adds that, it is only in the later stages that the ants attend the caterpillars, or any fluid is excreted from the median gland, though the organs are certainly present at an earlier stage. HKdwards further finds the attendance of the 32 BRITISH BUTTERFLIES. ants to be confined to the summer broods of caterpillars of Cyaniris pseudargtolus, and, even then, to those on Cimicifuga, and suspects that the larve feeding on Cornus or Actinomeris cannot exude so sweet a fluid, the flower of Cuimicifuga being of exceeding sweetness, whilst that of Actinomeris is bitter to the taste. Saunders notes (Can. Ent., x., p. 14) that the larve of Rusticus scudderit (closely allied to the European Plebeius argus) are accompanied by ants, and that the discovery of the larvee was made comparatively easy from the invariable presence of these active attendants. The ants were observed actively running about the leaves on which these caterpillars were found, and repeatedly over the caterpillars themselves, which did not seem in the least disturbed by them. Edwards also gives a most interesting account of the connec- tion between ants and the larva of another Lycenid, Rusticus melissa (Papilio, iv., pp. 92-3). He notes that, on June 9th, he introduced a small ant to a larva of this species, which was confined in a glass tube. The ant soon discovered the larva and ran about it in great excitement, caressing it with its antenne. Immediately the tubes, not hitherto seen, began to play, and one or the other, or both together, were exposed for some minutes, and, indeed, so long as the ant was near. Sometimes the tubes were fully protruded, with the tentacles expanded, at other times they were partially withdrawn, in that case coming together in a pencil just as has been observed in C. pseudargiolus. The ant always ended its caresses by putting its mouth to the orifice of the gland on the 7th abdominal segment; and, by its motions, evidently found the fluid it sought. Next day, two ants of a larger species were turned in at the same time; they ran about the glass, alarmed at finding themselves in confinement, and accidentally one soon touched the larva; at once a drop of green fluid bubbled out of the orifice before the tubes made any movement. The ant saw it, rushed at it, and then the tubes began to play although they had been quiet for fully five minutes before; they now played intermittently for two or three minutes, the tentacles fully expanding and then partly retracting. The ants drank of the drops four times and then desisted, running about the glass again; then they were liberated, and a small ant of the species experimented with the preceding day, was introduced ; almost at once it found the larva, caressed it gently, and was favoured with the coveted nectar, the tubes being all the time in motion. On June 12th, the larva now being mature, another ant was introduced ; as usual, as soon as the manipulations began, the tubes commenced to play, and, presently, a large drop issued ; in ten seconds another followed, but for some time after there was no more, though the ant begged urgently for it. The ant left the orifice, ran up and down the body of the larva, caressing the anterior segments, and then returned to the orifice and begged again; this was repeated several times, but the larva was obdurate, probably it was exhausted, being near pupation. The solicitations are made by the antenne alone, which fly about drumming here there and everywhere, the ant manifesting sreatexcitement. HKdwards notes that he was observing larve of Cyaniris pseudargiolus at the same time, and the behaviour of the two species was identical. Nicéville observes that many Lycenid larve in India are provided with an oval opening on the dorsal line of the 7th abdominal THE ASSOCIATION OF ANTS WITH BUTTERFLY LARVA. 33 segment, with lips ike a mouth. These lips can, at the will of the larve, be somewhat protruded, and a drop of sweet liquid exuded. The larve possessing this gland are greatly affected by ants of different species, which, in return for the food they obtain from the larve, act as their most efficient guardians. He says that he has found as many as four species of ants attending one species of larva. Ant- tended larve are most easily found by looking for the ants. The larve are usually coloured like the leaves, buds, flowers and seedpods on which they feed, and, for other reasons, are not easily seen; but the restless red or black ants are very conspicuous. Curetis larve, which are not attended by ants, have a highly-developed eversible organ on either side of the 8th abdominal segment, apparently for protective purposes; in other larve, attended by ants, the organs on the 8th abdominal segment are smaller than in Curetis, and are, one supposes, gradually becoming aborted, probably because, the ants having constituted them- selves their defenders, there is no further use for them for defence, but Kdwards possibly correctly surmises that in their aborted condition they serve as signals to the ants to examine the 7th abdominal segment for the sweet fluid emitted by the larve. Doherty has recorded (Journal As. Soc. Beng., lv., pt. 2, p. 122) some interesting observations on the same subject ; so also has Mrs. Wylly (Journ. Bomb. Nat. Hist. Soc., iii., p. 164). Not only do the ants attend the larve from their very first and smallest stages (some ants were found attending larve of Rapala schistacea only 4 in. long), until they are fullgrown, but they often cause the larve to change to pupe within their nests, in this manner protecting them from harm from the time they emerge as minute caterpillars from the egg, until they assume the pupal stage. Nicéville also mentions that Aphnaeus vulcanus is attended by the black ants, Pheidole quadrispinosa and P. cremastogaster, and that Gerydus symethus and Tarucus theophrastus are also attended by ants. Green says, ‘‘ The larve of a Cingalese Lycenid, Aphnaeus lohita, Horsf. (=lazularia, Moore), frequent the nests of Cremastogaster, on Acacia and Grevillea trees, upon the foliage of which they feed. These larve carry a dorsal honey-gland near the posterior extremity of the body (7th abdominal segment), and are cultivated by the ants on this account. They are herded in special shelters built by the ants, are driven out at night to feed, and brought back to their shelters each morning” (Fint., xxxv., p- 202). this subject, Doherty writes (Journ. As. Soc. Beng., lv., pp. 122- 173) : “Dr. Thwaites (mn Moore’s Lepidoptera of Ceylon) says, ‘ Nature, however, finds a protection for these helpless Lycenid larve, in the instincts of an ant, Formica smaragdina, Fab., which, finding a sub- stance most palatable to it, secreted naturally from a glandular defined spot upon the body of the larve, takes possession of them as cows, surrounding each separate one, and the leaf on which it feeds, protect- ing them jealously and attacking most fiercely any living thing intruding upon them.’ Besides a remark of Herrich-Schiffer’s, quoted in Distant’s Rhopalocera malayana, that Gerydus symethus inhabits ants’ nests, I have met with no other mention of this singular habit. I have, however, myself observed it in quite a number of Indian Lycaenidae, belonging to several distinct groups, and feeding on the leaves of various trees and herbs. The larve in question, are all very helpless and inactive grubs, slug-like in shape, tapering at both ends, 34 BRITISH BUTTERFLIES. pubescent-green or brown, with a very small pretractile head. On each side of the 8th abdominal segment above, there is a short pro- tuberance, from which can, in most cases, ¢.g., Tarucus theophrastus, be extended a brush of hairs, and apparently absent in Azanus ubaldus. This is, I have no doubt, a scent-gland, and may be intended to attract the notice of the purblind ants. On the dorsal line of the preceding segment, there is another short tubercle exuding a viscid juice. It exists in all the Lycaenidae known to me, whether they are maintained by ants or not, and from it issues a gummy thread, by the aid of which, I believe, the caterpillars sometimes swing themselves from branch to branch, or attach themselves to leaves. But, though in all probability acquired for such purposes, it is peculiarly attractive to ants, which, at all hours, surround the caterpillar, and, -by stroking and tickling it with their antennz, induce it to yield up this sweet (?) liquid. I have not yet found any caterpillar in the possession of web- making or arboreal ants, such as Formica smaragdina, and no restraint, such as Dr. Thwaites mentions, was placed upon any larva observed by me ; but the ants would always remain near the caterpillar, and would always fly fiercely to the rescue if anything molested it. When it had attained its full-growth, the ants, forming a circle round it, would forcibly drive it down to their nest at the foot of the tree. This sight is rather an amusing one, the caterpillar often showing the greatest reluctance to leave its pasture ground, and manifesting strong doubts as to the intentions of its escort. 1 was struck with the forbearance and patience of the ants which carefully abstained from any violent use of their formidable jaws, though the journey was sometimes pro- longed to six or seven hours. Having arrived at the foot of the tree, the ants deposited the caterpillar in an open space just within the mouth of the nest, whereupon the latter would attach itself to the bark, and there commence its transformations. I have counted as many as thirteen chrysalids of Azanus ubaldus so attached, in one nest, at the foot of a kind of babul tree, Acacia leucoplaea. The instinct which induces the ants to preserve these caterpillars in their nests, thus sacrificing a large present supply of food to the possibility of a future supply of the sweet juice they are so fond of, strikes me as one of the most remarkable things in nature.”’ Nicéville gives (Journ. Bomb. Nat. Hist. Soc., i., pp. 164-168) further details on this subject, observing that, in Calcutta, he has found the larve of over a dozen Lycaenidae affected by ants. The most important part of the paper, however, consists of a series of observations on Tarucus theophrastus, Fab., by Mrs. Wylly, who writes: ‘‘The larve of YVarucus theophrastus are cultivated and protected by the large, common, black ants of Indian gardens and houses. ‘The caterpillar, which varies in colour from light pure green to a dark reddish tint, is about 75 in. in length, louse-like in shape, and slow in movement, feeding on Zisyphus jujuba, with an edible astringent yellowish fruit. On the dorsum of the 7th abdominal segment is a small slit from which the larve exude a small drop of a juice of some sort, eagerly sought by the ants, and which they can generally procure by stroking the larve gently with their antenne. The ants set up what appears to be merely a temporary nest at the foot of the tree the better to carry on their operations. Just before the rains set in, about the middle of June, great activity among the THE ASSOCIATION OF ANTS WITH BUTTERFLY LARVE. B5 inhabitants of a Zisyphus tree may be observed. The ants are busy all day long running along the branches and leaves in search of the larve, and when they meet one fullgrown and ready to pupate, they drive the caterpillar down the stem of the tree towards their nest. As a rule, the larve are docile and easily led, and, having got him into his proper place he undergoes transformation into a pupa. If one gently scrapes away the loose earth piled up at the base of the tree, one will see some hundreds of larve and pupe in all stages of develop- ment.arranged in a broad even band all round the trunk and lightly covered with earth. The ants object to their being uncovered, and will immediately set to work to recover them, and, if one persists, they will remove all the chrysalids and bury them lower down. ... . A larva ofa species of Catopsilia (one of the Pierinae) given to the ants as aN experiment, was immediately set upon and torn to pieces in a second by the ants. A larva of T. theophrastus, taken from a tree, was introduced into the pathway of another company of the same species of ants, which lived on our verandah, but kept no “ farm.”’ It was odd to see the ants come tumbling over headlong to fight the intruder, and the sudden way they cooled down on investigation of the foe. None attemped to harm him, and he was politely escorted across their boundary, the ants running alongside, and feeling him all over with their antenne. This must have been instinct as they could have had no former knowledge of him as a “ milk-giver.”’ The ants distinguish between dead and living pups, the dead chrysalids being carefully removed and thrown away outside.” Distant mentions that the larve of the genus Amblypodia are attended by Formicasmaragdina, Fab. Bethune-Baker says that the larve of almost all the species ofthe Australian genus Ogyris are probably attended by ants, some apparently by different species in different neighbourhoods, whilst Dodd records that O. zosine is attended by two species of Camponotus, and also by a small black ant (Trans. Ent. Soc. Lond., 1905, pp. 269-270). Of O. zosine, Bethune-Baker writes (op. cit., p. 279) that the larve feed on Loranthus linophyllus and L. celastroides, and hide in the cracks of the bark of the host tree . . . . coming out at dusk and feeding at night, at which time the ants associated with them are likewise said to be on the move. The species that Dodd has found them with most commonly is Aicophylla virescens, but several other species also associate with them. They evidently protect the larve, and have been observed to milk them; in one instance, an ant was observed to approach a larva and wave its antenne over its terminal segments, and then to lightly touch it with its foreleg, when a small globule of Jiquid was emitted from a small, retractible, nipple-like organ on the dorsum, which was at once sucked up by the ant. Lyell and Fricot (Vie. Nat., xxi., pp. 166-167) have confirmed these observations, and state that, in order to test the action of the attendant ants, one or two larve were placed a couple of feet or so away from a tree; they were, however, soon discovered, and dragged carefully back to the tree by the ants, at a pace much more rapid than their own rate ; pupz were likewise carried back to the tree. It is recorded also that “ants are always found with the larve of O. abrota. Anderson further observes (Victorian Butterflies, pp. 101-102) that the larve of the genus Ogyris are greatly attractive to ants, which tend them with great care, never leaving them. Raynor notices (in 36 BRITISH BUTTERFLIES. litt.) that, at Paramatta, N.S.W., from 1877-1880, he frequently found the larve of Lalmenus ictinus, a Theclid superficially resembling our Zephyrus quercis, feeding on Acacia decurrens; at first, he obtained them by beating, but afterwards by searching, when he was greatly surprised to find numbers of ants running excitedly backwards and forwards over the larve. He says that it seems to him most interesting that the power of exuding an attractive secretion should extend to the far-off region of Australia. Anderson says (Vict. Butts., pp. 98-99) that the ants affiliated to the larve of J. ictinus are particularly large and fierce. He also observes that the larve of the allied J. evagoras are gregarious, and invariably attended by ants. The connection of at least one British species, Lycaena arion, with ants, has been observed and commented on. Frohawk noticed that the butterflies of this species showed a preference for laying their eggs upon thyme plants growing on the nests of Formica flava, and suspected some connection between the ants and larve. He placed a living larva of L. arion, that had passed its 8rd moult, into a box with four examples of F’. flava. They immediately ran to it, and, waving their antenne over and upon it, apparently smelt and licked it, and seemed particu- larly attracted to the hinder part of the back, about the 10th segment, i.e., the 7th abdominal segment. First one and then another of the ants would run over the larva, and then stop to lick that part of its back. He then noticed a tiny bead of moisture appear, and one of the ants touched it with its mouth, which instantly caused the bead to disappear. Hxamination of the larva and ants under the microscope showed a small elongated transverse gland on the dorsum of the 10th seoment. Examination of another larva in the same stage showed the eland which kept throbbing while the larva was feeding. The ants were placed close to the larva, and they soon ran over it. Directly a foot touched the gland, or a place very near it, it immediately throbbed more violently and swelled up, and then ejected a globule of clear white liquid, which was immediately licked up by an ant. In a few seconds a foot again touched the gland, and another bead of liquid oozed out, which was at once again licked up by an ant. An interest- ing fact is, that the larva unheeded the ants running over and around it while it kept feeding, but the gland is apparently exceedingly sensitive to the touch of an ant’s foot, and, although Frohawk several times touched the glands of several larvee with the point of a very fine sable- hair brush, they would at once wince and contract, but on no account could the exudation of the liquid be induced, yet directly an ant’s foot, or the claws of the foot, touched it, a bead would appear, and at once be imbibed by the ants. Although the larva was kept in a box with numerous ants, both workers and winged females, together with their pupe, the ants one and all acted precisely similarly; not one attempted to bite the larva, but, as soon as they touched it, they slowly closed the jaws and waved their antennz over and upon it. The gland is of peculiar construction, being formed of flexible tissue, and surrounded by numerous glassy-white pyriform processes varying in size; some are extremely minute, those bordering the edges of the gland are furnished with excessively small white bristles, each process bearing four or five; these are in the form of a fan with diverging points, and all are directed towards the central aperture, the whole forming a fringe surrounding the gland, and are obviously for the purpose of holding CARNIVOROUS HABITS OF BUTTERFLY LARVE. 37 the bead of liquid in place, and probably also serve as a protection to this apparently sensitive organ. The larve appear to be perfectly at - bome with the ants, neither molesting the other (Hntom., xxxvi., _ pp. 58-60). CHAPTER X. CARNIVOROUS HABITS OF BUTTERFLY LARVE. The fact that, under certain conditions, in confinement, lepidopterous larve will live on others of their own or different species, is well known, and the habit of the larva of Thecla w-album to leave its food and feast on the newly-formed pupz of its own species, has frequently been observed and recorded. That certain butterfly larve should, however, have a permanent carnivorous diet, is sufficiently unusual for us to devote a short space to the details of one or two of these cases. The best known of these speciesis Feniseca tarquinius, an American insect which Scudder makes a Chrysophanid, and which is undoubtedly a, Lycenid in sens. lat. The larva of this butterfly is purely carnivorous. The eggs are laid in the midst of a group of aphides, or near thereto, and, for protection, are coated during deposition with a thin coagulated albuminous deposit, which, on hardening, covers them like a thin but irregular veil, the egg-stage lasting only three or four days. The larva appears to live entirely on plant-lice, particularly affect- ing the species Schizoneura tessellata (on alder), Pemphigus frawinifolit (on ash), P. imbricator (on beech), all of which produce much flocculent and saccharine matter; it has also been fed on aphides from willow and plum, in confinement. The young larva eats a hole through the summit of the egg, and pushes its way under the larger aphides, and forthwith begins to spin for itself a loose web, not close enough to conceal it from view were the aphides away, but sufficient to keep the aphides from walking over the body, and to protect it when a moult ig approaching, and the skin sensitive. The web seems to be just about the length of the larval hairs from the body. The aphides may be seen running over it, and often get their legs fast in the meshes, and are apt to be devoured as a consequence; the larve appear to pass both the first and second moults beneath this web, but, after this, seek fresh supplies of food, devouring the aphides from the underside, their backs covered with wool from their victims (Butts. New Engl., i1., pp. 1022 et seq). The most remarkable fact, however, connected with this larva, is that the ants, which nurse other Lycenid larve, are its sworn enemies, for, by feeding on the aphides that. the ants keep, they destroy the source of supply of the ants’ sweet food (secreted by the aphides), and are furiously attacked and killed by the latter; it appears only to be in their later stages, when feeding largely exposed, that the ants are able to successfully deal with them. Kershaw gives details (Trans. Ent. Soc. Lond., 1905, pp. 1-4) of the connection between the Chinese Gerydus chinensis and aphides, chiefly those frequenting various species of bamboo, the eggs being laid among a crowd of aphides, and often hidden under _a@ mass of them. They hatch in about four days, the larva at 38 BRITISH BUTTERFLIES. first being cylindrical, and later slug-shaped, the head retractile. The larve feed on the aphides, pressing them against the plant with head and forelegs, sometimes holding them in the forelegs quite away from the plants; a few bites dispose of an aphis, and the larva then licks and cleans its legs, just as a mantis does; some aphides must have a better flavour than others, as the larve pick and choose, moving their heads up and down over the backs of the insects, evidently smelling them; as a rule, the creatures seem to make little attempt to escape till they are actually bitten, when struggling is useless. When not engaged in feeding, the larve rest among the aphides, or crawl about between, or over, them, and the aphides do likewise, the larvee sometimes covered with them. The larve have been observed to feed only on two kinds of aphides, one slate-coloured with white efflorescence, the other greenish, with four dark green patches, some of them being fringed with white, probably moulted skin. The larval state lasts about fifteen days, and it was reckoned on the average, from first to last, that a larva ate some twenty aphides per day, but it would require many larve to make much impression on the crowds of aphides one sees, for often a yard of bamboo stem, two or three inches in diameter, is absolutely covered with these insects. One of the features of this larva is the calm way it moves about among the aphides and selects its prey, and the indifference with which the latter apparently accepts its fate. Although Kershaw notes that the aphides are overrun by a host of ants of the two species, Polyrrhachis dives and Dolichoderus bituberculatus, he notes no signs of enmity between the ants and larve, possibly because of the abundance of the aphides. Holland records (Can. Fint., xix., pp. 61-62) the receipt of a large ? of Liphyra brassolis, captured in Penang, upon which was, at the time of capture, a quantity of fluff-like mildew, particularly thick on the abdomen and underside of the wings. This fluff was proved to be the mealy covering of certain ‘‘shield lice,’ a few specimens of a large species of which were in the same consignment. Holland at once concluded that—(1) The captor had caught the specimen of Liphyra near a colony of scale insects, which were so large as to attract attention, and lead him to put a few into papers. (2) This 2 was engaged in oviposition just before she was captured, and that (8) the mealy-white deposit described as “ fluff,’ and which was compared with mouid or mildew, was nothing less than the fragments of the white covering of the scale-insects over and among which the butterfly had been flying whilst engaged in the act of laying her eggs. Dodd observes (Hnt., xxxv., pp. 158 et seq.) that, in July, 1900, he noted a 2 of Liphyra brassolis depositing eggs upon a tree in complete possession of the wonderfully interesting green tree-ant, U’cophylla smaragdina, which exists in vast numbers in Tasmania, on the coast and mountain scrubs. Upon this tree were several nests of the ants, and several eggs were deposited singly on the tree, on the underside of branches, or protected side of the trunk. Searching the ants’ nests for larvee was unsuccessful, until a half-erown larva was found accidentally, whilst searching an ants’ nest for other insects. After this, other larvee were found, the lozenge-shaped body peculiarly flattened, the head, legs, and claspers, in a groove, the edges of the body closing down tightly all round, except during progression, when the body is raised a little, On one occasion a larva was seen to deliberately seize a half- OARNIVOROUS HABITS OF BUTTERFLY LARVA. 39 grown ant-grub, which, however, was released when the larva was turned over. Even giving them small ants’ nests in confinement was not successful, and they had to be returned to the nest in order to feed-up successfully to pupation stage. Dodd’s further remarks are to the effect that the larve move from nest to nest, that they are so tough- skinned that the mandibles of the ants can make little or no impression upon them, and to protect its head and legs the larva just lowers its sides, and is secure. Although the evidence here offered is sufficient to lead us to accept the fact that the larva lives in ants’ nests, it is by no means satisfying that it lives on the grubs of the ants. Green observes (Hnt., xxxv., p. 202) that the evidence that they are really carnivorous appears to be proved by the fact that they seized, and attempted to eat, some of the grubs, but he further observes that they do not appear to have been satisfied with that diet, and asks whether it is not possible that their proper food may be some species of Coccid enclosed in the ants’ nests. He says that, ‘‘in Ceylon, the arboreal nests of this same ant almost invariably include colonies of Cocczdae, Aphididae, and Aleurodidae, and adds that there is, in Ceylon, also, a coccidophagous Lycenid larva, viz., that of Spalgis epius, which he has, on more than one occasion, found inside nests of another tree-ant, Cremastogaster dohrni, feeding upon ‘mealy bugs,’ Dactylopius sp., enclosed therein.” Green communicated his observations on the carnivorous habits of Spalgis epius to Nicéville, who published them in his Butts. of India, vol. ili., pp. 55-56. He says that Spalgis epius has been several times reared from a larva that associates with, and feeds upon, the mealy- bug, Dactylopius adonidum. The larva is dull olive-green above, with numerous minute dark bristles and a lateral fringe of brown hairs ; beneath pale green, slightly suffused with pink on anterior segments. It partially covers and conceals itself with the mealy secretion from the Dactylopius. Independently, Aitken gives (Journal Bomb. Nat. Hist. Soc., vili., pp. 485-487) an account of the carnivorous habits of the same species, a ? of which he saw in December, 1891, flying about a bush absolutely infested with ‘‘ mealy-bug,’’ some of which appeared to be suspiciously large, and which, when the white woolly secretion was brushed off, proved to be Lycenid larve. A lateral fringe of bristles, continued round the prothorax, was immediately used by the larva to shovel a quantity of the white stuff on to their backs and clothe their nakedness, after being.denuded. They were then seen to be feeding on the mealy-bugs, burying their heads in the down covering them. A number, secured and placed in pill-boxes, fed up on the mealy-bugs, and pupated in due course, and, in a fortnight, imagines of Spalgis epius emerged. In 1891, Holland received (Psyche, vi., pp. 201-202) larve of Spalgis s-signata from Kangwe, on the Ogové River, in West Africa, collected by Good, who found the dark brownish larve on the leaves of a Frangipanni, the body all covered over with a whitish substance, and which was assumed at once to consist of the remains of plant-lice, with which the undersides of the leaves, on which the larve were found, abounded. Suspicion was at once aroused that the caterpillars must have fed upon these white plant-lice, because no leaves appeared to be eaten. The white fluffy substance was readily rubbed off, but there was sufficient left. to prove that it was really the remains of the plant-lice, 40 BRITISH BUTTERFLIES. and Holland says that, ‘‘examined under a powerful microscope, this adhering matter is seen to present a peculiar shining appearance, and to thickly cover the hairs with minute granulations, as if each hair had been dipped in some substance like a solution of sugar or salt, and had then been dried.’’ There appears to be no doubt of its aphidivorous habits. Holland further states that he believes Lachnocnema and Euliphyra to have similar carnivorous habits. CHAPTER XL. COLLECTING BUTTERFLY LARVS. The larve of some butterflies are, in certain seasons, from the gardener’s point of view, too abundant. Such is the case with the larve of Pieris brassicae and P. rapae, which are occasionally sufficiently numerous to do considerable damage to the cabbage crops. The larve of Aglais urticae and Vanessa io are gregarious on nettles, the former usually abundant enough in their thickly populated silk nests, the latter much more irregular in their appearance. Other gregarious larve are Melitaea aurinia and M. cinwia, but, although the black spiny larvee of the former can usually be taken in the haunts of the species, those of the latter are to be found only in a few places in the Isle of Wight, it having been almost exterminated as a British insect. The larvee of Papilio machaon can only be sought with success in their local haunts in the fens of Cambridgeshire and the Norfolk Broads; and those of Strymon (Thecla) pruni in the woods of Huntingdonshire. In the southern counties, the larve of Gonepteryx rhamni can be readily found on buckthorn, and those of E'uchloé cardamines on Cardamine pratensis and Alliaria officinalis. The larva of Apaturairisis confined to sallowin woods, and that of Limenitis sibylla to honeysuckle, although these, especially the former, are to be obtained more frequently by beating than by searching. Beating is also pursued for larvee of Bithys (Zephyrus) quercis on oak, Ruralis betulae on blackthorn, andStrymon (Thecla) w-album on elm. Sweeping at night, with a strong sweep-net, by the grassy sides of woods, hedges, the sheltered hollows of chalk-hill slopes, ete., will produce larve of the Satyrids, and occasionally of the Argynnids and Urbicolids, in spite of the fact that the latter live in silken nests, whilst grassy hollows on the mountain-sides will give larvee of Hrebia aethiops, and, in its haunts, the local Melampias epiphron. We have already dealt with this phase of our subject, at length, in Practical Hints for the Field Lepidopterist, in which the collecting work to be carried out each month, for lepidoptera in all stages, is set out at considerable length. The following are suggestions of work that can be done in the various months, in collecting butterfly larve, whilst the reader can, by a careful study of the systematic part of our work, especially of the paragraphs ‘Habits of Larva,” etc., gather much more information similar to the following. Fesruary AND Marou.—The larve of Rumicia (Chrysophanus) phlaeas are sometimes very common on Rwmea acetosa in February and March ; they are difficult to see, as their bodies are about the same size as, and COLLECTING BUTTERFLY LARV&. 41 the crimson dorsal line and broader spiracular stripe render them very like, the young curled-up leaves in the centre of the plant. The hybernating larve of Polyommatus icarus are sometimes to be found on Lotus corniculatus, when searching for cases of Coleophora discordella. In March and April the larve of Aricia (Polyommatus) astrarche feed on the undersides of the leaves growing on the young tender shoots of Helianthemum vulgare, making marked brown blotches where they feed, and thus betraying their whereabouts; they are fullfed from the middle to the end of May. Have a plant of Hippocrepis comosa ready to place the larve of Plebetus aegon upon, as soon as they leave their eggs, which they always do either in the last few days of February, or the very first days of March. The larve of Pararge megaera are to be obtained, feeding on grasses on the outskirts of woods, by wild hedgesides, etc., in March, the larve being fullfed, and pupating, in early May. Aprint.—tIn late April, the nearly fullfed larve of Polyommatus icarus are to be found on Lotus corniculatus and Ononts arvensis. The larve of Agriades (Polyommatus) bellargus are to be obtained on Hippocrepis comosa in April and May; they pupate about the middle of May, and emerge in June. The very earliest larve of Celastrina (Cyaniris) argiolus are to be found just hatched at the end of the month, feeding in the buds or flowers of holly ; later, in May and June, they attack the young tender leaves and shoots, upon which they thrive. The larve also feed well on young ivy leaves, and on the tender leaves, and young green berries, of Rhamnus frangula. The larval colonies of Melitaea aurinia are found, in April and early May, on scabious and honeysuckle, but the eggs always appear to be laid on the former. The larve feed up well in confinement on honeysuckle. In confinement, the larve of Melitaea aurinia appear to be very susceptible to warmth, collecting in the hottest part of the cage, and becoming lively when the sun ison them. ‘They are much better fed up, however, on a growing plant than in a breeding-cage. The larve of Melitaea cinxia are to be obtained in their restricted haunts in the Isle of Wight, towards the end of April; they are gregarious, and hence the capture of one or two at this time usually means the capture of a brood. The larve of Argynnids—Dryas paphia, Argynnis adippe, A. aglaia, Brenthis euphrosyne, B. selene—should now be sought on various species of Viola. They feed in the daytime, but are usually well-hidden. The larva of Brenthis selene feeds on Viola canina, appears to have an aversion to the sun’s rays, reposing either on the undersides of the leaves, or on the stems shaded by the leaves, selects always the youngest and tenderest leaves until near maturity, eating out large portions of them, and making its whereabouts conspicuous. The larve of Brenthis euphrosyne, approaching full-growth in April and early May, are to be found by searching the leaves of Viola canina and primrose, where there is much sign of the plants being eaten ; they generally hide, and are to be found on the underside of a leaf, but, when the sun is shining, love to bask in it, and are very active, retiring, how- ever, aS soon as. the sun disappears. Hedgesides, and the ridings of woods, will give larve of Pararge 42, ‘BRITISH BUTTERFLIES. egertia, P. megaera, Enodia hyperanthus, Epinephele tithonus, etc., in their respective localities. These are best obtained by sweeping, or searching with a lantern by night. Larve of Melanargia galatea, Hipparchia semele, etc., can similarly be obtained in their known haunts. The larve of H'rebia aethiops may be collected in abundance at night, with a lantern in the local haunts of this species. The hybernating larve of Pararge egeria are to be obtained, in early April, by the sides of the ridings and paths in woods, feeding on grass (Dactylis glomerata) ; the most. forward spin up before the end of the month, and the imagines usually appear in May. The larvee of Coenonympha typhon can be obtained on the moors, in the early spring, on the beaked-rush (Rhynchospora alba), on which they slowly feed up, being fullfed in early June. The larva of Hipparchia semele should be swept for, in its known habitats, in April and May, by night, when it comes up to feed ; it hides by day, often beneath the surface of the ground. The young larve of Polygonia c-album may be sleeved out on hop, stinging-nettle or currant. They must, when they first leave the egg, be fed on quite fresh young leaves, if they are to be reared successfully in confinement. The young hybernating larva of Limenitis sibylla begins to move in early April, moults almost at once, becomes reddish-brown in colour, and spiny, feeds on the fresh bursting honeysuckle-buds, and, by the middle of May, has usually assumed a miniature resemblance to the adult larva. In the last week of April and early May, search for the larva of Limenitis sibylla; it is then in its brown stage, and rests on the brown stem of the honeysuckle just below the green shoot, generally low down on the bush, in a sheltered position. Sometimes the larve may be found on the green leaves, where they are much more conspicuous than on the stems. Young larvee of Limenitis stbylla, reared in confinement, should be fed on young and tender shoots of Lonicera periclymenum. They nearly always commence to feed at the top of a shoot, and eat their way down- wards, being especially fond of the sun, and always eat greedily when the sun is shining on them. The young larve of Apatura wis may be beaten from sallows. The lowest and most unpromising-looking bushes are often the most productive, when working for this species (see Hnt. Record, vi., pp. 146-147). In the spring, the larve of Apatura iris feed up quickly; they eat at night, and rest in the daytime on the midrib of the upperside of a leaf, the head turned towards the base of the leaf. One ought to be able to find them by the leaf hanging down with the weight of the larva ; otherwise they are almost undiscoverable. May.—The larvee of Thymelicus acteon are to be found in May and June, on the sea-slopes from Swanage to Weymouth, feeding on the leaves of Brachypodium pinnatum. In confinement, they will eat Triticum repens and allied grasses. The presence of the larva of Thymelicus acteon is best told by the wedge-shaped pieces which they eat out of the sides of the blades of Brachypodium pinnatum. When such traces are observed, search for COLLECTING BUTTERFLY LARV2. 43 the tubes in which they hide, and which are made by spinning the two edges of a leaf together, so as to enclose themselves therein. The fullfed larve of Augiades sylvanus are to be found in early May on Luzula pilosa, the edges of a leaf of which are folded over, and lined with silk, to form a puparium, in which the larva changes to a chrysalis. The larve of Aricia (Polyommatus) astrarche var. salmacis are to be taken in late Mayandearly Juneon Helianthemum vulgare, in the northern counties of England. From pupe, formed from larve obtained June 8rd, 1877, near Hartlepool, three imagines— apparently salmacis, . artaxeraes, and astrarche, on the upperside, but more like salmacis on the underside, emerged. The larve of Aricia (Polyommatus) var. artawerxes are to be found on the undersides of the leaves of Helianthemum vulgare throughout May, their colour assimilating remarkably well with that of the underside of the leaves of the foodplant; the larve pupate towards the end of the month in a nearly perpendicular position amongst the stems of the Helianthemum, and slightly attached thereto by a few silk threads near the ground. The larve of Strymon (Thecla) w-album are sometimes to be obtained in numbers at the end of May, and in early June, by beating elms. Searching is recommended, and an interesting account of this mode of capture is given in H’nt. Rec., x., p. 187. In early May the larve of Melitaea athalia are to be found on Melampyrum pratense, Plantago major, and P. lanceolata, the first-named foodplant being apparently preferred. The larve of Limenitis sibylla are more readily found in May than in April. Look out for freshly-eaten leaves, and then search the stem round, being careful not to overlook the trailing branches. They want close work, being only about half-an-inch in length at the com- mencement, but almost fullfed at the end, of the month. Beat sallows through this month for larve of Apatura iris—sallows that stand high and dry in the middle of a marsh even furnish larve —for the ? wanders very far in search of sallows, and you never know on what stunted little bush may be feeding the horned head that is so dear a prize. In May search carefully the terminal buds of a buckthorn bush, Rhamnus frangula or R. catharticus, for the larve of Gonepterya rhamni ; several may often be found on one bush. In late May and throughout June, the flowering stems of Cardamine pratensis and Sisymbrium officinale should be searched for the young larve of Euchloé cardamines: they are very like the seed-pods of their foodplants. In confinement, the larvz of Pieris napi will feed well on horse-radish. They also eat Nasturtium officinale, Barbarea vulgaris, ete. Towards the end of the month, plants of Vicia cracca and Lathyrus tuberosus, growing by hedges or on the borders of woods, should be searched for the young larve of Leptosia sinapis. JunE.—The young larve of Nisoniades tages are to be found, at the end of June and throughout July, in little hollows, formed by drawing together three leaflets of Lotus corniculatus ; the two outer ones being drawn close together, and the third one bent over like a curved roof; the structure looks almost exactly like a leaf not quite expanded. In early June the larvee of Adopaea flava (thaumas) are to be swept 44 BRITISH BUTTERFLIES. from the soft grass, Holcus lanatus, with the colour of which their tints assimilate remarkably well; they may also be swept from Brachypodium sylvaticum. The larva of this species is often found in one’s net, when one is working for micro-lepidoptera among the long grass at dusk, in open places in woods, etc. When still small, the larve of Cyclopides palaemon make tubular homes in the leaves of Brachypodium sylvaticum, leaving an opening at each end, whence they emerge to feed on those parts of the plant near their domiciles. In mid-June the young larve of Polyommatus icarus feed on the leaves of Lotus corniculatus, eating into the substance of a leaf either from the upper- or underside, leaving the opposite skin as a white spot, although they sometimes eat the flowers, the petals of which they devour entirely. The larve of Agriades (Polyommatus) corydonare to be found on Hippo- crepis comosa through June. The larva of this species can only be distin- suished from that of Agriades bellargus by its having the ground colour ofa lighter, brighter green (a green with more yellow in it), and the hairs light brown, whilst that of P. bellargus has the ground colour deeper - green, with the hairs or bristles black. The larvee of Aricia (Polyommatus) astrarche are to be found in late June and July, on the underside of the leaves of Helianthemum vulgare. The feeding of the smaller larve makes small spots on the upper, dark ereen, surface of the leaves, the spots becoming larger and browner, until, at last, almost the whole undersurface of the leaves is entirely eaten, although, with an indefinite supply of food, they rarely remain long enough on one leaf to more than blotch it very markedly, before moving to another. At the end of June and the beginning of July, search the underside of holly leaves for larvee of Celastrina (Cyaniris) argiolus. The leaves affected have the appearance of being mined. At the end of June and in early July, the larve of Callophrys rubi can be beaten from broom, Genista tinctoria, and many other plants ; bramble, after which the species was named, appears to be very rarely chosen. The larvee of Ruralis (Zephyrus) betulae always sit on the underside of a leaf of blackthorn, along the midrib, and are most difficult to see in this position. The blackthorn bushes are, therefore, better beaten than searched, for larvee of Ruralis betulae; stunted ones are often the more prolific. An umbrella is better than a tray for this purpose, as it can be fitted into the structural irreeularities of the blackthorn bushes more successfully. The low branches of oak, with their growth of foliage, on isolated trees, often prove the best, when one is working for larve of Bithys (Zephyrus) quercus. Search the tray carefully, as the half-grown examples imitate the fallen bud-sheaths exactly in colour. During the first week of June, beat low elm-trees on the outskirts of woods, or on the borders of rides of woods. Large numbers of larvee of Strymon (Thecla) w-album may sometimes thus be obtained. In early June, the eggs of Hamearis (Nemeobius) lucina can be found fairly readily, in the localities where the species occurs, on the under- side of primrose (more rarely cowslip) leaves. The young larve eat COLLECTING BUTTERFLY LARVZ. 45 little holes in the leaves, but later they devour large pieces of the leaves, and their whereabouts become conspicuous. The young larve of Brenthis selene usually divide into two sections in this country, one very small part feeding up rapidly and producing a few imagines in August, the others hybernating when about 10mm. long, and going through the winter in this stage. The fullfed larva of Argynnis aglaia is to be found on Viola canina in June; it is difficult to discover, and is best obtained when feeding, as its movements are rapid and may attract attention ; when not feeding, it usually hides below the leaves of the plant which it has been eating. The fullfed larve of Dryas paphia are to be found in early June, feeding on the leaves of Viola canina, freely exposing themselves, according to Buckler, on the violet plants. The larve of Pyrameis cardui are, in years when an immigration has taken place in May or early June, most abundant, in their little globular homes of spun-together thistle leaves, or other of their food- plants, in late June and early July. The gregarious larve of Vanessa io are to be found in considerable companies in late June and early July, spread out over beds of stinging- nettles by roadsides, behind hedges, or sunny corners on the edges of woods. The gregarious larve of Hugonia polychloros should be sought on elm, willow, sallow, aspen, etc. ; the eggs are laid in spring by hyber- nated 9s, and the presence of hybernators in April and early May in a locality should lead to a search for larve in June. The fullfed larve of Hipparchia semele require light soil, peat, or similar material, in which to burrow. ‘They hide therein by day, feeding by night, and, when mature, form their puparia just beneath the surface of the ground in a manner altogether different from every other British butterfly. The fullfed larvee of Apatura iris are to be found in June on sallow; they eat rapidly, are easily alarmed, when they draw themselves in, and are difficult to detect on the leaf on which they are. They are usually obtained by beating. In late June and July, on Wicken Fen, stand over a plant of Peucedanum palustre, and look most carefully, if you wish to see the little black larve of Papilio machaon. In confinement, the larve of Papilio machaon will feed very freely on the leaves af garden carrot, on Angelica sylvestris, and other umbellifers. The larve of Colias edusa, obtained from eggs laid by immigrant 2s in June, will feed up well in confinement on Trifolium repens and Lotus corniculatus, pupating in July, the imagines emerging in August. In late June and early July, the larve of Gonepteryx rhamni are to be found on Rhamnus frangula and R. catharticus; stunted bushes in sheltered nooks on the outskirts of a wood are usually good localities for them. They are sometimes very abundant; and it will add to one’s success 1f one places oneself so that the sun falls across the Rhamnus leaf, showing the shadowed side of the larva, when it is at once discovered ; otherwise it so exactly resembles the midrib along the centre of the leaf (where it rests) that it will easily escape notice. 46 . oe BRITISH BUTTERFLIES. The larvee should be searched for early (especially in a recognised early season) as many of them appear to wander away to pupate. In late June, carefully search the seedpods of Cardamine pratensis, Sisymbrium officinale, garden rocket, etc., for larve of Huchloé cardamines. Particularly examine those where the growth of the seedpod seems irregular, which will be owing to the feeding of the larvee, and the latter will be found closely imitating the growth there. Immigrant 9s of Pontia daplidice lay their eggs occasionally, in June, on Reseda luteola, on which the larve feed in July, producing imagines in August or September. [The species is quite unable to winter in our climate. | In June (and August) the larve of Pieris napi may sometimes be found in numbers feeding on Nasturtium officinale and Barbarea vulgaris. They may also be found on Hesperis matronalis, ete. They grow very rapidly, and are fullfed in early July, pupating during that month, and emerging towards the end of July or in early August. Tome. —In July, the larve of Nisontades tages, no longer able to hide within the little caves formed of the leaflets of Lotus corniculatus, which they use when young, make longer ones, but their feeding soon exposes their bodies partly to view. They repeatedly change. their habitations, always, however, by night, and are most retired in their habits. They are fullfed at the end of the month, when they spin silken hybernacula, in which they remain invisible, not pupating until the following April or May. The young larvee of Hesperia malvae are to be obtained, in July, on Potentilla fragariastrum, P. reptans and. Rubus fruticosus; the larve that are on bramble seem to be found chiefly on stunted bushes with small leaves; the large juicy leaves of strong bushes apparently offer no temptation to the female. The larvee of H. malvae appear to choose the upper side of a leaf on which to rest, and, stretched along the midrib, spin several silken threads overhead for a covering, feeding therein by eating away the upper part of the leaf. When a larva has cleared this, and made a blotch of considerable extent, it repeats the work on another leaf. The larger larvee pull down a second leaf over the first, fastening the edges with silk, and these form a hollow in which they live, coming out therefrom occasionally to feed on the surrounding leaves. In July, the young larve of Augiades sylvanus feed on cock’s-foot- grass, couch-grass, etc., resting in the middle of a blade and fastening its edges across with five or six distinct little ropes of white silk. The young larvee of Adopaea flava (thaumas) leave the eggs in late July or early August, and spin little silken ropes across the blades of erass; but, although they feed until November before hybernation, they are not then more than about 2mm.-3mm. in length, almost the whole of the growth being done in the spring (Hellins). In the early part of July, collect the flower-heads of Anthyllis vulneraria for larvee of Cupido minima. They eat little holes through the calyx and corolla so as to get into the flowers, when they feed on the immature seed-vessels, leaving one floweret when cleared and entering another. As they get older, their bodies cannot be wholly contained in the corolla, and they may be then seen with the fronts of their bodies thrust into the flowers, the hinder parts hanging out, but COLLECTING BUTTERFLY LARV. 47 still difficult to distinguish among the dense inflorescence of the flower-head. In confinement the larvee of Cupido minima are fullfed before the end of July; they then take up a position as if for pupation, but remain quite still and immovable until the following May, when pupation takes place. In July, the little larvee of Polyomimatus icarus make small, pale, transparent blotches, on the leaflets of Lotus corniculatus, Ornithopus perpusillus, etc., the paleness being due to the eating away of the soft part of the leaf, and leaving only the transparent skin. In late July the now nearly fullfed larvee may be more easily found. When fullfed the larva of Argynnis aglaia will spin together several of the large leaves of its foodplant into a hollow, tent-like enclosure, and, in this, suspend itself before changing into a chrysalis. The larve of Dryas paphia leave the egg towards the end of July, the ege-stage only lasting about a fortnight, the larve feeding very little (or not at all) on Viola canina, before hybernation, being only about 3mm. long in spring (March) when they commence to feed. The larve of Pyrameis atalanta are to be found in July and August in little chambers, formed by drawing together the leaves of Urtica dioica and Partetaria officinalis; they generally hang up and pupate within these larval chambers. In July, the larvee of Pyramets cardui fasten together the leaves of Onopordon acanthium and other thistles, with a few tough silken threads, eating out the thick fleshy parts of the enclosed leaves. They generally hang up and pupate within these larval chambers. (Larvee also feed on Hehiwm vulgare, Malva, etc.) The young larve of Apatura iris are not difficult to rear in leno- sleeves on a healthy sallow-bush; the eggs are laid from about July 20th-August 10th, and the egg-stage lasts only about eight days. The young larve of Melampias epiphron are not easy to rear in confinement; they feed on Aira praecow and A. caespitosa, growing to the length of about 4in. before winter; they then hybernate until the end of February, when their food-plant should be attended to. The larve of Coenonympha tiphon may be reared in confinement on the beaked rush, Rhynchospora alba; a plant of this should be care- fully potted, and the young larve may then be left without much attention ; they go on feeding, as a rule, until the hybernating stage, with little trouble, provided that care be taken not to allow the active little fellows to escape. The larve of Huchloé cardamines are to be found, in July, on many cruciferous plants, of which WHesperis matronalis, Sinapis arvensis, Cardamine pratensis, Sisymbrium officinale, Alliaria officinalis, Turritis glabra, and, in gardens, garden-rocket and horse-radish appear to be the most frequently selected. The larve from the eggs laid by the summer @s of Leptosia stnapis will feed up on Vicia cracca or Orobus tuberosus, and are generally fullfed in early September. In July, the young larvee of Papilio machaon may be searched for with every prospect of success, the black larva, with its white saddle, being very easily found when once the eye of the searcher is in. Until then, it is most difficult to detect, although many may be on the 48 BRITISH BUTTERFLIES. plant under examination. The habit of repose, with the neck arched something like the larva of a Sphingid, is very striking. Aueust.—In August, the larve of Cyclopides palaemon feed within long cylindrical tubes made of the leaves of Brachypodium sylvaticum, quickly, however, eating out their domiciles and forming fresh ones; they first eat the lower part of the leaf below the tube, all but the midrib, then devour the top of the leaf above the tubular part, and, lastly, the tube itself, until, by degrees, it becomes too short to shelter them, when they desert it and cut through the midrib, causing the tubular remains to fall away, after which they select a fresh leaf for the construction of another tube, as above. The young larve of Celastrina argiolus are to be found feeding on the tender young ivy leaves and flowers, and pupate in early September. The eggs are usually laid in early August, beneath the flower-heads of the umbels of ivy. The newly-hatched larve of Colias hyale feed up slowly from about mid-August, until about October or November, on Trifolium repens, Medicago lupulina, M. sativa, etc. They then hybernate till March, and will usually pass this period successfully, provided that they are not exposed to a really low temperature, are kept quite clear of decaying leaves, and have a perfectly dry spot to rest on. They must be supplied with fresh food very early in the spring, and should be given as much sun and air as possible, but not exposed now to a low temperature. The larve of Colias edusa, obtained from eggs laid by ? s captured in August and September, will feed up well in confinement on Trifolium repens and Lotus corniculatus. They will try to feed up the same year, and must be carefully nurtured. They might, indoors, be induced to partially hybernate until early March, but then would have to be kept perfectly free from damp, and well away from any decaying leaves of their foodplant. The larve of H'pinephele tithonus appear after the eges have been laid about three weeks, and feed well on Poa annua, Dactylis glomerata, and other common grasses; they hybernate when exceedingly small, but nibble in winter when the weather is mild, not feeding very much, however, till mid-March. The young larve of Melanargia yalathea leave the eggs in August, and feed well, in confinement, on almost all the common garden erasses—Dactylis ylomerata has been noted as a specially favoured one. The larvee hybernate from about the end of October, feeding occasionally when the weather is mild, going ahead more rapidly in March and April, and being fullfed in June. The larvee of Hipparchia semele can be reared in confinement on Triticum repens, Aira praecov, and many other grasses; they are very sluggish, hide low down among the foodplant, nibble slowly most of the winter, feed only at night, and often bore under the ground, if at all suitable, by day. They are fullfed about mid-June, when pupation takes place. The fullfed larve of Papilio machaon are to be found in August, usually resting in an almost vertical position on a stem of the food- plant, or on a plant near; although such a large conspicuous caterpillar, when separated from its food, it is not at all easy to see when COLLECTING BUTTERFLY LARVZ. 49 surrounded by the herbage of the fen districts to which it is almost absolutely restricted in Britain. SEPTEMBER.—The young larve of Polyommatus icarus make litile, pale, transparent blotches on the leaflets of Lotus corniculatus, Ornithopus perpusillus, etc., the paleness being due to the eating away of the soft parts of the leaf, and leaving only the transparent skin. Still more common in July. Sleeve larve of Apatura iris out on sallow, so that they can rest on a thick branch ; they must be removed from the sleeves every day, till they settle down on a twig, as none ever hybernate successfully if left on the sleeve. _ Larve of Pyrameis cardui and P. atalanta, found in September and October, will pupate and emerge the same year; they must, therefore, be kept under artificial conditions, and care taken of them, both as regards temperature and food, to ensure success. The species will not hybernate as larvee or pup under any conditions. The larve of Pararge egeria can be reared, in confinement, on Dactylis glomerata, etc. ; they appear to nibble throughout the winter, and to pupate as soon as there is any mild weather in the spring, often at the commencement of April (sometimes this species passes the winter as pupa). The young larve of Colias hyale (obtained from eggs laid by confining 2s on clover plants, exposed to the sun), feed slowly the end of October, when they become dormant and hybernate. In confinement, they do this best by removal from the foodplant, and by being placed in a chip-box covered with muslin ; they must be protected from frost, and kept at a temperature of about 40°F.-45°F. By the middle of February, the larve are again on the move, and should then be placed on growing plants of clover, with plenty of young leaves, when they will commence to feed again. They feed on slowly through March and April, pupate in May, and the imagines emerge in about a month. Ocroser.—In October, the larva of Cyclopides palaemon draws a leaf of Brachypodium sylvaticum into tubular form around itself, lining the inside carefully with white silk, and thus forms the hybernaculum, in which it spends the winter. Tn late October and November, the hybernating larve of Augiades sylvanus, about 12mm. long, are to be found in their long, silken, narrow, tough, close-fitting hybernacula, formed by spinning together the edges of the green blades, the opaque webs being not much bigger than the larvee; in confinement, riband-grass forms a useful substitute for the finer grasses; the larve commence feeding again in March, and are fullfed about the end of May. Small larve of Polyommatus bellargus are to be found in October (and July) on the underside of the leaves of Hippocrepis comosa, eating out the undersurface for a small space, but leaving the upper skin untouched, which then turns white; these little white dots or spots show, therefore, where the larve are at work; they feed slowly through the winter, and the blotches are much larger by early February. In March, the leaflets are eaten from the edge, and often demolished entirely. The hybernaculum, in which the larva of Limenitis sibylla passes the winter, may be placed three or four buds down from the tip of the 50 BRITISH BUTTERFLIES. twig, shooting out from the main stalk of a large honeysuckle-bine ; it is made of a honeysuckle leaf, which has been first partly bitten through near its axil, and then securely fixed by its two edges, for about half its length, to the twig from which it grows, and across which its edges are firmly bound together with a spinning of strong silk; just at the point where the leaf meets the underside of the twig, there is a circular aperture, apparently designed for the egress of the larva in spring ; as the leaf withers the hybernaculum becomes puckered, and little more than half-an-inch in length, and has the appearance of a small shrivelled leaf, clinging to the dry stem, and would thus easily escape ordinary observation. The larvee of Hrebia aethiops commence to hybernate in October, when exceedingly small, hiding in the thickest parts of the tufts of grass with which they may be supplied. They commence to feed again in early spring, as soon as the grass commences to grow, and are fullfed in May and June. The larve of Hipparchia semele hybernate small, remaining on the erass all the winter, and show no tendency to burrow or hide; they feed a little all the winter in suitable weather, but do not grow perceptibly till the spring. The larvee of Hnodia hyperanthus, E'pinephele ianira, Coenonympha pamphilus, etc., require considerable attention in confinement during the winter; they appear to nibble occasionally, during mild weather, from November to March, growing very slowly, or not at all, but making good progress later. Hnodia hyperanthus, in particular, appears to hide as much as possible from the daylight during the hybernating period. These ‘‘hints,’’ as to obtaining eges of butterflies, are extracted from our work Practical Hints for the Field Lepidopterist, and are inserted as illustrations of the various details to which the attention of the seeker for, and breeder of, the larve of butterflies must be directed. CHAPTER XII. THE SILK-SPINNING HABIT IN BUTTERFLY LARV. The silk-spinning habit is common to almost all lepidopterous larve. The larval spinneret appears to be homologous with the hypopharynx of insects of other orders. The homology of the different parts, Packard says, is apparently identical, the common duct of the silk-elands or sericteries opening at the end of the hypopharynx, which here forms a complete tube, or proboscis, extending beyond the end of the labium, and being then modified into the spinneret in adaptation to its use as a spinning organ. The silk thread which issues from the spinneret was discovered by Leeuwenhoeck to consist of a double ribbon- like band, due to the silk-glands each forming a cylindrical thread of silk, surrounded by a layer of gum; the two threads having passed into the common duct receive the secretion of Filippi’s gland, where the silken fluid is formed, and, passing into the common canal, enter the orifice of the spinning canal, almost completely divided into two by the sharp THE SILK-SPINNING HABIT IN BUTTERFLY LARV. 51 edge of the rachis. The threads each pass into one of the two grooves surrounded by gum, and are pressed by the powerful contractions of the muscles of the plates, so that each is compelled to mould itself in the groove it occupies, and to take its shape. The spinneret thus compresses the thread and diminishes its diameter, whilst the constant compression of the thread as it passes through the press keeps it in a certain state of tension so as to allow the caterpillar, while spinning, to firmly hold its thread ; the press does not act directly on the silken thread, but through the gummy layer which transmits over the whole surface of the silken fluid the pressure exerted on it. After having overcome this difficult passage, the silk thread has acquired its definite form and passes out of the spinneret. [Full details of the structure of the spinning organs, the formation of silk, the drawing of the thread from the spinneret, etc., are to be obtained from Packard’s Textbook of Entomology, pp. 389 et seq.| These preliminary details will perhaps enable the young lepidopterist to understand some of the observations he is certain to make when watching the larve of butter- flies, for the silk-spinning habit is to be observed in a greater or less degree in all butterfly larve. It is remarkable how variable in degree is the silk-spinning habit developed in the larve of different butterflies. Usually, but not at all necessarily, the larve of allied butterflies have a somewhat similar habit in this respect, e¢.g., the larve of most Ruralids, and many Pierids, spin very little silk, whilst those of others spin much. The larve of certain Vanessids, Melitzids, etc., spin threads wherever they walk, being apparently unable to move about in their early stages, at least, without so spinning. All larve, however, whatever be their general habit, spin a silken web on which to fix themselves during the period of moulting, the old larval skin being attached thereto before the larva in its new dress withdraws itself therefrom. Some larve construct nests in common, and live gregariously, especially when young, others fold up one or more leaves with silk, and live in the hollow thus constructed singly; others live quite exposed, resting on a leaf, stem, or other part of the foodplant, and, at the most, spinning a few silken threads as a pad on which to stand. As a rule, butterfly larve live an exposed life. Of those that hide, however, in the fashion of so many other lepidopterous larve, by making a tent of one or more leaves in which to secrete them- selves, the Urbicolid larve are the most remarkable. All our British species do this, as will be seen by reference to our notes on the “habits of the larve’’ (posted, pp. 98, 108, 120, etc.), and the habit is common to the superfamily. There is no need to redescribe the habits of our British species in this respect, but we may note that Scudder, speaking of the American species, says that ‘“‘the ‘skipper’ larve form a nest of a single leaf, in early life, folding over a little piece of leaf, and fastening the edge to the opposite surface by a few loose strands of silk; to effect this, they first bite a little channel into the leaf, at just such a place as to leave a fragment of leaf, neither too large, nor too small, to serve as a roof when they shall have turned it over; often they have to cut two channels in order to procure a flap sufficiently small for their purpose; and it is curious to watch one of these tender creatures, just as soon as it has devoured its eggshell, struggling with a tough oak-leaf, to build 52, BRITISH BUTTERFLIES. for itself a house. These nests are usually very firmly made, the silken fastenings being composed of many strands, often very tough. On leaving one nest to construct a larger one, the caterpillar always appears to first bite off the threads of the old nest, and thus give the flap a chance to resume its position, which, however, it rarely fully does. When older, many of these same ‘skipper’ larve find a single leaf of their foodplant too small to conceal them, and so they draw several leaves together, just as they grow upon the plant, and, retaining them in the desired place by silken bands, the larve live within the silken bower. This mode of construction is adopted almost from the first by the Pamphilids, which feed on grasses, the proximity of adjoining blades near the base affording a good chance to attach them together, while a cluster of blades furnishes a similar chance to construct the somewhat tubular nest they require when they have grown large and fat.’ As to the value of these homes, Nicéyille observes (Butts. of Sumatra, p. 894) that the larva of a large ‘‘ skipper” butterfly, Hidari trava, and that of a Nymphalid, Amathusia phidippus, live, at the same time, on the leaves of Cocos nucifera, and he remarks that, owing to their general abundance, the two species often have a severe struggle to live together, in which the more robust Hesperiid, which secures a shelter for itself by spinning the leaves together, is generally victorious. Even among the Papilios, the larve do not disdain to use this mode of protection, and whilst that of Jasoniades glaucus merely spins silk on the surface of the leaf on which it rests, so that the edges of the leaf curl up and conceal its sides, the larva of Huphoeades troilus spins the leaf completely over, so that the opposite edges touch, and itself thus becomes quite hidden. The purpose of gregarious nests appears to be twofold. In one case, ~ e.g., that represented by Aporia crataegi, the nest is very definitely intended for the purpose of concealment; in the other case, e.g., that represented by Euvanessa antiopa and Aglais urticae, the silken web appears, especially after the larvee have reached the second stadium, to be merely a means of keeping up a connection between the various parts of the eregarious company, and to be little used for the purpose of hiding. Scudder, referring to the larve of Huvanessa antiopa, notes that ‘they move about from place to place, spinning wherever they go, so that, at last, the line of movement, by successive strands thrown across every angle that a twig makes with the larger stem, forms a sort of veil of silk over which they crawl with extreme rapidity, but without which their movements are greatly retarded.’”’ Although our single British Apaturid species has a solitary larva, those of one of the American species, Chlorippe clyton, are gregarious in their first three stadia, and use their web much in the same manner as that just described as usual for Huvanessa antiopa. They feed side by side in rows, eating the leaves from the tips backward, but leaving the stouter ribs; they form a pathway of silk wherever they go, but make no special structures for concealment. The larve of the allied C. celtis lives solitarily, but lines the upper surface of a leaf of Celtis with silk in such a manner as to cause the sides to curl slightly upward, and thus partially conceal it from view. Similarly, the larva of Anaca andrica lines the upper surface of Croton with silk, bringing the upper edges together without fastenings, and thus makes a nest like that of Euphoeades (troilus), within which it lies concealed, eating the base of THE SILK-SPINNING HABIT IN BUTTERFLY LARVZ. 538 the leaf; when this becomes too small, it makes a similar nest from another leaf, but goes outside to feed on neighbouring leaves, generally towards evening. The tiny larve of Aporia crataegi, spin a web over two or three leaves of their foodplant, within which they hide, coming out only to feed, and extending their web as they increase in size. A specially tough retreat is built for a hybernaculum, and, in this, a whole community winters in safety. Our other Pierid larve spin threads of silk on which to walk, but, otherwise, none of the allies of A. crataegi, in Europe, spin similar webs. Scudder, however, notes that a Mexican Pierid constructs a web nearly as close as parchment. Belonging to an entirely different group of butterflies, but having almost exactly the same gregarious habit as Aporia crataegi, the larvee of Melitaea aurinia spin a silken tent over the young leaves of scabious, in which they live, feeding only on the undersides of the leaves; they feed very slowly, and are still exceedingly small when they make their hybernaculum, after leaving which, in the spring, they appear to live singly and fully exposed, making, however, a silken cocoon by drawing together several culms of grass, when fullfed, in which they pupate. The larve of M. aurinia are said to leave their hybernacula very regularly, about March 1st in Co. Cork, in Ireland, although, in the early season of 1893, they had already donesoin mid-February. The larve of M. cinxia similarly pass their early lives and the winter gregariously in a tent formed by a compact web, leaving its shelter in the spring for another slighter structure. The hybernating web is larger than that under which they feed, and is woven of silk, with grass and plantain stems inter- mixed, and is well-roofed, so that the inner grass is quitedry. Luff says that, in Guernsey, he had noticed the very young larve of M. cinwia on the webs, spun on their foodplant, in August and September, but that, when he searched for their hybernacula in December and January, in the same spot, he could not find them until he came accidentally across a winter-nest whilst searching for beetles; this was in the centre of a tuft of grass, close to the roots; 1t was pear-shaped, and, with the larve of M. cinaia were a number of larve of Anthrocera trifolit, hybernating with them. Luff says that they spin another web in spring on their foodplant, but this isless compact than the winter-nest, although larger. Most of the larvee leave the nest and live singly, when nearly fullgrown, others, however, live more or less gregariously, even up to the time of pupation. This mode of life is evidently not confined to the Melitzids of the Old World, ‘‘for Scudder notes a similar habit in some of the Meliteids (Cinclidia, Huphydryas, etc.) of North America, which, living in company, cover at first a few leaves, then the whole head of the plant, and eventually, sometimes, the whole plant, in a tolerably firm web, within which the company feed, until the whole becomes a nasty mess of half-eaten and dying leaves, and all sorts of frass, including their own excrement and cast-off pellicles, everywhere tangled with web. Within such a nest they hybernate, but not until they have strengthened it with denser web, and drawn the leaves of the head more tightly, so that it becomes a mere bunch which one may cover with his hand, and which contracts the more apparently as winter approaches. In the spring, they evidently have had enough of this sort of communal life, and live, thereafter, in the open air.”” He further notes that, whilst the larve of Huphydryas phaeton hybernate 54 BRITISH BUTTERFLIES. as thus described within a web on Chelone glabra, those of Cinclidia harristi, on Aster umbellatus, live similarly until the time for hybernation, when they desert the nest and conceal themselves in crevices, where they pass the winter. The gregarious character exhibited by such Meliteid larve as those just mentioned, is almost paralleled by that of certain Vanessid larvee to which, already, reference has been made. But the less gregarious species among the Vanessids also spin hiding-places, usually several on one plant, but only one larva in each little nest. Of these, the spun-together thistle-leaf (or leaves) that often forms the hiding-place of the larva of Pyrameis cardui, and the spun-together nettle-leaves that similarly form the home of the larva of Pyrameis atalanta, will be known to everyone. Scudder, writing of these, says that-‘‘ the most common form of nest is that in which different parts of the same leaf, or adjacent parts of different leaves, are fastened together by silken strands. The simplest and weakest of these are made by the cater- pillars of Polygonia faunus and Vanessa (Pyrameis) atalanta, which fasten together very weakly the opposite edges of a single large leaf, so as just to make them meet; but the threads are so slight that they are ruptured with the slightest effort. The caterpillar within, having thus secured a shelter, seems loth to leave it, and makes its meals from its own dwelling, until, having literally eaten itself out of house and home, it is forced to venture forth and construct another. When, however, V’. atalanta is more than halfgrown, it finds it easier to attach neighbouring leaves of the thickly-growing nettle, than to find one sufficiently free to use it only, so that fully one-half of the nests of the larger caterpillars are made from a number of leaves; the nest is always roomy, capable of housing several caterpillars, though never containing more than one. ‘The nesting-habits of V. atalanta are shared by the other species of Vanessa with certain slight variations. In early life, V. cardui tries to make the stiff and crenulated edges of thistle-leaves meet together, but with indifferent success, and so fills in the interstices with an exceedingly thin web, in no way concealing it from sight. In after life it forms an oval nest of the size of a pigeon’s ego, by fastening adjoining leaves together very slightly, and filling all the interstices with a similar flimsy web, upon which it fastens, or into which it weaves, bits of eaten leaf, or parts of the inflorescence of the plant, still imperfectly concealing it from sight; and, sometimes, it hangs itself up for chrysalis within the same narrow, and by this time very filthy, apartment. V. hwntera makes a similar, but rounder, nest on the Gnaphalium, and conceals itself very effectually by com- pletely covering the more compact, but still very slight, web, with the inflorescence of the plant. At first this is merely composed of the silky hairs of the foodplant, mixed with much silk, forming a dense white mat, beneath which they devour the parenchyma, then enlarge the nest, never leaving it for food, but enclosing larger and larger areas, until, finally, many leaves are drawn together, the bitten-off inflorescence of the Gnaphalium interwoven with the web, and a nest formed as large as a pigeon’s egg, only in the last few days of their life do the larve leave the nest and devour the entire leaf.” The hybernating period is a serious one for those silk-spinning larve that hybernate as larvee. Limenitis sibylla for its hybernaculum, spins a honeysuckle leaf to a twig of the plant, which it securely fixes, THE SILK-SPINNING HABIT IN BUTTERFLY LARVA. 55 and prevents from falling, by silk spun over its extremity and the twig on which it grows. The leaf is then spun carefully together and _ becomes crumpled as it dries during the winter. Into this the larva crawls, and, in this, it hides during the period of hybernation. So firmly is the hybernaculum fixed to the twig, that it can have no independent movement, and it resembles so exactly a dead leaf clinging to the stem, that it is sure to escape observation. When Apatura iris spins its hybernaculum, it not only covers the twig with a silken pad, to which it may firmly cling, but also envelopes the hinder part of its body in a silken covering. The hybernaculum of the larva of Basilarchia archippus appears to be as interesting. Scudder says that the larva hybernates when partly grown, and provides for the occasion a winter residence, which is occupied only during the cold season. For this purpose, it eats the side of a willow-leaf nearly to the midrib, for about one-third the distance from the tip, ordinarily selecting for the purpose, a leaf near the end of a twig; it brings together the opposite edges of the leaf, and not only fastens them firmly with silk, but covers this nest outside and inside with a carpet of light-brown, glossy silk, so that the leaf is nearly hidden, nor is this all, it travels back and forth on the leafstalk and around the twig, spinning its silk as it goes, until the leaf is firmly attached to the stalk, and, in spite of the frost and wind, it will easily hang until spring. Following the projecting midrib, the caterpillar creeps into this dark cell, head foremost, and closes the opening with its hinder segments, all abristle with spines and warts. The other species of the same genus, B. arthemis and B. astyanax have similar habits, the former feeds on birch, and, if one examines these trees in early spring, one can hardly fail to be struck by the deceptive resemblance that these hybernacula bear to the opening buds and curving terminal shoots of the very twigs on which they occur; the colour of the soft down of the buds, and the enveloping silk of the hybernaculaareassimilarasare their forms, and this mimetic resemblance is doubtless as effective as it is interesting. The larva of B.arthemis spins its hybernaculum when about half-grown, and, selecting a growing leaf of birch, it eats away the apical third or fourth, excepting the midrib and a narrow flange on each side of it, or it uses the leaf it has been eating, already trimmed in this fashion; it then draws together, above, the outer edges of the uneaten portion to construct a tube, which it lines very heavily with brown silk, within and without, and further binds the leafstalk to the stem with repeated bindings of silk to prevent its falling to the ground in winter; by means of the ledge formed by the projecting midrib, it then enters the tube head foremost, and completely fills it, so that the opening is just closed by the roughened end of the body. Although nothing to do with the silk-spimning of the hybernaculum, we may here mention the peculiar habit of Basilarchia arthemis, of retiring, after a meal (made of a birch leaf), to the stripped midrib to rest, fastening to it, however, minute bits of leaf with an abundance of silk in order to strengthen it, whilst Chapman describes the much more complicated silk-covered platform, made by the larva of Charazes jasius, and which, in the Ksterel, it spins on the south side of an Arbutus tree, low enough down to have the upper part of the tree as a protection and shelter. The larva either clothes the surface of a leat with silk, or fastens together several leaves, which it then similarly covers with 56 BRITISH BUTTERFLIES. silk, and on which it obtains a firm foothold, resting thereon the ereater part of the day, basking in the sun, the leaves being usually at such a slope that they get an almost vertical exposure, whilst to feed they often prefer to go to a neighbouring spray, so as not to interfere with their carefully prepared resting-platform. Of the protective value of this particular resting-position, Chapman says (Hnt. Rec., ix., p. 1938): “The larva at rest, seen from whatever direction, exactly imitates some aspect of leaves or buds under the different effects of light and shade, and it is thus possible for an untrained eye, in many instances, to look at it, and for it, some time before seeing it. The yellow lateral line resembles the midrib of the leaf seen from above or below, accord- ing to light; the colour and apparent texture of the skin are the same as those of many leaves. The extraordinary head, with its coloured jaws and spines, suggests, in many aspects, the little group of buds at the extremity of the branches. One has often to look a second time at certain leaves and branches, as well as at the buds, to be sure that they are parts of the tree and notalarva. The curiously-coloured circles on the back of the abdominal segments 3 and 5, which are more brilliant with their blue and yellow than anything in an Arbutus leaf, neverthe- less produce exactly the effect of certain little rings of fungus, or decay, that are very common on the leaves.’’ But the larva of Bastlarchia arthemis is said, by Scudder, to further use its silk-spinning habits for protective purposes in a most remarkable way, for he says that the young larva makes a loose ball about the size of a small pea, out of bitten scraps of leaf held together by strands of silk, and this it attaches by a thread to the stripped midrib, on which it is resting, as described above, so that it is moved by every breath of wind, a device, perhaps, to distract from itself the attention of an enemy, for, by constant removals, it is always kept close to the eaten edge of the leaf, while the posterior of the larva is as far out on the stripped midrib as it can find a good footing; after the second moult it no longer makes this remarkable little packet. Although less usual among butterfly larve, some species use the means of escape that is so frequent among the larve of certain groups of moths, viz., when disturbed, of allowing themselves to drop rapidly from their position of rest by a silken thread attached to a leaf. Such. among our British species are Chrysophanus dispar, Hesperia malvae, ete., and Scudder observes that, in America, Strymon titus and Hypatus bachmaniti have the same habit. It is certainly more frequently observed among young, than in older, larve. The final stage usually brings out the silk-spinning possibilities of the larva to their greatest extent. For the purpose of pupation, almost all butterfly larve, however little silk-spinning they do during their larval life, do some spinning at this _ period. Few, except the Urbicolids and Parnassiids, spin silken cocoons, i.é., Silken webs in which to pupate, but almost all spin at least a thick silken pad to which the pupa is attached by its anal cremastral hooks, and many spin, in addition, a_ silken eirth, or support, round the body, which acts as a girdle when pupation takes place. There are, however, many intermediate stages between the coarse, but slight, silken cocoon of the Parnassiids, in which the pupa hes loosely, and the merely suspended butterfly pupa, whose larva has spun a silken pad before pupation, and from which it THE SILK-SPINNING HABIT IN BUTTERFLY LARVA. 57 hangs freely in order to change to the pupal state. Some of the most interesting of the puparia spun by butterfly larve are those of certain _Vanessids, Argynnids, and Meliteids. Among the former, one may note the large open umbrella-like puparium of Pyrameis atalanta, formed by the larva spinning nettle-leaves together, yet quite open beneath, in order to facilitate the escape of the imago. There is also the often almost globular silken puparium spun by the larva of P. cardui, and a somewhat similar one spun low down, near the ground, by the larva of Argynnis aglaia. Many of the Melitzids, too, spin silken puparia, and the larve of Melitaea cinaia, when fullfed, do not always lose altogether the gregarious habits of their younger stages. We have noted (Ent. Rec., iv., p. 169) that, on one occasion, three larve of this species had spuna common silk tent, in which they had changed to pup. It is quite common to find a number of larve of this species spun up for pupation in close proximity to each other. Newman says, dozens of chrysalids are often suspended to the plantain almost close to the ground, in company. The larva of M. aurinia pupates alone, spinning a large quantity of loose, flossy silk, from which it suspends itself for pupation. We have already noted (anteda, p. 58) on the gregariousness of certain Pierid larvee, and the nests spun by Aporia crataegi, and the Mexican species, Huchetra socialis. A note by Anderson and Spry (Victorian Butterfites, pp. 86-37) on another Pierid, Delias harpalyce, whose larva feeds on a species of mistletoe (Loranthus pendulus), suggests that the web spun by the gregarious larve of this species is more or less utilised as a basis for the pupation pad. They write: ‘‘The larve, after the second moult, spin a silken footing for themselves wherever they go, and these spinnings, from so many larve feeding eregariously, form, eventually, quite a web-like habitation, which, no doubt, is of great service in enabling the caterpillars to maintain their footing during high winds or storms. When fullfed, they spin more than ever, and then, attaching themselves to the web, turn into pupe.’’ Floersheim, how- ever, particularly insists (Hint. Rec., xvil., pp. 310-311) that the larval habitations of Pyrameis atalanta are not, as often stated, used as puparia, but that the pupa, as noted above, is attached to the underside of a kind of umbrella, formed by drawing nettle-leaves together, and quite open beneath; this kind of puparium is what we know. We opened a great number at Val Tournanche in August, 1905, each containing a pupa. The belief that the silk-spinning habit of certain larve is some- times utilised in forming protective homes against adverse weather conditions—particularly of wind and rain—and this not only in the direction of making puparia, but also in earlier larval life, has often been expressed, and it is possible that this idea is well-founded, e.y., the silk-spinning habits of the larve of Hugonia polychloros are almost identical with those of Huvanessa antiopa, both tree-feeding species, and are apparently entirely different from those of Aglais urticae, Vanessa 10, etc., allied species feeding on low plants, the silken habitation made by the larve of the two first-named species being much more ex- tensive and permanent than in the case of the two last-named. Do these more extensive webs give a greater degree of safety on trees ? Chapman observes that “the young larve of Hugonia polychloros and Euvanessa antiopa cover their eggs, and the neighbourhood where they were deposited, with a silken web, not spun, as it were, of set 58 BRITISH BUTTERFLIES. purpose, but the result of journeying to the nearest leaves to feed and returning to the central position for resting. They appear often to feed in turns, one lot going out to feed whilst others have just returned to rest. As they get larger they move their headquarters, again, apparently, according to such exigencies as may occur, from the form of the branch they are on, to make another position more central to the available food, than to any instinct that makes them move at any particular stage or instar. Different broods seem to vary a good deal as to how far they remain gregarious in the last instar or become quite solitary. If food remains at hand, few larve wander far off until they do so for pupation, but they cease to go to and fro so much, and so, though still spinning silk to walk upon, do not increase the considerable webs spun during the earlier ‘stages. Hssentially, perhaps, Vanessa 10 and Aglais urticae do the same as Huvanessa antiopa and Hugonia polychloros, but the abundant web they spin in their earliest instars is but slightly added to (comparatively) in the intermediate stage, and, in the final instar, gregariousness seems to have ceased.