(Ir-^ BY THE SAME AUTHOR. Eoyal 8vo. In 4 parts, each, with 25 Coloured Plates, 15s., or complete in one vol. 63». CONTEIBUTIONS TO THE FLOEA OF MENTOKE, WINTER FLORA OF THE RIVIERA, Including the Coast from Marseilles to Genoa. BY J. TEAHEENE MOGGEIDGE, F.L.S. L. REEVE & CO., 5, HENRIETTA STREET, CO VENT GARDEN. HARVESTING ANTS AND TKAP-DOOE SPIDERS. HARVESTING ANTS AND TRAP-DOOR SPIDERS. NOTES AND OBSBEVATIONS ON THEIE paMts rab gfatllxtrgs* J. TRAHERNE MOGGRIDGE, F.L.S, LONDON : L. REEVE & CO., 5, HENRIETTA STREET, CO VENT GARDEN. 1873. LONDON : SAVILI., EDWARDS AND CO., PRINTERS, CHANDOS STREET, COVENT GARDEN. CONTENTS. PART 1. PAGE HARVESTING ANTS 1 PART 11. TEAP-DOOR SPIDERS 71 EXPLANATION OF PLATES. PART I.— HARYESTING ANTS. Plate I., p. 21, fig. A. — View of the entrance to a nest of Atta barbara, showing part of a train of ants bearing seeds, the conical mound of refuse thrown out, and some seedlings, which have sprung up from seeds accidentally dropped by the ants ; B, one of the larger workers of this species, of the natural size, andB 1, its abdomen and pedicle, with two nodes, magnified ; C*, one of the smaller workers, of the natural size ; C, a male, of the natural size ; D, a female, of the natural size ; D 1, wing of the same, magnified; I) 2, mouth organs of the same, magnified, with the mandibles removed, the two outer pieces being the maxilhie and their palpi, and the lozenge- shaped piece the labium, from the upper part of which the labial palpi spring, while behind the labium is the true tongue ; D 3, one of the mandibles, magnified ; E, a larva, of the natural size, and E 1, the same, magnified. Plate II., p. 22, fig. A. — A trowel containing earth, in which a granary full of seeds is lying almost undisturbed, of the natural size ; B, the crater-like entrances found at the mouths of the nests of Atta str actor, reduced to one-half the natural size. Plate III., p. 23. — The floors of three granaries of Atta barbara, surrounded by the much coarser gravelly earth, of the natural size. Plate IV., p. 31. — A mass of earth pierced by roots, in which the ants {Atta barbara) have made their gi-anaries and galleries. The galleries were full of seeds when first laid open. Of the natural size. Plate V., p. 33, fig. A. — Galleries and terminal cells of a nest of Atta barbara, excavated in the living sandstone rock, drawn in situ, of the natural size ; B, part of a cylindrical gallery from another rock-nest, and B 1, the same gallery seen in front, of the natural size. Plate VI., p. 35, fig. A. — A sprouting hempseed, part of the radicle of which has been gnawed by the ants, of the natural size ; A 1, the same, magnified, rad. radicle ; A 2, an entire sjjrouting seed of the same, magnified ; B, a sprouting pea, part of the radicle of which has been gnawed off; B 1, the same, magnified; B 2, the same stripped of its EXPLANATION OF PLATES. coat, and showing the two seed leaves; C, a sprouting "canary-seed" (the grain of Phalaris canarioisis) , part of the fibril of which has been gnawed off; CI, the same, magnified, rad. the radicle which remains tmdeveloped, and /6. the fibril or first rootlet; C 2, an unmutilated sprouting " canary seed ;" D, a mass of earth taken out of the heart of a nest of Atta barbara, in which a spherical cell, made of hardened earth, was buried. It contained grass seeds, among which 1 found ants at work, and seeds of the same grass still in their husks lay in the gallery leading up to the entrance of this cell ; D 1, the same, further freed from the earth, and having part of one side removed, so as to show the interior and the small lower opening leading out from the bottom of the cell. PART II.— TRAP-DOOR SPIDERS. Plate VII., p. 88, fig. A. — The nest of Cteniza fodiens, the lower part of which is seen in section lying in the earth, the door is artificially repre- sented as partly open; A 1, surface of the door viewed from above; A 2, the spider ; A 3, the spider deprived of its legs, from a specimen preserved in spirits [figs. A, A 1, A 2, and A 3, are of the natural size] ; A 4, the spider viewed sideways, with the legs removed ; A 5, the eyes, viewed from above and in front ; A 6, the cephalothorax and falces ; A 7, the left hand falx, viewed from the inner side ; A 8, the fang of the same ; A 1), the tarsal joint of the foremost right leg ; A 10, one of the two larger and the smallest claw of the same [figs. A 4, 5, 6, 7, 8, 9, and 10, all magnified]. Fig. B, the door of a nest of the same kind, concealed by lichens, below which, on the left hand, the doors of two miuute nests of Nemesia meridionalis are seen ; B 1, the same, with the doors open ; C, the door and mouth of tube of a nest similar to that at A ; C 1, the upper surface of this door, which is slightly convex. Plate VIII., p. 94, fig. A. — The nest of Nemesia ccvmentaria ; A 1, the door of the same, partially open; A 2, the spider; A 3, the same deprived of its legs, from a specimen preserved in spirits [figs. A, A 1, 2, and 3, of the natural size]; figs. A 4, 5, 6, 7, 8, 9, and 10 as in Plate VII., and magnified ; B, a moss-covered lump of earth, in which the door of a nest of the same type as that at A lies concealed ; B 1, the same, with the door open ; C, the door and mouth of another similar nest, showing the claw marks on its imder surface ; D, the closed door of a third nest of the same kind ; D 1, the same, opened. Plate IX., p. 98, fig. A. — Thenestof Nemesia meridional is; A 1, the open sur- face-door and mouth of the tube of the same ; A 2, the inner and upper surface of the lower door ; A 3, the spider ; A 4, the same deprived of its legs, from a specimen preserved in spirits [figs. A, A 1, 2, 3, and 4 are of the natural size] ; A 5, the spider viewed sideways, with the legs EXPLANA TION OF PL A TES. xi removed ; A 6, the eyes, viewed from above and in front ; A 7, the ce])halothorax and falces; A 8, the left hand falx viewed from the inner side; A 9, the fang of the same; A 10, the tarsal joint of the foremost right leg; All, one of the two larger and the smallest claw of the same [figs. A 5, 6, 7, 8, 9, 10, and 11, magnified] ; B, a mass of earth containing the minute nest of a young spider (A'', mcridionalis) ; B I, the lower door of this nest; B 2, the spider [figs. B, B 1, and 2, of the natural size]. Plate X., p. 100, fig. A. — Part of a nest of N. mcridionalis ; B, the new and larger upper door of a nest of this spider, with the former and smaller upper door partially united to it ; C, another example of enlargement in the upper door of the same spider, showing traces of two previous doors DOW incorporated. [All the figures are of the natural size.] Plate XI,, p. 105, fig. A. — The upper part of a nest of N. meridionalis con- cealed in a plant of Ceterach fern ; A 1 and A 2, a minute cork-door, closed and open, which I saw constructed by a very young spider [either Cteniza fodiens, or, more probably, Nemesia ccementaria] at the mouth of a hole in the mass of earth containing the nest of N. meridionalis figured at A. This hole may be seen on the right of the fern. B, the door of a small nest of N. meridionalis, as seen from above, in its natural position in a steeply sloping bank ; B I, part of the same nest placed in an upright position, and showing the surface door open and the lower door closing the branch j B 2, the same with the lower door pushed across so as to close the main tube ; B 3, 4, and 5, different views of this second door. [All the figures in this plate are of the natural size. ] Plate XII., p. 106, fig. A, — The nest of N. Eleanora with the surface door artificially represented as being open ; A 1, the outer side of the surface door of the same nest into which mosses of two kinds are woven ; A 2, the second door of the same nest ; A 3, the spider ; A 4, the same deprived of its legs, from a specimen preserved in spirits [figs. A, A 1, 2, 3, and 4 are of the natural size] ; fig. A 5, the spider viewed sideways, with the legs removed ; A 6, the eyes viewed from above and in front ; A 7> the cephalothorax and falces ; A 8, the left-hand falx viewed from the inner side ; A 9, the fang of the same ; A 10, the tarsal joint of the foremost right leg ; A 1 1 , one of the two larger and tlie smallest claw of the same [figs. A 5, 6, 7, 8, 9, 10, and 11, magnified]; fig. B and B 1 , the upper part of the tube and door of a nest of N. Eleanora which partially projected beyond the surface of the earth and was clothed with living moss. [Figs. B and B 1 are of the natural size.] PART I. HARVESTING ANTS. PART I. HARVESTING ANTS. It was in May, 1869, that Mr. Bentbam in his presidential address to the Linnean Society called attention to the want of reliable information as to the existence of such subterranean accumulations of seeds as are popularly supposed to account for the sudden appearance on railway cuttings, gravel from deep pits, and the like, of crops of weeds hitherto unknown in a district. He suggested that it might repay the trouble if some accurate observers were to take this in hand, and investigate the matter both by examining samples of undisturbed soil taken from various depths,^ — when, if any seeds of moderate size were present and un- decomposed, it would be tolerably easy to distinguish them, — and also by ascertaining what means of transport exist by which seeds may be scattered over exposed surfaces, and thus explain the difficulty without having recourse to liypothetical supplies of sound though long-buried seeds.* * M. Kerner of Innspruck has lately adduced some facts bearing on the question of the transport of seeds by the wind, having examined the collec- tions of animal and vegetable substances found on the icy surfaces of glaciers and the plants growing on moraines. Judging from the facts thus obtained, he attributes but a small influence to this agency, as the specimens dis- B 2 4 HARVESTING ANTS, As I listened, the question occurred to me whether the ants, which I had observed carrying seeds to their nests at Mentone, might not be unconscious agents on a small scale, both in the distribution and the subterranean storing of seeds. When at a later time I made this suggestion to some of our leading naturalists, I learned with considerable surprise that the unanimous opinion of our highest modern autho- rities on the subject is opposed to the belief that European ants ever do systematically collect and make provision of seeds, and that the instances of such occurrences in tropical climates remain as isolated thouq-h undoubted facts which it is difficult to ex- plain. I was not then aware that towards the middle of last century the ancient belief, dating from the time of Solomon, that ants habitually show forethought and husbandry in the collection of supplies of seeds and grain had begun to be called in question, and that our most able observers, such as Huber, Gould, Kirby and Spence, and at the present day Mr. Frederick Smith, had by close scrutiny of the habits of these creatures proved that, wherever personal investigation had enabled them to put the matter to proof, no trace of harvesting was found.* covered belonged to the fauna and flora of the immediate vicinity, and not one of these specimens must needs have come from a distance. See alistvact of his pajjer in Gardener's Chronicle, Feb. 3, 1872, p. 143, and in 'Nature' for June 27, 1872, p. 164. * I have myself on many occasions thrown seeds in the track of the com- mon English ants, and my experience was, up to the past summer (1872), similar to that of the above-named naturalists, but I have lately, by the merest chance, become acquainted with a curious exception to this rule. It happened as follows. I was gathering some fresh capsules of the common sweet violet in a garden at Richmond, near London, and in pouring the seeds HARVESTING ANTS. 5 However, just as the ancient writers, judging from their own experience and from the reports of others, had erred in attributing to ants in general the habit of seed-storing possessed by certain species commonly found in the south, so have modern naturalists fallen into the mistake of denying it to any of the European species. The older authors who lived in Greece and Italy, and the mediseval authors who drew their information in great measure from the former, being familiar with the fact that some ants habitually collect large supplies of seed, went so far as to assert, or to imply, that all European ants do so; the authors of the present day, on the other hand, generalizing too freely from their experience of ants found near their northern homes, maintained and maintain the very reverse. So long as Europe was taught natural history by southern writers the belief prevailed ; but no sooner out of my hand into the paper hag made to receive them, a few were spiUed on the ground. In a short time afterwards I was greatly surprised to see some of these spilled seeds in motion, being carried by the common black ant (Formica nig>a) into its nest. On seeing this I hastened to get some more fresh violet seeds, and also a quantity of seeds taken from ant 3 granaries at Mentone, and scattered these where the other seeds had lam. After watching for half an hour a few of the violet seeds were carried in, but not one of the granary seeds was removed, though these were examined with some curiosity. I repeated this experiment twice afterwards on a dis- tinct colony of ants of the same kind and obtained exactly the same result. I opened the nest of the former colony on the day after they had carried in the seeds, but failed to find these or any stores of other seeds. I am incUued to think that the ants took these seeds believing them to be larvffi of other ants which they might eat ; for fresh seeds of violet are not very unlike the larva of certain ants, as, for example, those of Aita barbara, ficrured at Plate I., Fig. E.. p. 21, the semi-transparent membranous appendage partly concealing the seed and giving it a fleshy appearance. I think this the more likely because on two occasions the seeds which had been carried into the nest were subsequently thrown out by the ants, which had I believe discovered their mistake. 6 HARVESTING ANTS. did the tide begin to turn, and the current of infor- mation to flow from north to south, than the story became discredited. It is interesting now to recal a few of the allusions to the harvesting ants made by ancient authors, some of which contain tolerably accurate accounts of what was to them a familiar sight or a universally accepted fact. The passages in Proverbs* are the following : " Go to the ant, thou sluggard : consider her ways and be wise ; which, having no guide, overseer, or ruler, pro- videth her meat in the summer, and gathereth her food in the harvest." " The ants are a people not strong, yet they prepare their meat in the summer." Hesiodf speaks of the time " When the provident one (the ant) harvests the grain." OTi r' icfjOif awpov df^arai. Horace I also alludes to the foresight of the ant, who is *' haudignara ac non incauta futuri." Virgil § compares the Trojans hastening their departure to harvesting ants, and the passage has been thus rendered by Dryden : — ' ' The beach is covered o'er With Trojan bands, that blacken all the shore : On every side are seen, descending down. Thick swarms of soldiers, loaden from the town, Thus, in battalia, march embodied auts, Fearful of winter, and of future wants, * vi. 6-8 and xxx. 25. + Works and Days, 776. X Satires I. i. 33. § ^neid, Bk. iv. I. 402. " Ac velut ingentem formicae farris acervum Quum populant, hiemis memores, tectoque reponunt : It nigrum campis agmen, pra?damque j^er herbas Convectant calle angusto ; pars grandia trudunt Obnixffi frumenta humeris ; pars agmina cogunt, Castigantque moras ; opere omnis semita fervet." HARVESTING ANTS. ? T' invade the corn, and to their cells convey The plundered forage of their yellow prey. The sable troops, along the narrow tracks, Scarce bear the weighty burden on their backs ; Some set their shoulders to the ponderous grain ; Some guard the spoil ; some la?h the lagging train ; AU ply their several tasks, and equal toil sustain." Indeed, it would seem that among the people in- habiting the shores of the Mediterranean it was almost as common to say " as provident as an ant as it is with us to say " as busy as a bee." Plautus* introduces a slave who, when attempting to account for the rapid disappearance of a sum of money of which he had charge, says, " Confit cito Quam si tu objicias formicis papaverem." " It vanished in a twinkling, Just like poppy seed thrown to the ants. ' ' Any one who has seen the eagerness with which certain southern ants seize upon seeds thrown in their path will appreciate the correctness of this simile. Claudius Ji^lianus, who lived in the time of Hadrian, gives a detailed account of the habits which he attri- butes to ants,t from which the following is a transla- tion : " In summer time, after harvest, while the ears are being threshed the ants pry about in troops around the threshing floors, leaving their homes, and going singly, in pairs, or sometimes three together. They then select grains of wheat or barley, and go straight home by the way they came. Some go to collect, others to carry away the burden, and they avoid the way for one another with great politeness and consi- deration, especially the unburdened for the weight * Trinummus, Act ii. sc. 4, 1. 7. f ,^ian, De Natura Animalium, ii. 25. 8 HARVESTING ANTS. carriers. Now these excellent creatures, when they have returned home, and stored their granaries with wheat and barley, bore through each grain of seed in the middle ; that which falls off in the process becomes a meal for the ants, and the remainder is unfertile. This these worthy housekeepers do, lest when the rains come the seeds should sprout, as they would do if left entire, and thus the ants should come to want. So we see that the ants have good share in the gifts of nature, in this respect as well as others." Further on* he gives a very interesting account of their mode of collecting and preparing the grain, many details of which I can myself substantiate from personal observation, though I have never seen ants actually at work upon the ears of corn. "But when the ants start a foraging, they follow the biggest, who take the lead as generals. And when they come to the crops, the younger ones stand under the stalk, but the leaders ascending gnaw through the culms, as they are called [ovpayovq, " the stalk ends on which the ears grow " (Lid. and Scott, Gr. Lex.), probably meaning that they detach the separate spikelets of which the ears are composed], of the ears [Kapiri/nwv], which they throw to the people below. These busy themselves with cutting away the chaff and peeling off the envelopes which contain and cover the grain. So the ants, though they need no threshing time, nor men to winnow for them, nor an artificial draught of wind to separate corn and chaff, yet have the food of men who both plough and sow for it." ^Elian appears also to have heard reports of the habits of ants in • Lilian, De Nat. Anim., lib. vi. chap, xliii. HAR VESTING ANTS. 9 tropical countries, for he says,* " Certainly the Indian ant is also a wise creature They leave one open- ing at the top (of the nest), by which they have their exits and entrances, when they come bearing the seeds which they collect." I have never myself found seeds bored through the centre in the way recorded above, but it is possible that different species of ants may treat the seeds in other ways than those observed by me ; or, on the other hand, ^lian may have mistaken the gnawing off the radicle of the seed, a process which I shall describe from personal observation below, and imagined that the seed itself was pierced. ^ Aldrovandus, writing in the sixteenth century, speaksf of the ants as storing seed and of their gnaw- ing, " illud principium sen acumen grani, e quo germen emitti a tritico solet" — that is to say, the radicle. But it is not clear whether Aldrovandus treats of what he has himself seen or refers to the account given by a cer- tain Bishop, Simon Mariolus, who, he says " in his most pleasant and learned work, introduces a philo- sopher as taking his walks abroad and examining an ant's nest with its seed store," &c. The lively fable of the ant and the grasshopper, as related by La Fontaine, has done much towards fami- liarizing and keeping alive in the minds of many of us the idea that ants habitually provide stores against the winter ; but we must not infer from this narration that the witty French author had ever cared to exa- mine for himself whether the fable, which he borrowed from ^sop, had its foundation in fact or not. The * Id. lib. xvi. 15. I Aldrovandus, De Insectis, lib. v. (de Forinicis). 10 HARVESTING ANTS. following translation from, the Greek original* bears in a much higher degree the impress of personal and accu- rate observation. Mvpfx-))K^Q KOI Tfrri^ : The Ants and the Grasshopper. Once in winter time the ants were sunning their seed- store which had been soaked by the rains. A grass- hopper saw them at this, and being famished and ready to perish, he ran up and begged for a bit. To the ant's question, "What were you doing in summer, idling, that you have to beg now ? " he answered, " I lived for pleasure then, piping and pleasing travellers." " 0, ho ! " said they, with a grin, " dance in winter, if you pipe in summer. Store seed for the future when you can, and never mind playing and pleasing tra- vellers."! It would be easy to multiply instances in which the older authors allude to this habit, but enough have been given to afford a sample of what may easily be found repeated elsewhere, and I will now quote a few instances which illustrate the more modern belief, utterly opposed to that so long main- tained by the ancients. Messrs. Kirby and Spencej discuss the matter in the following terms : — "When we find the writers of all nations and ages unite in affirming that, having deprived it of the power of vegetating, ants store up grain in their nests, we feel disposed to give larger credit to their assertions. Writers in general have taken . . . (this). . . for granted. But when observers of nature began to examine the manners and economj'" of these * For this translation and all the foregoing extracts from ancient and mediaeval authors T have to thank my brother, M. W. Moggridge. f ^-Esopicae Fabulae (Tauchuitz edition), p. 92. X Entomology, ed. 7 (I85G), p. 313. HARVESTING ANTS. 11 creatures more narrowly, it was found, at least with respect to the European species of ants, that no such hoards of grain were made by them ; and, in fact, that they had no magazines in their nests in which provi- sions of any kinds were stored up." They then proceed to explain how easily the white pupa^, which the ants carry about in their jaws, may have been mistaken for grains of wheat, and to inform us that the accurate observations of Mr. Gould, pub- lished in 1747, were among the first which led to the correction of this error. " However," they continue, " it may be otherwise with exotic ants, for although during the cold of our winters they are generally torpid and need scarcely any food, yet in warmer regions, during the rainy seasons, when they are pro- bably confined to their nests, a store of provisions may be necessary for them." The author of the article on ants in Smith's Dic- tionary of the Bible says, in reference to the assertion that ants store seed, that " observation of the habits of ants does not confirm this belief." Latreille* denies it in the following emphatic terms : *'!N'attribuons pas a la fourmi une prevoyance inutile: engourdie pendant I'hiver, pourquoi formeroit elle des greniers pour cette saison ?" Huber again throws the weight of his great autho- rity into the scale against the ants, when he says,t " I am naturally led to speak in this place of the manner in which ants subsist in the winter, since we have relinquished the opinion that they amass wheat and other grain, and that they gnaw the corn to prevent * Hist. Nat. des Fourmis, 1802. + Huber, on Ants, translated by J. E. Johnson, 1S20. 12 HARVESTING ANTS. it from germinating." He then goes on to show how the ants are frequently torpid during the winter, and that when it happens that a few warmer days wake them up to life, they can always find a few a23hides also on the alert ; for, strange to say, the same degree of warmth which rouses the ants calls forth the aphides also. It would appear that ants in the northern parts of Europe feed on the honey-dew of aphides, and on animal matter when they can get it ; and up to the present time the belief prevails among our modern naturalists that they are limited to the same diet in all parts of Europe. It is now well known, however, that exceptions must probably be made in tropical countries, for the observations of Lieut. -Col. Sykes* and Dr. Jerdonf have shown that many ants in India collect grain in large quantities, robbing the crops and i:)lants cultivated in gardens, and even stealing seeds put away in drawers, the inference being that they employ them for food. The same observers have re- corded how the ants may be seen after wet weather bringing out the grain to dry in the sun. Dr. Lincecum has also given a very interesting account J of the habits of the " agricultural ant" in- habiting Texas, Mp-mica {Jlfa)barbata, which not only stores the grain of a particular rice-like grass, but is said * Lieut.-Col. Sykes, Description of New Indian Ants, in Trans. Ent. Soc. Lond., i. 103 (1836), where a single species of ant, which he names Atta providens, is described, and its habit of harvesting recorded. f Dr. Jerdon, Madras Journal Lit. and Sc. (1851), where three species are stated to harvest seeds on a large scale — namely, CEcodoma (or Atta) providens, CEcodoma diffusa, and Atta rufa, all of which belong to the same section of ants as our Meutouese harvesters, Atta barbara, Atta structor, and J-'/ieidole (or Atta) ineyaccphala. These very interesting observations of Dr. Jerdon's, as well as tliose of Lieut. -Col. Sykes, will be found in Appendix B. t Published in the Journal of the Liunean Society of London, vol. vi. p. 29. 1861. HARVESTISG ANTS. 13 to maintain a clean crop of tins plant around its nest, suffering no weed to appear among it, and harvesting the crop in its proper season. The Sauba ant {(Ecodoma cephalotcs) has been seen by Mr. Bates plundering baskets containing mandioca meal (an impure form of tapioca) in Brazil, and this in so wholesale a manner as shortlj^ to threaten the loss of the entire supply ; and Dr. Delacoux records * the presence in New Granada of a monstrous ant, called by the natives Arieros, a word which, I am informed, is of Arabic extraction, and means the carrier, which emptied an entire sack of maize belonging to him in a single night. It seems strange that while travellers have reported the seed-storing habits of ants in far distant countries, our naturalists at home should have not only remained unaware of its existence inEurope, but even strenuously denied it. It is certain, however, that naturalists and others in southern Europe are more or less aware of the fact, but I have been unaljle to learn that any accurate account of the habits of harvesting ants has hitherto been published, or that any one has taken pains to discover what becomes of the seed so laboriously obtained. It is true that in the Enciclopedia PopoIare\ extracts are given from the remarks made by M. Genej on the subject, in which he assumes that the fact that ants collect and carry to their nest large supplies of grain and seed is well known, but states that he is at * Notice sur les Mceurs et les Habitudes de quelques Especes de Formi- ciens des Climats Cliauds. Eev. Zool., Mai, 1848, p. 1849. + Article Formica, vol. v. p. 143-4. (Turin, 1845). % Memorie per servire alia Storia Naturale di alcuni imenotteri, iniLlislied at Modena, in 1842. 14 HARVESTING ANTS. SL loss to conceive how tliej employ tliem, unless it may be that they use them as materials for the con- struction of their galleries, for they cannot eat such hard substances, all their food being either liquid or of the nature of juices, " gli alimenti sono sempre materie liquide o materie sugose. Quanto ai corpi duri e secchi che le formiche raccolgono, io non so altrimenti riguardarli che come materiali di costruzione." It will be understood, I think, from what Las gone before, that thus far nothing has really been ascertained as to the exact state of the case ; for though the Italian author just quoted was aware that certain ants in the Medi- terranean region do store seed, his knowledge went no further. Nor am I aware that any French author has published an account of this habit and its object ; and in a recent abundantly illustrated volume founded on a work by M. Emile Blanchard,I find, on the contrary, the following very emphatic denial of its existence : — " The curious idea which appears to have commenced in very remote times, and to have been carried down by tradition, and which was assisted by the results of careless observations, concerning the habits of the ants in collecting and storing up provisions, as it were under the influence of a wise foresight, is evidently incorrect."* There was, therefore, clearly an opening here for close observation, and this I determined to do my best to supply. When I set out again from England in October, 1871, on my way to Mentone, I had obtained an idea of some of the leading points which needed to be * The Transformations of Insects : an adaptation for English readers of M. Emile Blan chard's Metamorphoses, Mcjeurs, et Instincts des InsecteS; p. 196. London. 1871. HARVESTING ANTS. 15 cleared up, and I was greatly encouraged in my attempt by the interest exj^ressed in the subject by several of our leading naturalists, among whom I may especially mention Mr. Frederick Smith.* Plainly the first thing to do was to determine whether the seeds which I had watched the ants carry to their nests were separately stored in sub- terranean granaries, as they would be if the ant really provides for the future ; or whether they were merely strewed here and there, or used as building materials. Next I must, if possible, obtain conclusive evidence as to the use to which the ants put the seeds thus collected ; whether they eat them or turn them to some other account. Again I must observe whether the seed-collecting ants also search for aphides, and what other kinds of food they obtain. Then another very interesting question remained — namely, whether all southern ants uniformly collect seed, and to the same extent, or whether the habit is peculiar to certain species. These, and many other subjects of inquiry con- nected with them, readily suggested themselves to my mind, and it will now be my endeavour to show how far I have been able to throw light upon them. The habits recorded in the following pages refer exclusively, unless special notice is given to the contrary, to Atta barbara, the black ant represented on Plate I. We have, as far as I am aware, only four bona fide harvesting ants on the Biviera — namely, Atta barbara under two forms, the one wholly black * I am very greatly indebted to Mr. Smith for much kind assistance, and especially for having named the specimens which I collected. 16 HARVESTING ANTS. the other red-lieaded ; Jlfa strucfor, a creature very similar to harhara, but of a claret-brown colour ; and a minute yellow ant, the large workers of which have gigantic heads, named Pheidole (or Atta) megacepliala. My renewed observations at Mentone were carried on from October, 1871, to May, 1S72, and I was able during that interval to become a frequent visitor to a warm and sheltered valley, which lay but a few minutes' walk from the house in which 1 lived, and in which thirty nests of the most active of the seed-storing ants were to be found. Full therefore of my intention to resolve this difficulty if possible, I set out on October 29, 1871, immediately after my return to Mentone, to revisit this valley, where, in the previous May, I had seen the ants busily engaged in cutting, carrying, and sorting their harvest. ' The spot in question was a rough slope of soft sandstone rock, with accumulations of sandy soil in the hollows, covered with a sparse and scrubby vegetation, composed of Cistiis {C. sahifolms), pot- herb thyme, black lavender {Lavandula stadias), spiny broom {Cahjcofome spinosa), overshadowed here and there by a few scattered stone and maritime pines, and intermixed with coarse grasses and some smaller plants. Cultivated lemon terraces lay on the edge of the wild ground lower down in the valle}^, and at this season, as also in the late spring, these terraces were overgrown with a rank crop of weeds, most of which were in seed. I had scarcely set foot on the garrigue, as this kind of wild ground is called, to distinguish it from HARVESTING ANTS. 17 meadows or terraced land, before I was met by a long train of ants, forming two continuous lines, hurrying in opposite directions, the one with their mouths full, the others with their mouths empty. It was easy enough to find the nest to which these ants belonged, for it was only necessary to follow the line of ants burdened with seeds, grain, or entire capsules, which had their heads turned homewards, and there, sure enough, at about ten yards distance, and partly shaded by some small Cistus bushes, lay the nest, to and from the entrances of which the incessant stream of incomers and outgoers kept flowing. The proceedings of the ants were the same as those previously observed in the late spring (April and May), the workers usually seeking their harvest at some distance from the nest, and going in search of it as far as the cultivated ground, where the crops of weeds were more abundant and more varied. In a few cases, however, where the terraces were too far distant, they contented themselves with plundering the grasses, pea-flowers, honeywort, and the other denizens of the garrigue. In one case I was able to follow the thread-like column of workers from the nest to the weedy terrace where the plants grew from which they were gathering the seeds, and found that the nearly continuous double line measured twenty- four yards. Even this gives but an inadequate idea of the number of ants actively employed in the service of this colony, for hundreds of them were dispersed among the weeds on the terrace, and many were also employed in sorting the materials and in attending to the internal economy of tlie nest. Still 18 HARVESTING ANTS. this affords some evidence of the systematic and extensive scale on which foraging is carried on by this ant, and of the high importance which these creatures attach to their provision of grain. It is not a httle surprising to see that the ants bring in not only seeds of large size and fallen grain, but also green capsules, the torn stalks of which show that they have been freshly gathered from the plant. The manner in which they accomplish this feat is as follows. An ant ascends the stem of a fruiting plant, of Shepherd's-purse {CapseJIa Bursa pasforis) let us say, and selects a well-filled but green pod about midway up the stem, those below being ready to shed their seeds at a touch. Then, seizing it in its jaws, and fixing its hind legs firmly as a pivot, it contrives to turn round and round, and so strain the fibres of the fruit-stalk that at length they snap. It then descends the stem, patiently backing and turning upwards again as often as the clumsy and disproportionate burden becomes wedged between the thickly set stalks, and joins the line of its companions on their way to the nest. In this manner capsules of chickweed {Alsine media) and entire calyces, contain- ing the nutlets of Calaminth, are gathered ; two ants also sometimes combine their efforts, when one stations itself near the base of the peduncle and gnaws it at the point of greatest tension, while the other hauls upon and twists it. I have never seen a capsule severed from its stalk by cutting alone, and the mandibles of this ant are perhaps incompetent to perforin such a task. I have occasionally seen ants engaged in cutting the capsules of certain plants droj3 them and allow their companions below to carry them away; and this corre- BAR VESTING A NTS. 1 9 sponcis with tlie curious account given by iElian* of the manner in which the spikelets of corn are severed and thrown down " to the people below," rJ ^hfXM rJ KaT(o. If the incoming and weight -carrying column of ants be closely examined it will be found that though the great majority of workers are bringing seeds in some form to the nest, a few are burdened with other and more miscellaneous materials. Occasionally one or two may be detected carrying a dead insect, or crushed land-shell, the corolla of a flower, a fragment of stick, or leaf, but I have never seen aphides brought in to the nest or visited by this ant or by Atta structor. It sometimes happens that an ant has manifestly made a bad selection, and is told on its return that what it has brought home with much pains is no better than rubbish, and is hustled out of the nest, and forced to throw its burden away. In order to try whether these creatures were not fallible like other mortals, I one day took out with me a little packet of grey and white porcelain beads, and scat- tered these in the path of a harvesting train. They had scarcely lain a minute on the earth before one of the largest workers seized upon a bead, and with some difficulty clipped it with its mandibles and trotted back at a great pace to the nest. I waited for a little while, ray attention being divided between the other ants who were vainly endeavouring to remove the beads, and the entrance down which the worker had disappeared, and then left the spot. On my return in an hour's time, I found the ants passing * Vide supra, p. 8, 20 HARVESTING ANTS. unconcernedly by and over the beads which lay where I had strewn them in apparently undiminished quantities ; and I conclude from this that they had found out their mistake, and had wisely returned to their accustomed occupations. I have often amused myself by strewing hemp and canary seed or oats, all of which form heavy burdens for the ants, near their nests ; and it is a curious sight to watch the eagerness and determina- tion with which they will drag them away. It is interesting also to note how on the following day the husks of these seeds will appear on the rubbish- heaj), or sometimes, after a shower of rain, they will be brought out by the ants with the point of the little root (the radicle or fibril as the case may be) gnawed ofi" (see Figs. A, B, C, Plate VI., p. 35). It frequently happens that on the wild hillside the position of a nest of Jtta harhara is indicated by the presence of a number of plants growing on or round the kitchen midden, which are properly weeds of cul- tivation, and strangers to the cistus- and lavender- covered banks of the garrigue. These have sprung from seeds accidentally dropped by the ants, and which they had obtained from the lemon terraces. Thus when you see little patches of ground from one to three feet long and broad, covered with such plants as fumitory [Fumaria), oats [Avena), nettles {JJrtica mem- hrariacea), four species of Veronica, chickweed {Alsine media), goosefoot {CUenopodiwii), Riimex Bucephale- phoriis, wild vnSiYi^oXdi {Calendi/la arvensis), Antirrhinum Oroniiurn, Linaria simplex, and Cardamine kirsuta, you may confidently expect to find a colony of these ants close at hand. f^ l> ^^4 w V.U 'W 0'% J\ ' \ -. li.^^ S2> ■E^ HARVESTING ANTS. 21 These plants are sometimes found along the sides of miniature gullies and crevices in the rock, where they have been washed by little runlets of water formed in seasons of heavy rain, and thus these interloping plants are occasionally dispersed and brought into competition with the rightful occupiers of the ground. Aita structor and A. barbara do not employ any materials in the construction of their nest, simply excavating it out of the earth itself, or occasionally out of the sandy rock, and the large mounds, in great part composed of vegetable matter, which may fre- quently be found at the entrances of their nests, are nothing more than the rubbish heaps and kitchen middens of each establishment. These consist in part of the earth pellets and grains of gravel which the ants bring out from their nest when forming the sub- terranean galleries, but principally of plant-refuse such as the chaff of grasses, empty capsules, gnawed seed- coats, and the like, which would occupy much space if left inside the nest (see Plate I., Fig. A.). While an army of workers are employed in seeking and bringing in supplies, others are busy sorting the materials thus obtained, stripping off all the useless envelopes of seed or grain, and carrying them out to throw away. Thanks to the unwearied activity with which this divided labour is carried on the kitchen middens speedily rise in the harvest season, and in places where they are not exposed to the action of wind and rain, often acquire a considerable size, so much so that sometimes, if collected, one alone might fill a quart tankard. It was the sight of such a refuse mound, and an ex- amination of the materials which composed it, — many 2% HARVESTING ANTS. of which show that they were once parts of seeds, &c. . the albuminous contents of which had been extracted through holes gnawed in the side, — that gave me the conviction that large stores of seed must lie hidden below in the nest ; for if it were true, as some have suggested, that the ants employ the grain and seeds which they collect as materials for the construction of their nest, they would certainly not reject such parts as the chaff of grasses and the like, which are admirably suited for the purpose, and are actually used for this end by other species of ants. It was therefore with the greatest confidence as to the result that I opened the nests of Atfa harhara in search of granaries and seeds. My first attempt was made upon a nest lying in a hollow where there was a rather deep bed of soil, and the galleries extended so far on either side and in a downward direction tbat, though I removed enough soil to fill a wheelbarrow, I failed to reach the arcana of the nest, and saw neither chambers nor granaries. Yet I frequently encountered workers carrying seeds downwards along the subterranean passages. I then selected a nest where the coarse and hard rock lay much nearer to the surface, barring their downward course, and compelling the ants to extend their nest in a horizontal direction. Here, almost at the first stroke, I came upon large masses of seeds carefully stored in chambers prepared in the soil. Some of tliese lay in long subcylindrical galleries, and, owing to the presence in large quantities of the black shining seeds of amaranth {Amaranihis Biitum, &c.), looked like trains of gunpowder laid ready for blasting. Fig. A, Plate II. represents a trowel- -^' '-^Ja^ E^ Pla/^ III. ■:<■&■: m :v.i:^ X,7 I ,:, v-.WricV.Vt- Vincent Bicol^s Day&Son.lmp L.R-eeve & C"^ Puilisiers. HARVESTING ANTS, 23 ful of eartli taken from this nest, and lifted with care so as to leave the seeds almost in situ. Others were massed together in horizontal chambers, having a concave roof and a flat and carefully prepared floor. The texture of the floor usually differs markedly from that of the surrounding soil, and the fine grains of silex and mica which are selected for its construction are more or less cemented together, so that the floor will sometimes part, when dry, from the soil about it, as caked and dry mud separates from a gravel path (see Plate III.). On carefully examining a quantity of the seeds, grain, and minute dry fruits taken from the granaries, I found that they had been gathered from the following plants : fumitorj^ {Fumaria Capreolata, &c.), amaranth {Amar- anihus Blitum, &c.), Setaria, and three other species of grasses, honeywort {Alyssum maritimiim), Veronica^ and from four unrecognised species, one of which was a pea-flower. There were therefore in this nest seeds, &c., which had been taken from more than twelve distinct species of plants, belonging to at least seven separate families. The granaries lay from an inch and a half to six inches below the surface and were all horizontal. They were of various sizes and shapes, the average granary being about as large as a gentleman's gold watch. I was greatly surprised to find that the seeds, though quite moist, showed no trace of germination, and this was the more astonishing as the self-sown seeds of the same kinds as those detected here, such as fumitory for instance, were then coming up abun- dantly in garde as and on terraces. The seeds of Odontites lutea afford a curious test of the presence of 24 HARVESTING ANTS. moisture in the granaries, and it will usually be found that, when they are recently taken out of the nest, they are of a greenish colour and semi-transparent horn- like texture, which changes on exposure to the air to a chalky white and opaque appearance, due to the drying of the coat of the seed. The fact of the sound condition of the seeds in these granaries seemed to me so very strange and difficult to explain that I determined to pay special attention to the subject, and with this view collected and carefully examined large quantities of the grain and seeds taken at different times from the stores of twenty-one distinct nests, the first of which was opened on October 29th, and the last on May 5th. In these twenty-one nests out of the thousands of seeds taken I only found twenty-seven in seven nests which showed trace of germination, and of these eleven had been mutilated in such a way as to arrest their growth. The sprouting seeds were found in the months from November to February, while in the nests opened in October, March, April, and May, no sprouted seeds were discovered, though these latter months are certainly highly favourable to germination. It is therefore extremely rare to find other than sound and intact seeds in the granaries, and we must conclude that the ants exercise some mysterious power over them which checks the tendency to germinate. Apparently it is not that moisture or warmth or the influence of atmospheric air is denied to the seeds, for we find them in damp soil, in genial weather, and often at but a trifling distance below the surface of the ground ; and I have proved that the vitality of the seeds is not affected by raising crops of young HARVESTING ANTS. 25 plants, such as fumitory, pellitory. Polygonum avi- culare, and grasses, from seeds taken out of granaries.* I have frequently remarked that it is the seeds last collected before a fall of rain which are brought out in a sprouting condition from the nest ; for T have observed in cases where I had recentl}^ scattered seeds near wild nests, that it is these which are carried out from the nest and placed to dry after a wet night ; and so in the case of a nest which I kept in captivity, when a variety of different seeds had been successively sup- plied to the ants, it was the cabbage, lettuce, and chicory seeds, given the day before the nest was watered, that reappeared after having been carried below, and not the hemp, canary, and mixed seeds of wild plants previously strewed on the nest. It seems possible that the process, whatever it may be, to which the ants subject the seeds which are to remain dormant may require some time, and the construction of the gra- nary chambers is doubtless a long affair, so that when ■unusually large supplies of grain, &c., are brought in by the workers some part of them may not find the necessary accommodation and attention. When the seeds do germinate in the nests, and it is my belief that they are usually softened and made to sprout before they are consumed by the ants, it is very curious to see how the growth is checked in its earliest stage, and how, after the radicle or fibril — the first growing * This experiment was tried by me on two occasions, in the former case the seeds were taken from a granary about four inches below the surface of the ground, on November 10th, and sowed two days afterwards, and several of these were up on Dec. 1st. The second trial was made on seeds found at only one and a half inch below the surface, on Dec. 29th, 1871; these were sowed iu England on June 18tli, 1872, and the young plants made their ap- pearance in large numbers ten days afterwards. 26 HARVESTING ANTS. root of dicotyledonous and monocotjdedonous seeds — has been gnawed off, they are brought out from the nest and placed in the sun to dry, and then, after a sufficient exposure, cariied below into the nest. The seeds are thus in effect malted, the starch being changed into sugar, and I have myself wit- nessed the avidity with which the contents of seeds thus treated are devoured by the ants. Figs. A,B,C, in Plate VI., p. 35, illustrate the manner in which the ants mutilate the germinating seeds and check their growth. Thus, at Fig. C 2 of Plate YI. a sprouting but uninjured canary seed {Phalaris cana- riensis) is drawn, magnified, and at Figs. C and CI the same of the natural size and magnified, after the ants have gnawed its fibril (fib.), which in this case pierces the undeveloped radicle (rad.). Fig. A 2 represents a sprouting hemp-seed, magnified,* and Figs. A, A 1, the same of the natural size and magnified, mutilated, the tip of the radicle being removed. At Figs. B, B 1, B 2, the same process is shown in the case of a small wild pea. It is, however, certain that though a few individual seeds may sprout in the nests from time to time either with or without the concurrence of the ants, the great mass remain for many weeks, or even months, quite intact, neither decaying nor germinating, whereas every one knows that if a quantity of seeds are placed in the soil in a moist and warm place, all the seeds that are of one kind will almost simultaneously begin to grow after the lapse of a fixed interval. Now if this took place in an ant's nest, the provisions * Properly a nut, for it comprises the seed and the enveloping coat of the ovary. The canary seed also, spoken of above, is a grain containing a seed. HARVESTTNG ANTS. 27 would have to be rapidly consumed at stated periods and to be frequently renewed ; but this is not the case. This is easily shown by an examination of the seeds contained in the nests in April or May, many of which will prove to belong to plants which fruit in the autumn and are not to be found later than November. Thus, for example, on May 5th at Cannes, I discovered nutlets of C^no^Iossum jnrtinii, which can scarcely have been col- lected later than the preceding October or November. Besides, during the time from the middle of January to the middle of ]\Iarch, scarcely a seed is collected under ordinary circumstances, there being extremely few wild plants in fruit at that season, and yet the granaries will be found well filled if a nest is opened at the end of this period. A knowledge of the fact that ants in warm climates accumulate large and very varied stores of seeds re- taining their power of germination, might at times be of service to travellers, by enabling them to obtain, by a stroke or two of the spade, an interesting col- lection of the seeds and the seed-like fruits of the country, when time and opportunity failed for ob- taining them in a more satisfactory manner. The following list of plants, the grain, seeds, and small dry fruits of which I have found in the subter- ranean granaries of Atta structor and A. barbara, es- pecially the latter, shows that the ants probably col- lect almost indiscriminately from any fruiting plant that falls in their way. Fumitory {Fumaria, three species), honeywort {Alyssum maritivium), narrow-leaved sun rose {Fumaria viscida and F.Spac/iii), Oxalis corniculata, 8ilene,Linum gallicum, mallow {Lavatera crefica ?), mQdi\Qk{Medica(^o), 28 HARVEST INO ANTS. wild lentil {Ervum), spiny broom {Cyiisiis sjnnosui)^ Valerianella carinata, Centaurea aspera, Odontites lutea^ Calamintha Nepefa, Polygonum convolvulus and P. avicu- /f/;^, amaranth {JmaranthusBUtum andpatulus), pellitory {Parietaria), Euphorbia, pine {Pinus), wild sarsaparilla {Smilax aspera), Setaria verticillata and >S'. italica, An- dropofjon Jschcemuiu, and of eight other plants of which J do not recognise the seeds. This list, comprising plants belonging to eighteen distinct families, might be greatly prolonged if I were to add to it the names of the seeds which I have seen the ants carry towards their nests, but have not actually detected in the granaries. Thus I have seen trains of ants burdened with the long'beaked, spirally-twisted fruits of crane's bill [Erodiuni], and, as above mentioned, with capsules of chickweed {Alsine media) and shepherd's-purse {Capsella Bursa pastoris), with whole orange pips, and even haricot beans, seeds of the New Zealand veronica (F! Andersonii), of Silenepseudoatocion, and many other garden plants, also with nutlets of the plane tree and seeds of the cypress. Pliny mentions* incidentally having watched the ants carrying away cypress seeds, and comments upon the fact that so small a creature should be able to interfere with the growth of such a noble tree. I have little doubt that tlie seed stores of the ants in botanic and other gardens, where rare plants are cultivated in southern Europe and in warm climates generally, contain samples taken from the fruits of a great many of the rarer and more interesting species as well as of the weeds and native plants. Indeed 1 * Pliny, Nat. Hist., xvii. 14, 3- HARVESTING ANTS. 29 have been told that this is the ease by my friend Dr. Bornet, who complains of the depredations committed by the ants in the gardens of the Villa Thuret, at Antibes. They go so far as to plunder the seed bags which are hung from the branches of the trees and shrubs, unless these are securely closed and tied with string ; they carry off wholesale the grass and anemone seeds,* which are scattered when the lawns are resown ; and Dr. Bornet has seen the seeds of Acacia refinoides lie heaped up by the handful at the entrances of their nests, and disappear below after a few hours. M. Germain de St. Pierre has observed similar facts at Hyeres, where he has detected large stores of cereals in the granaries of the ants, and considers that the robberies committed by these creatures are suflBcient in extent to cause a serious loss to cultivators. It is difficult to estimate the amount of seed stored in a single nest by a colony of ants both on account of the extent of these nests, and because of the number of seeds which are always lost in digging. The nests themselves also vary greatly in size. Perhaps I shall not be very far from the mark however, if I conjecture that average-sized nests contain during the winter months about half a pint of seeds. Atta structor is more frequently found near houses and in gardens than A. harhara, the latter usually living on wild ground adjoining cultivation. There was a flourishing colony oi structor in the main street of Mentone, cleverly placed at the lintel of the door of a corn chandler's store, where they were ever on the look out for stray grains of oats and wheat, which * Properly grass grain and anemone achenes. 30 HARVESTING ANTS. might chance to fall from the sacks. Another nest, in a different part of the town, got its principal subsistence from the grains of canary seed, which were scattered by the birds occupying a cage hanging outside a shop window at a little distance. Vertical section of an ant's nest. The horizontal lines represent inches of depth. The granaries of ^. sfructor are arranged in the same way as those of ^. barbara, and may, in like manner, be 0 ^ HARVESTING ANTS. 31 found stored with, seeds, and lying at depths below the surface, varying from one to twenty inches. A diagram is given in the preceding woodcut of a vertical section of a nest of barhara lying in soil sixteen inches deep, the granaries being at 1^, 2, 4, 6, 9, and 12^ inchesj as determined by actual measure- ment on the spot. In some cases, and especially where the soil is shallow, the galleries and granaries are much crowded together, as is shown in Plate IV., which represents a small mass of earth, pierced by the roots of plants, taken out of a nest of harbara, lying at two inches below the surface. When first opened all these granaries were filled with seeds. The shape of the granary chambers varies con- siderably, as may be seen by reference to the draw- ing of three floors given in Plate III., p. 23, and that shown diagrammatically in the woodcut on next page, where the white space represents the granary floor, and the dark circular spot in the centre, the aper- ture of a gallery leading downwards. I once had an opportunity of seeing a large portion of a nest of the red-headed variety of barbara laid bare by a cutting recently made through a bank at Cannes in digging the foundations of a house, which exposed a very extensive and complicated series of galleries and granaries. The lowest point at which I detected the workings of the ants was at twenty inches below the surface of the ground, and here granaries containing seeds in abundance were present, and the galleries and granaries extended over a space measuring 5ft. 9in. in a horizontal direction. In two cases I have found nests of Alt a barbara at Mentone which were carried 32 HARVESTING ANTS. far into the living rock in places where it happened to be of an even grain, and not gritty or pebbly as it fre- quently is. It was quite by chance that I first dis- covered this very interesting fact, having tracked a train of seed-bearing workers to a part of the sand- stone rock where steps had quite recently been hacked out leading to some terraces. Tioite V. .-j^ •'^•'liiBN ■-*!■>>' VincentBrooko Diyft Soa Imp LKeeve & C° Fiiblisliers . HA Pi VESTING A NTS. 33 I soon saw that the ants entered and came out from three or four small passages in the cleft surface of the rock, and that their nest actually lay in the sandstone itself. Havino^ contrived to wedg-e off several larsre flakes of the rock, which was soft in most places and might be scooped out with a strong knife, I discovered that though some of the passages of the ants followed the lines of cleavage and the cracks made by the fine wiry fibres of the bushes growing on the surface, others were frequently made in the form of tubular tunnels through the living rock. Without the aid of hammer and chisel it was not possible to follow the galleries and to secure specimens of the mined rock ; but on the next day (Dec. 7th) I returned armed with tools, and with the assistance of a friend * quarried out a portion of the nest, tracing it down eventually to twenty-three inches below the surface of the rock in a vertical, and to about sixteen inches away from the surface in a horizontal direction. At one point where the rock was almost entirely solid and without flaw or crevice, and where it was clear that the passages were entirely the work of the ants, we measured a tunnel by worming a straw down it, and found it to be ten inches in length. We subse- quently traced this tunnel or rock gallery down until it communicated with a chamber filled with winged ants and seeds of several kinds. This granary was hori- zontal, and merely an enlargement of an ordinary gal- lery of a compressed spindle-shape, flattened from above downwards, measuring as nearly as I could estimate three inches in length, by a trifle less than * I take this opportunity of expressing my tliauks to Mr. Holier t I.ightbody for help on this and other occasions D 34 HARVESTING ANTS. an inch in breadth, and half an inch in height. Tlie walls were tolerably smooth, but not prepared or glazed in the way that certain small terminal cells which I shall shortly describe were. The surfaces, however, had a very different appearance to that of the surrounding sandstone, being of a darker and brownish colour, and seeming to be coated with some kind of dressing or cement. , One of these tunnels at first took a horizontal course for two and a half inches, then descended vertically for an inch and a half to a point where it made two hori- zontal branches, and from these latter several other vertical galleries descended, two of which we were able to trace until they expanded into a cluster of small pear- shaped cells, the walls of which were quite smooth and very carefully laid with plates of mica and cement. I was able to draw this on the spot, Fig. A, Plate V., while Mr. Lightbody worked it out piecemeal with hammer and chisel. It was unfortunately impossible to se- cure more than very imperfect fragments as specimens. These terminal cells were empty when we came to them, but it is quite possible that the ants may have conveyed away larvae or winged ants from them, having received abundant notice of the coming danger from the continued jarring of the chisel-work. One entrance to this nest lay in a small accumula- tion of soil in a hollow of the rock, and it was at this point that the refuse from the nest was cast out. In- - deed, had it not been for the accidental circumstance of myhaving traced the ants to the newly hewn step in the sandstone, I might never have discovered the fact that the nests are sometimes carried deep into the living rock. With this to guide me, however, I succeeded in find- ing a second nest of the same kind, and here I was PlaU VJ ■ \ ■' ■<- r 2 '-a ' ftb.-jll ^ 'H Vim.eiitBrooksDa.yi Sou, Imp L. Reeve & C? Publishers HARVESTING ANTS. 3& able to secure better specimens of the tunnels for draw- ing (Figs. B, B 1, Plate V.,p. 33). These drawings may be taken as representing also the size and shape of the tunnels in the former nest, which were for the most part like these, beautifully cylindrical, as is shown in the front view of the tunnel at B 1. In one nest of harhara I found a curious hollow spherical dome, about an inch in diameter, the walls of which were constructed of hardened earth about two lines thick, and having a hirge circular aperture at the top and a very small one below (Figs. D and D 1, Plate VI.). This dome was imbedded below in earth which adhered to it, but it was otherwise easily separable from the soil ; its inner walls were smoothed with great nicety. It has been suggested to me that this spherical chamber was originally the work of a scarabseus, which had chanced to bury the ball containing its eggs close to the nest of the ants, and that the latter had appro- priated it after the departure of the beetle grubs. This may perhaps have been the case, but the dome was rather larger than the ball usually formed by the scarab beetle, and I have never seen one of these balls surrounded by a hardened case. The chamber thus constructed was employed as a granary, and filled, as well as the adjacent passages, with the grain of a grass {Tragus racemosus), still enclosed in the husks, among which I detected several ants at work, and also some minute white semi-transparent creatures, like spring-tails [Podunis], which abound in these ants' nests. Besides this spring-tail it is common to find in the galleries and granaries of Atta structor and A. har- hara, certain silky j^ellowish-white " silver fish" {Le- jpisma), a small white woodlouse which does not D 2 86 HARVESTING ANTS. roll itself into a ball, and at times the larvae of an elater beetle. I have observed- on more than one occasion that when in digging into an ant's nest I have thrown out an elater larva, the ants would clus- ter round it and direct it towards some small opening in the soil, which it would quickly enlarge and disap- pear down. At other times, however, the ants would take no notice of the elater, and it is my belief that the attentions paid to it on former occasions were purely selfish, and that they intended to avail them- selves of the tunnel thus made down into the soil, with a view of reopening communications with the galleries and granaries concealed below, the approaches to which had been covered up. I have frequently watched the ants make use of these passages mined by the elater on these occasions. At one time I suspected that the elater larvse might consume the seeds stored by the ants, and I therefore confined some of them in a tumblerful of earth and seeds ; but at the end of three weeks, though the larvse were strong and healthy-looking, I could not detect that any of the seeds had been touched, and even those which had sprouted remained uninjured. I have searched in vain for the beetles and staphylinidse which are known to inhabit certain ant's nests. In one nest I found (on Dec. 28) a quantity of small spherical, egg-like galls, slightly larger than but resembling the fruit of Fumaria cajyreolata, spotted with pink-brown on a yellowish or greyish ground. There was a dark spot at the point at which the mature insect would emerge, and one did escape from the egg-like cocoon while I was watch- ing, and proved to be a Ci/nijjs of very small size, but furnished with a terrible dart for puncturing its prey. HARVESTING ANTS. 87 It seems difficult to understand how it comes that these galls are systematically placed among the seeds, for it was evidentl}^ no chance occurrence, and I can only conjecture that the worker ants may have brought them in and stored them under the impres- sion that they were really seeds ! Even ants make mistakes, and of this I have given an example above (p. 19). Though I have frequently found colonies of several distinct species of ants inhabiting nests made in the earth traversed b}^ the widespread galleries of Atta strudor and barbara, I have never detected any inter- mixture of species in the chambers of a nest,* and but rarely found even the galleries and entrance used in common by more than one species. On one occasion when opening a nest of structor I cut through a colony of the tiny, large-headed, yellow ant Fheidole mcr/acejjJiala, lying in the midst of, though distinct from, the former. AVhen, however, it chanced that one of the sfructors fell from the crumbling earth into the midst of the P/ieidoIes, it was curious to see how fiercely it would be attacked, and with what terrified speed it would scamper off, without attempt- ing any resistance, and often carrying two or three Fheidohs haniicino: on to its leg's. Accidentally in this way battles do sometimes take place between ants of different species ; but by far the most savage and prolonged contests which I have witnessed were those in which the combat- ants belong to two different colonies of the same species. * Except in a few cases wbere I have seen one or two structors in nests of harhara and viceversd, and in the curious instance to be mentioned below, where one colony cousisted of nearly equal parts of structor, iarbara, and the red-headed variety of barbara. 38 HA R VESTING A NTS. Atta barbara, Formica cruentaia, F. erratica, and espe- ciall}^ Myrw'ica ccBspitiim may sometimes be seen fight- ing in this desperate fashion. Rival colonies of Mynnica ccBsjpitum often gather for tlie battle into dense masses three or four inches deep, and the place of conflict will be seen on the following day strewn with the dead, and this though the majority of the slain are carried oft* for food by the victors. But the most singular contests are those which are waged for seeds by A. barbara, when one colony plunders the stores of an adjacent nest belonging to the same species, the weaker nest making prolonged though, for the most part, inefficient attempts to recover their property. In the case of the other species of ant which I have watched fighting, the strife would last but a short time — a few hours or a day — but A. barbara wdll carry on the battle day after day and week after week. I was able to devote a good deal of time to watching the progress of a predatory war of this kind, waged by one nest of barbara against another, and which lasted for forty-six days, from Jan. 1 8 to March 4 ! I cannot of course declare positively that no ces- sation of hostilities may have taken place during the time, but I can affirm that whenever I visited the spot, and I did so on twelve days, or as nearly as possible, twice a week, the scene was one of war and spoliation such as that which I shall now describe. An active train of ants, nearly resembling an ordi- nary harvesting train, led from the entrance of one nest to that of another lower down the slope, and fifteen feet distant ; but on closer examination it appeared that though the great mass of seed-bearers HARVESTING ANTS. 39 were travelling towards the upper nest, some few were going in the opposite dii-ection and making for the lower. Besides this, at intervals, combats might be seen taking place, one ant seizing the free end of a seed carried by another, and endeavouring to wrench it away, and then frequently, as neither would let go, the stronger ant would drag seed and opponent to- wards its nest. At times other ants would interfere and seize one of the combatants and endeavour to drag it away, this often resulting in terrible mutila- tions, and especially in the loss of the abdomen, which would be torn off while the jaws of the victim retained their indomitable bull-dog grip upon the seed. Then the victor might be seen dragging away his prize, while its adversary, though now little more than a head and legs, offered a vigorous though of course ineffectual resistance. I frequently observed that the ants during these conflicts would endeavour to seize one another's antennae, and that if this were effected, the ant thus assaulted would instantly re- lease his hold, whether of seed or adversary, and appear utterly discomfited. Ko doubt the antennae are their most sensitive parts, and injuries inflicted on these organs cause the greatest pain. It was not until I had watched this scene for some days that I apprehended its true meaning, and dis- covered that the ants of the upper nest were robbing the granaries of the lower, while the latter tried to recover the stolen seeds both by fighting for them and by stealing seeds in their turn from the nest of their oppressors. The thieves, however, were evidently the stronger, and streams of ants laden with seeds arrived safely at the upper nest, while close observa- tion showed that very few seeds were successfully 40 HARVESTING ANTS. carried on tlie reverse journey into the lower and plundered nest. Thus when I fixed my attention on one of these robbed ants surreptitiously making its exit with the seed from the thieves' nest, and having overcome the opposition and dangers met with on its way, reaching after a journey which took six minutes to accomplish, the entrance to its own home, I saw that it was violently deprived of its burden by a guard of ants stationed there apparently for the purpose, one of whom instantly started off and carried the seed all the way back again to the upper nest. This I saw repeated several times. After March 4 I never saw any acts of hostility between these nests, though the robbed nest was not abandoned. In another case of the same kind, how- ever, where the struggle lasted thirty-one days, the robbed nest was at length completely abandoned, and on opening it I found all the granaries empty with one single exception, and this one was pierced by tlie matted roots of grasses and other plants, and must therefore have been long neglected by the ants. Strangely enough, not one of the seeds in this de- serted granary showed traces of germination. No doubt some very pressing need is the cause of these sj^stematic raids in searcli of accumulations of seeds, and there can be little doubt that the require- ments of distinct colonies of ants of the same species are often different even at the same season and date. Thus these warring colonies of ants were active on many days when the majority of the nests were com- pletely closed ; and I have even seen these robbers staggering along, enfeebled by the cold, and in wind and rain, when all other ants were safe below ground. HARVESTING ANTS, 41 It may be that unusual exertions are necessitated by some exceptional demands made by the condition of the larvae of the v/inge(i male and female ants, and I have observed that these latter appear at very various periods. Thus I have seen winged males and females in the nests of Barbara on November 10, December 6, Februar}^ 2, and March 10 ; and in those of structor on February 23, 29, March 13, and April G. Though structor and fjartjara make seed collecting the business of their lives, they will, at least in times of scarcity, eagerlj^ devour animal food if it happen to fall in their way, and in the harvesting trains a few ants may occasionally be seen carrying small dead in- sects and the like. Once I threw a dead grasshopper down close to a nest of barbara ; it was immediately seized upon, and — after strenuous efforts had been made to dismember it above ground, some ants straining back the lejjs and wins^s, while others rushed in to gnaw at the muscles wdiere the tension was greatest, — carried down below. On the following morning the wings of the grasshopper were to be seen on the rubbish heap in front of the nest. Dead house-flies and the larvse of bees or wasps were at times readily devoured by my captive ants {barbara). I have also seen large numbers of strudors engaged in picking the bones of a dead lizard, and was once a witness of the following singular contest between a soft-bodied, smooth, greyish caterpillar, exactly an inch in length, and two medium- sized barbara ants. The ants were mere pigmies in comparison of their pre}^, for as such I believe they regarded the caterpillar, but they gripped its soft body with set mandibles, showing the most savage determination not to loose their hold. "When I first detected the group the war was being 42 HAR VESTING A NTS. waged in a tuft of grass over one of the entrances to the ants' nest, and the caterpillar was striding along the leaves, or thrusting itself between the culms in the hope to shake off or brush away its little persecu- tors. lYom time to time the caterpillar would turn viciously round and endeavour to pluck away its assailants, but though it actually succeeded in strip- ping off by means of its forelegs and mouth five of the six legs of one of the ants which was within its reach, they never once released their hold. At length a chance movement of mine shook the grass leaf on which they were, and ants and cater- pillar rolled together down a steep and rocky slope to about four feet distant. They tumbled over and over several times, but still the ants gripped their prey as firmly as ever. The last endeavour of the giant victim was to rub off the ants b}^ burrowing into the soil, but on un- covering its retreat, I saw that their positions were still the same. After watching this struggle for twenty minutes, time failed me, and I returned home, carrying with me, however, the combatants ; and when on my return I opened the box in which they were im- prisoned, these bull-dog ants were clinging with man- dibles locked as firmly as ever, and now as I write, in death they are clinging still, drowned in a sea of spirits of wine. During the winter and spring I kept two colonies of harbara captive in the house, placed in separate glass jars, each of which might perhaps hold half a gallon. The former of these colonies was taken on December 18 ; but neither the queen ant nor larvae were found, though there probably were larva3 in some HA R VESTING ANTS. 43 unexplored part of the nest, and the ants were always restless and miserable, unceasingly trying to escape, and dying in large numbers. On February 12 I found that all these ants, though abundantl}^ supplied with seeds and all other kinds of food, were dead. Two other colonies of ants, however, which had been taken in a torpid state in the masses of earth which formed part of tlie original nest, were alive and well, though still torpid. The second captive colony, taken on December 28, with the wingless queen ant and quantities of larvae, formed a strong contrast with the previous one. Here the ants at once set to work upon the construction of galleries and safety places for the larvae below the even surface of garden mould on which I had placed them within the jar; for in this case I did not attempt to preserve any portion of their own nest. This was done at 3.80 p.m., and by 9 that evening I found the ants most busily at work, having in less than six hours excavated eight deep orifices leading to galleries below, and surrounded these orifices by crater-like heaps, made of the earth pellets which they had thrown out. I have observed some- what similar structures raised b}^ harhara after the nests have been closed on account of rain, -dMi^strucior frequently raises still more elaborate and distinct craters, such as those represented at Fig. B, Plate II., p. 22 (reduced one-halt). On the following morning the openings were ten in number, and tlie greatly increased heaps of ex- cavated earth showed that they must probably have been at work all night. The amount of work done in this short time was truly surprising, for it must 44 HARVESTING ANTS. be remembered tliat, eighteen hours before, the earth presented a perfectly level surface, and the larvae and ants, now housed below, found themselves prisoners in a strange place, bounded by glass walls, and with no exit possible. It seems to me that the ants displayed extra- ordinary intelligence in having thus at a moment's notice devised a j)lan by which the superabundant number of workers could be employed at one time without coming in one another's way. The soil contained in the jar was of course less than a tenth part of that comprised within the limits of an or- dinary nest, while the number of workers was pro- bably more than a third of the total number belonging to the colony. If therefore but one or two entrances had been pierced in the soil, the workers would have been for ever running against one another, and a great number could never have got below to help in the all-important task of preparing passages and chambers for the accommodation of the larvie. These numerous and funnel-shaped entrances admitted of the simultaneous descent and ascent of large numbers of ants, and the work progressed with proportionate rapidity. After a few days only three entrances, and eventually'' only one remained open. Yet for weeks this active work went on, and the ants brought up such quantities of earth from below that it became difficult to prevent them from choking up the bottle containing their water, which they repeatedly buried up to the neck. On January 10 the surface of the earth was raised from an inch and a half at its lowest, to three inches at its highest point above its original level, and this ])ulk of excavated earth .BA R VESTING ANTS. 45 represented the amount of space contained in their galleries and chambers constructed below. Tt was not, ho^vever, until nineteen daj's after their capture that the ants began to form systematic trains to carry down the seeds which I placed for them on the surface, and I suppose that they had required this time for the construction and consolidation of the granary chambers. From this time forward the ants came out repeatedly in greater or less force to gather in the various seeds with which I supplied them. Indeed, throughout the whole of their cap- tivity they seemed to be perfectly contented with their lot and free from disease, remarkably few ants dying or appearing feeble, and as far as the limited space would permit they reproduced most of the habits Avhich I had noted as belonging to them in a wild state, .such as the formation of a rubbish heap ; bringing out refuse materials, gnawed and empty seed-coats, the ends of radicles, and root fibres which had penetrated their nest, and laying sprouted seeds in the air to dry after having gnawed off the radicle in order to arrest their growth. I was also in this way able to see for myself much that I otherwise could not have seen. Thus I was able to watch the operation of removing roots which had pierced through their galleries, belonging to seedling plants growing on the surface, and wdiich was performed by two ants, one pulling at the free end of the root, and the other gnawing at its fibres where the strain was greatest, until at length it gave way. Again the habit of throwing sick and appa- rently dead ants into the water, the object of which was in part, I imagine, to be rid of them, and partly 46 HARVESTING ANTS. perhaps with a view to effecting a possible cure, for I have seen one ant carry anotlier down the twig which formed their path to the surface of the water, and, after dipping it in for a minute, carry it labo- riously up again, and lay it in the sun to dry and recover ; thirdly, the stripping off the ccats and husks of seed and grain swelling and on the point of sprouting, previous to eating it ; and finally, the actual eating of the contents of the seed. Most of these operations are usually performed below ground, and even in my captive nest it was but rarely that I could get a glimpse of their subterranean life, as they avoided the glass as much as possible, though it was carefully covered with flannel and black paper ; and it was only by having the nest constantly before me on my table, and thus becoming a witness of their operations day and night during four months, that I detected them in positions which permitted me to watch these actions of theirs. The ants were in the habit of coming out in num- bers of an evening to enjoy the warmth and light of my lamp, and it was on one of these occasions that I first observed them in the act of eating. I perceived that in the midst of the black mass of ants gathered together on the side of the glass jar one was holding up a white roundish mass about as big as a large pin's head. Having turned a stream of bright light passed through a condenser on this group, and being per- mitted by the ants to make free use of my pocket lens, I was able to see the details with great precision. The white mass appeared to be the floury portion of a grain of millet, and I could see that two or three ants at a time would scrape off" minute particles with their HARVESTING ANTS. 47 toothed mandibles, and take them into their mouths, re- peating the operation many times, before giving place to other ants, and often returning again. It certainly appeared to be a botidjide meal that they were making, and not merely an act performed for the benefit of the larvse, as when they detach crumbs from a piece of bread and carry tliem below into the nest. However, I must own that, though I subsequently dissected ants taken in tliis act, which I suppose to be that of eating, I was unable by the use of the iodine test to ^detect starch grains in their stomachs. Still it seems quite possible that this failure may have been due to my not having allowed the ants sufficient time to swallow their food, as I killed them almost immediately after disturbing them at their meal. After having twice observed the ants eating as above described, I made some experiments in feeding them myself. They immediately seized and set to work upon a minute ball of flour which I cut out from the centre of a grain of millet, taken from a heap in front of a nest of A. structor, which had begun to sprout and been deprived of its radicle and dried. A similar ball taken from a sprouting grain of millet, but the growth of which had not been arrested, was also partially eaten ; but the hard, dry flour taken from a grain of the same in its natural state, not moistened, was at once rejected and thrown on the rubbish heap. The fat, oily seed leaves of the hemp, however, were eagerly taken, though not softened by water, their peculiar texture allowing the ants to scrape off" f)articles, as in the case of the ball of flour of the sprouted millet. 4S HARVESTING ANTS. Under ordinary circumstances the hard shell of the hemp-seed, and the coats of most other small fruits, grain, and seeds, would prevent the ants from getting at the contents while dry, but in the earliest stage of sprouting the shell parts of itself, allowing the radicle to protrude, and then they find their opportunity. (See Figs. A, A 2, Plate VI., p. 35.) It has always been siipposed that ants, from the delicate nature of their mouth organs, were only able to lap up liquids or to swallow very soft animal tissues, and one of the great difficulties in the way of admit- ting that they might collect seeds for food, lay in the apparent impossibility of their eating such hard sub- stances. But it has generally been overlooked that not only are all seeds soft when moistened with water and ready to grow, but also that there are certain kinds of seeds the contents of which are naturally soft. The most important organs in an ant's mouth are shown in Fig. D 2, and D 3, in Plate I., p. 21 . D 3 re- presents one of the horny, toothed mandibles, which serve admirably for scraping off particles of flour from the seeds. Within these are the parts shown at D 2, where the outermost pieces are the maxillae and their four-jointed palpi or feelers, and the innermost piece the labium and its three-jointed palpi, between which the end of the delicate membranous tongue appears. I repeatedly placed leaves from the orange trees covered with cocci and aphides from rose-bushes and pine trees, all of which are eagerly sought by several other kinds of ants, in the captive nest, but the ants never looked twice at them, and this corresponds with the fact that I have never seen either strudor or HARVESTING ANTS. 49 harhara attending on or searching for apliides and the like. These captives took part of a small quantity of honey which I placed in the nest, but displayed no eagerness about it, and soon neglected and allowed it to be covered up with earth thrown out from the nest. Tlie ants work very frequently at night during the dark,* and this is the case in the wild as well as in the captive nests. A friend, at my request twice visited a nest of sfructor ants in the garden of an hotel at Mentone, when it was quite dark (in March, between seven and eight o'clock p.m.) and no moon, but the light of a candle showed that the workers, both large and small, were busily engaged in carrying into the nest seeds which had been purposely scattered in their neighbourhood. I have myself seen Pheidole mcgacephala similarly engaged at about nine p.m. on a warm night in April, when it was perfectly dark, not even the stars showing ; but in this case the ants were collecting from the weeds in the garden. On the same occasion I also observed long and active trains r)f Formica emargitiaia [a rather small dusky ant, with a yellow thorax], making for the orange-trees in search of cocci and apliides, just as if it were broad day. Before leaving Mentone, on May 1, T turned out this second captive nest, and found that the colony appeared perfectly healthy, and did not seem to have diminished materially in numbers. The queen ant and the larvse seemed to be in just the same state as when they were taken. The earth in the lower part of the jar was honeycombed with galleries, granaries, * This beara out the much-questioned assertion of Aristotle, though he only claimed that ants work "by night when the moon is at the full." — H'st. Anim., lib. ix. cap. xxvi. 50 HARVESTING ANTS. and cells, constructed quite as in the wild nests, but more crowded together. Tlie granaries were in many instances full of seeds, which, though very wet, [the surrounding soil being extremely moist on account of there being no drainage to carry off the water which I was obliged to sprinkle from time to time over the surface of the nest], still showed no trace of germination that I could detect. The ants were therefore able to exercise the same influence over these seeds, under the strange conditions of their captive state, that they do in their natural homes. The foregoing remarks, as has been stated above, refer for the most part to only one of the three kinds of harvesting ants which I have observed on the Eiviera — that is to say, to Af/a larhara, the jet- black ant. As far as the manner of collectino: and storinij the seed is concerned, all that has been said of Atta harhara applies with equal truth to A. stnictor. A. structor is, however, less frequently seen above ground from December to March than barhara, and is more frequently found in or near the streets and gardens of a town. The fourth species, on the other hand, the little Pheidole megacepliala, differs in several particulars. This ant appears to shun the daylight, and to be most active at night, when, in the warm weather at the end of April, it may frequently be seen carrying large quan- tities of seeds into its nest. I have rarely observed it at work in the daylight, so that my knowledge of its habits is but small. Nor have I succeeded in dis- covering its subterranean granaries, though I have opened several nests. Still, I believe that it is a HARVESTING ANTS. 51 true harvesting ant, and not merely a casual collector of" seeds. Of the habits o^ P//cidoIe paUldida, a very closely allied and similar species, but one less frequently met with, I cannot speak with certainty, though it is quite possible that it also may be a true harvester, in which case it would add a fifth species to this class. Both Pheidole megacephcda and Ph. paUidula appear to remain inactive, or nearly so, during the mouths from November to April, and it is probable that they are only to be seen in full activity during the sum- mer when I am not there to watch them. There can be little doubt that any naturalist who will take the pains to note the habits of ants on the shores of the Mediterranean through June, July, August, and September, might collect a most inte- resting series of observations on harvesting and other species, and add to, and perhaps modify, those which my limited opportunities have enabled me to make. There are three other ants* — namely, Formica emar- glnata, F.faaca, and Myrmica caspitmn, which may also occasionally be found carrying a few seeds, but this is tlie rare exception, as far as my experience goes, these species living on honey dew, sweet secretions, and animal matter, like the great majority of ants all over the world. I have never found seeds in the nests of any ants except those of Atta harbara and A. slrudor, though I have carefully searched for them in most of the nests of the sixteen species of ants whose habits I have watched. * For some details of the habits of the sixteen species of ants observed on the Riviera, see Appeudix A. E 2 62 HABVESTINO ANTS. There is every probability that these harvesting' ants will be found all round the shores of the Medi- terranean, but the only points at which I have posi- tively heard of the existence of the habit besides Men- tone, Cannes, and Marseilles, are Capri* and Algiers. I am indebted to Miss Forster for having, during a short visit to Algiers, devoted some time to watching the habits of the ants in a garden at that place. These observations were made in April last (IS 72), when the three following species were watched : — (1) Formica {Catcu/I/jpJiis) viatica, a large, long-legged, blackish ant, with orange-red and semi-transparent thorax, which never carried seeds, but lived on animal food, especially flies. (2) Formica {Tapinoma) nigerrima,\ a rather small dusky ant, which brought in some seeds to its nest, but principally " animal food, flies, small worms," &c., and which did not carry the hemp and canary seed strewed in their path, though on one occa- sion when Miss Forster scattered some split hemp seed, they eagerly fastened upon the contents, and ate some on the spot, while they transjDorted the greater part to their nest, and (3) Atta harbara, which, as on the Hiviera, was a true and most active harvester, and eagerly seized upon the hemp and canary seed when these were placed in its way. Recapitulation and Concluding Remarks. There are some points of interest suggesting open- ings for future observation, to which I will now allude, * Where a harvester, probably AUa harbara, has been observed by Mr. Buchanan White. See Appendix C. t Mr. Smith thinks that this ant is either F. nigerrima, of Nyhander, or a new species, but it was not possible for him to pronounce with absolute certainty as he had only two specimens of workers from which to judge. HARVESTING ANTS. 53 making at the same time a partial recapitulation of what has gone before. We have learned in the first place that the ancients had facts on their side when they said that the ant is one of the very few creatures which lays up supplies of food sufficient to last for months, or even perhaps, as Bochart says, for a whole year ; and though we cannot quite accept the statement that " there is no animal except men, mice, and ants, that stores its food,"* they were right in sajdng that the habit is a most singular and interesting one. It is probable, however, that the old writers may have fallen into the error of supposing that all ants were harvesters, though the truth appears to be, that even in hot climates, it is only a very small number of species that are so. The fact that certain ants in Southern Europe do store large quantities of sound seed in damp soil, and check their tendency to germinate, may be thought to favour the possibility of the exis- tence of those deeply hidden supplies of seed which, though they have never been detected, are popularly supposed to explain the sudden appearance of the crops of weeds on soil newly brought out from great depths. The argument may be stated thus : seeds remain for months undecayed, and still capable of germination, at depths varying from one to twenty inches below the surface of the soil in certain ants' nests, why should they not lie hidden for indefinite periods in ordinary soil? To answer this positively, experiments should be * Soiihian, quoted by Bochart in his Hierozoicon, ii. cap. xxi. p. 497. 54 HARVESTING ANTS. made* in order that we might learn whether these seeds can retain their vitality without sprouting in moist soil ; but the general belief is that under these conditions thej will do one of two things, they will either grow or rot. Be this as it may, one of the most curious points that we have learned about these ants, is that they know how to preserve seeds intact, even when within from one to three inches of the surface of the ground, that is to say, at the actual depth at which a gardener most frequently sows his seeds, though if these very seeds are taken out of the granary and sowed by hand, they will germinate in the ordinary way. It is possible that this may be in part due to the compact nature of the floors and ceilings of the granaries, these excluding air in some measure, though as moisture freely passes through them, and there are always two or three open galleries leading into tlie granaries, and which communicate directly with the open air, I can scarcely accept this explanation as complete. The seeds do occasionally sprout in tlie nest, though it is extremely rare to find instances of this, and then the ants nip off the little root, and carry each seed out into the air and sun, exactly as the old writers have described, and when the growth has been checked and the seed malted by exposure, the}^ fetch them in * In order to try the experiment fairly, seeds taken from ants' nests, or seeds of the same species as those which are habitually found in ants' nests, should be placed at different depths in the earth and examined after the lapse of six or eight mouths. Why it is that certain seeds resist the influences which destroy the vitality of other seeds of closely allied species is another and a very curious but coniplic£ited problem, the explanation of which may perhajjs lie in the different chemical 2)roperties of the seeds in question, in the more or less permeal'le character of their seed-coats, or their general texture. HA R VESTING A NTS. 55 again. It is in this condition that the ants like best to eat them, as I have proved by experiments among my captives. As the ants often travel some distance from their nest in search of food, they may certainly be said to be, in a limited sense, agents in the dispersal of seeds, for they not unfrequently drop seeds by the way, wliicli they fail to find again, and also among the refuse matter which forms the kitchen midden in front of their entrances, a few sound seeds are often present, and these in many instances grow up and form a little colony of stranger plants. This presence of seedlings foreign to the wild ground in which the nest is usually placed, is quite a feature where there are old established colonies oi Atta barbara, as is shown at Fig. A in Plate 1., p. 21, where young plants of fumitory, chickweed, cranesbill, Arabis ThaUana, &c., may be seen on or near the rubbish heap. It would be interesting to make a list of all these ant-imported plants, and I think it quite likely that, if a sufficiently large number of nests were visited, some seedlings of cultivated species might be found amongst them, for we have seen that garden plants are frequently put under contribution. One can imagine cases in which the ants during the lapse of long periods of time might pass the seeds of plants from colony to colony, until alter a journey of many stages, the descendants of the ant-borne seedlings might find themselves transported to places far removed from the original home of their immediate ancestors. It is a true cause, but at the same time it may be one which has, like many true causes, exceedingly small effects. One can scarcely look at the teeming population of an 5(i HARVESTING ANTS. ant's nest, without asking whether there are any checks to their increase, and if so, what these checks are. I know ver}^ little of what foreign enemies they may have, though I have occasionally seen them captured by lizards, Cicindela beetles, and spiders,* and it is well known that the females are eagerly sought for b}^ birds at the season when they are above ground, and about to found new colonies ; but I believe that ants are the ants' worst enemies, for fearful slaugh- ter and mutilation often result from the encounter of armies of the same race, but belonging to different nests. Harvesting ants have nothing to do, as far as I have been able to discover, with aphides, cocci, and the like, nor do they seek for any of those sweet secretions which are the staple food of the generality of ants ; they live, however, on very friendly terms with certain yellowish-white and satiny-coated " silver-fish " {Lepisma), which are found in the passages and chambers of the nests ; but what their relations are to these creatures and to certain beetles which have been found in the nests of Atta harhara in Spain and Syria is unknown. It is possible that by carefulty watching captive ants in company with these creatures under very favourable conditions, something further might be learned on this head. My captive ants constructed all their * I have seen tlie remains of ants at the bottom of the tube of trap-door sijitler nests, and watched a hunting sjiider, Lycosa, capture a large black ant [Formica xjubcscens) , by entangling it in threads, which it deftly spun about its limbs, while running rapidly r^und the struggling victim in a circle, and dodging out of the way of the ant's mandibles. In England one may frequently see ants caught in the spiders' webs among the rose- bushes, and Mr. Blackwall says, in his Spiders of Great Britain and Ireland, that Theridion riparium lives principally on ants. HARVESTING ANTS. 57 chambers, granaries, and almost all their galleries away from the glass, and in the interior of the earth, though I tried to tempt them to work in parts more accessible to sight by swathing the jar in flannel. There is much to be learned, I do not doubt, about the friends and enemies of harvesting ants ; and an- other great desideratum is further information as to the parts of the world in which they are found and what causes may be assigned for the limitation of the habit. What is the geographical distribution of the har- vesting species, and what the geographical distribution, of the habit? For instance, to quote Mr. F. Smith,* jitfa structor, though not " found in England, is scat- tered over a great part of Europe, having occurred in France, Italy, Germany, Austria, Dalmatia, and Swit- zerland ; it has also been found in Algeria" and Syria ; and A. harhara is almost as widely spread. May we then conclude that these species are harvesters wherever they are found, and that they store seed in Germany and Switzerland as freely as they do on the shores of the Mediterranean? If this be really so, then Huber, whose attention was specially directed to this point, and a host of laborious and scrupulous observers of the Continent, have had the very fact under their eyes, though they have been at considerable pains expressly to deny it. I cannot think that this is likely, but it is a matter Nvhich could easily be settled by those who travel or reside in Germany, Northern France, or Switzerland. It seems to me more probable, however, that tliey do store in the south, but not in the north ; for all the * Mr. F. Smith, On Some New Species of Ants from the Holy Laud, in Jouru. Linnean Soc, London, vol. vi. p. 35. 58 HARYESTIXG ANTS. difficulties wliicli attend the preservation of the seed in the granaries in the south would be greatly in- creased in the wet climates of Northern Europe, and there, moreover, the greater cold would render the ants torpid almost throughout the winter, when food would not be required. But the question is plainly an open one. We may also ask why it is that only a very few out of the many species of ants which inhabit the shores of the Mediterranean should possess this habit of collecting seeds, and differ so widely in their manner of living, from their neigh- bours ? If we wish to put ourselves in the way to answer these queries, the first thing we should do would be to examine and compare tlie structure of the digestive organs and parts of the mouth in harvesters and non- harvesters, with a view to seeing whether there may not be some capital difference here. These observations demand some skill in dissection and preparation, and in regretting that it has not been in my power to make them, I can only hope that some one more skilled than I am may undertake the subject. It seems probable, that in warmer latitudes there are many conditions which favour the rapid increase of ants, so that a given tract of country in Southern Europe, for example, must have on an average more colonies to support than a similar tract in the north, and that to meet this increase of population, it has therefore become necessary for these creatures to seek their subsistence from as many and as dissimilar sources as possible. The fierce conflicts over booty both between rival nests of the same and HA R VESTIXG A NTS. 59 of distinct species, tend to show tliat, even as things are, the}'" frequently have to fight for their food. Hitherto, as far as I have been able to learn, only nineteen true harvesting ants have been detected in the whole world, limiting this term to those species which make the collection of seeds the principal occu- pation of their outdoor lives, and are evidently in the main dependent upon this kind of food for subsis- tence. Now if we compare these nineteen species of ants* together a very curious fact forces itself upon our notice — namely, that all of them are closely related, so much so that not only do all belong to the same division of ants (the tribe MyrmicinecB), but that with one exception {Pseudou/t/rma) all would have been jDlaced by the great Fabricius in one genus, Atfa, and the one exception is not far removed from it. We must not forget, however, that, as has been stated, there are other ants which do occasionally collect seeds, and thus appear to show traces of this remarkable instinct ; but as far as I have yet seen, it is always possible to distinguish them readily from true harvesters. Still I think it verj^ likely that in hot climates the division between harvesters and non- harvesters may be bridged over by a complete chain of intermediates. Here two more questions suggest themselves for more complete future solution. (1) Do true harvesters which store seed in granaries ever * These 2ire Myrmica (Atta) harhata, from Texas and Mexico; (Ecohma {Atta) cephalotcs, fiom Brazil and Mexico ; CEcodoina (Atfa) proridens, from India ; (Ecodoma (Atta) dijusa, from India; Atta rufa, from India; Pheidole {^Atia) megacephala, from South France; Atta harbara, from South France, Capri, and Algiers ; Atta structor, from South France; and Fseiidomyrma ru/oni(/ra, from India. 60 HA R VESTING A NTS. attend upon aphides and seek for sweet secretions ? (2) Do occasional harvesters ever form granaries ? In any case the name of " the provident one " is only, I suspect, fully deserved by a limited number of ants, and ^Esop, in his well-known fable, might as properly have made the dialogue which ends in the recommendation to " dance in winter as he piped in summer," take place between two ants as between an ant and a grasshopper, as far at least as their respective foresight is concerned. Why it is that one ant should require stores of food in the winter of which other ants have no need, is one of the many problems which only patient watching and careful comparison and experiment can help us to solve. There are not wanting those among the many winter visitors of the south who have time in abun- dance or superabundance at their disposal, and might help to clear up these and many other mysteries, and to them I would strongly recommend the study of the habits of plants and animals as a pastime, if nothing more. The way is open : it is not difficult to follow, and it leads to very pleasant places. APPENDIX. The following are the species of ants which I have observed on the Riviera, and principally at Mentone ; the actual locality where my notes were taken being given in every case. Family Formicidce.* Tribe Formicinece. — Petiole (or stalk which unites the thorax and the abdomen) of one joint, and furnished with a single vertical scale, abdomen not contracted. (1) Formica fusca, Linn. — A rather large ant (.3 J to 4| lines long), of a blackish ash colour, with a satiny sheen on the upper half of the abdomen. Smells of formic acid when crushed. Lives upon sweet secretions and animal matter, and occasionally carries a very few seeds into its nest, which is made in the ground. (Mentone.) (2) F. emarginata, Latr. — Of medium size (2^ lines), brownish, with yellow thorax. Has a strong smell of honey when crushed. Lives principally upon sweet secretions, but occasionally carries a very few seeds also. Nest in the ground. (Mentone.) (3) F. (Camponotus) cruentata, Lat. — Large (5 to 6 lines), dusky brown, with orange red on legs and abdomen. Strong smell of formic acid. Lives on sweet secretions and animal * Ants have been divided iuto three tribes, the two first of which, Formicinece and Ponerinece, are distinguished by the latter having a contrac- tion in the abdomen not found in the former, and both are separated from the third tribe, Myrmicinece by having but a single scale on the petiole, while in Mi/rmicinece there are always two nodes or protuberances on the petiole. It is important to remember the difference between the first and the last named tribes, as we shall find that all the true harvesters belong to Mijrmicinece. I have not seen any of the representatives of the second tribe in the south. 62 HARVESTING ANTS. matter, and has never been seen by me carrying seeds. Nest in the ground. (Mentone and Cannes.) (4) F. (Camponotus) marginata, Latr. — Large (i to 5^ lines), black. Has no perceptible smell even when crushed. Lives principally on sweet secretions, and does not bring in seeds to its nest, which is made in the ground. I have seen this ant at Cannes ascending the cork oaks in search of certain cocci which resemble black and shining berries rather larger than a pea, and which exude sweet secretions. (Mentone and Cannes.) (5) Formica cursor, Fonscol. — A rather large but slender ant (8 to 4 lines long), nearly black, with a faint bronzy hue, legs very long. Smell not noted. Runs very swiftly, and is hard to catch ; feeds on sweet secretions, and does not carry seeds. Nest in ground. (Cannes.) (6) F. (species undetermined). — A large ant (5 to 6^ lines), black brown with yellow thorax and legs. In shape resembles F. onarginata. Strong smell of formic acid. Habits not observed. Nest found under a stone in a pine wood. (Cannes.) (7) F. (species undetermined). — A rather large ant (3 to 4^ lines), resembling F.fusca, but having the thorax yellow. Strong smell of formic acid. Feeds on sweet secretions, and does not carry seeds. Nest in ground. (Cannes.) (8) Forraicob {Tapinomo) erratlca, Latr. — Rather small (2 lines), nearly black. Has a strong and most disagreeable smell, something like rancid oil, which is emitted if the nest is disturbed or the insect crushed. Lives upon sweet secre- tions and animal matter, but rarely if ever carries seeds, and paj s no attention to them if placed in its path- It nests in the ground, and forms superficial covered ways, roofed in with a thin crust of earth and vegetable fibres cemented together. (Mentone, Cannes.) Tribe Myrmicinece. Petiole two jointed, furnished with two nodes (protuberances). (9) Crematogaster {Myrmica) scutellaris, Oliv. — Of medium size (3| to 4 lines), nearly black, with yellowish red head. Disagreeable smell like rancid oil when crushed. Erects the abdomen when excited, and runs about with it turned up at right angles to the body. Lives on sweet secre- APPENDIX. 63 tions, and does not carry seeds. When dissecting the abdo- men of this ant, I noticed that in freshly killed specimens a drop of poison appears at the extremity of the sting, which if brushed away will form again several times in succession. Nest in the bark and wood of sick or decayed trees. (Men- tone and Cannes.) (10) C. sordidulus, Mayr. — Very small (1| to 2 lines), reseml)les C. scutellaris, but is uniformly black brown. No perceptible smell. Lives on sweet secretions, and may fre- quently be seen inside flowers. Nest in earth. Behaves like C. scutellaris when excited. (Mentone and Cannes.) (11) Myrmica ccespitwrn, Latr. — Small (2 lines), brown. Faint smell like peat smoke. Feeds on animal food and sweet secretions, and may occasionally be seen collecting and carrying in seeds. Nest in the ground. (Mentone and Cannes.) (12) Pheidole {Atta or Myrmica) megacei:)hala. — Very small (1|- to 2 lines), yellow, the larger workers having enormous heads. Smell very peculiar, and a trifle like aniseed when crushed. Appears to be a true harvester, and not to seek for sweet secretions. Nest in ground. (Mentone and Cannes.) (18) Ph. {Atta or Myrmica) pallidula. — Very small (1^ lines), pale yellow, closely resembles Ph. megacephala, but is paler and more transparent, and the larger workers have less disproportionate heads. Smell not noted. Habits not fully observed. Nest in ground. (Mentone.) (14) Atta {Aphenogaster or Myrmica) structor. — E,ather large (2 to 4 lines), of a claret brown. No smell when crushed. A true harvester, and does not appear to seek for sweet secretions, though it will occasionally take animal food. Nest in ground or under stones. (Mentone and Cannes.) (15) Atta. {Aph. or Myrmica) harhara. — Rather large (2 to 4 lines), jet black. No smell when crushed. Habits of strudor. Nest in earth, and more frequently in uncultivated ground. I have twice seen a few ants coloured like structor in colonies of harhara. (Mentone, Cannes, and Marseilles.) (16) Atta {Aph. or Myrmica) harhara var. — A large ant (3 to 6 lines). The larger workers black, witli red or mahogany- coloured heads, the smaller most frequently black, and like those of Atta harhara, of wliich this is probably only a 64 HARVEST FNG ANTS. variety. It differs however in its smell, which, when the body is crushed, resembles that of Pheidole Tnegacephala, and is something like aniseed. Habits of structor and barhara. Nest in earth. On one occasion I opened a large nest at Cannes, where the colony was composed in about equal parts of ants which in colour and appearance might be said to represent the three forms, structor, barbara, and the red- headed variety of the latter. There were also a few ants with pale yellowish brown heads. (Mentone and Cannes.) B. The following Indian species are described by the late Dr. Jerdon as harvesters, in the Madras Journal Lit. and Sc. 1851 :— (p. 45). Atta rufa. — "Its favourite food is dead insects and other matter, but it also can-ies off seeds like the (Eco- dorna, chaff," &c. &c. (p. 46). (Ecodoma providens. — " Their common food I suspect to be animal matter, dead insects, &c. &c., which at all events they take readily, but they also carry off large quantities of seeds of various kinds, especially light grass seeds, and more especially garden seeds, as every gardener knows to his cost. They will take off cabbage, celery, radish, carrot, and tomato seeds, and in some gardens, unless the pots in which they are sown be suspended or otherwise protected, the whole of the seeds sown will be re- moved in one night. I have also had many packets of seeds (especially lettuce) in my room completely emptied before I was aware that the ants had discovered them. I do not know, however, if they eat them or feed their larvae on them, though for what other purpose they carry them otf I cannot divine. I have often observed them bring the seeds outside their holes, as recorded by Colonel Sykes, and this I think generally at the close of the rainy season ; but in some cases I had reason to beheve that it was merely the husks, of which I have seen quite heaps, and that the ants did not take them back to their nests. If any of the forementioned seeds be sown out at once in a bed, most likely in the morning the APPENDIX. 65 surface of the whole spot will be found covered over with little ridges, the works of these creatures, and the few seeds that perhaps remain, dug all round, and being carried off sometimes above ground, at other times under ground. Their galleries and subterranean passages are often very extensive, and it is no easy matter to dig down to their nest to see what becomes of the seeds." (Ecodoma diffusa has the same habits as (E. providens. Lieut.-Col. Sykes, Descriptions of New Indian Ants in Trans. Ent. Soc. Lond., i. 103 (1836). AttOj providens, Sykes. '' In illustration of the habits of this species of ant, I shall give the following extract from my diary: — 'Poena, June 19, 1829. In my morning walk I observed more than a score of little heaps of grass-seeds (Panicum) in several places on uncultivated land near the parade-ground ; each heap contained about a handful. On examination, I found they were raised by the above species of ant, hundreds of which were employed in bringing up the seeds to the surface from a store below ; the grain had probably got wet at the setting in of the monsoon, and the ants had taken advantage of the first sunn}'^ day to bring it up to dry. The store must have been laid up from the time of the ripening of the grass-seeds in January and February. As I was aware this fact militated against the observations of entomologists in Europe, I was careful not to deceive myself by confounding the seeds of a Panicum with the pupae of the insect. Each ant was charged with a single seed, but as it was too weighty for many of them, and as the strongest had some difficulty in scaling the perpendicular sides of the cylin- drical hole leading to the nest below, many were the falls of the weaker ants with their burdens from near the summit to the bottom. I observed they never relaxed their hold, and with a perseverance affording a useful lesson to humanity, steadily recommenced the ascent after each successive tum- ble, nor halted in their labour until they had crowned the summit, and lodged their burden on the common heap.' " (p. lOi). " On the 13th of October of the same year, after the closing thunderstorms of the monsoon, I found this species in various places similarly employed as they F 66 HARVESTING ANTS. had been in June preceding ; one heap contained a double handful of grass-seeds. It is probable that the Atta pro- videns is a field species of ant, as I have not observed it in the houses." c. After the appearance of a brief notice of a communication •which I sent in the winter of 1871-72 to the London Ento- mological Society, announcing the fact that certain ants harvest seeds in a systematic way at Mentone, two papers were pub- lished, in which confirmatory evidence of the existence of the habit in other parts of the world was set forth — one by Mr. Buchanan White, and the other by the late Mr. Home. Mr. Home's account of his observations was published in Eardwicke's Science Gossip, No. 89, p. 109 (for May 1, 1872), and runs as follows :* — " My notes carry me to the far East, where I have often watched this most interesting class of insects, and briefly recorded my observations — unfortunately cut short by illness, and the necessity of return to Europe, which must be my apology for their want of completeness. " But before transcribing, I would remind my general reader that all ants may be seen carrying off food to their nests for present consumption, and that this food consists of a great variety of substances. This is disposed of inside the said nest, being often masticated, and the juice extracted by the workers, and then given in an inspissated form from their mouths to the young grubs, which are in general tended by their nurses with the greatest care. It is indeed very curious to watch this feeding process ; but to proceed. " Under date Nov. 7th, 1866, I find in my natural history note-book as follows : — Mainpuri. This morning as I was walking across the ' Oosur/ or waste plain, where it was very sandy, being cut into small ravines, and clothed only here and there with fine grass disposed in clumps, thus forming little hillocks of sand blown by the wind, and arrested in its course * I omit the prelimiuary portion, in which my observations are erroneously stated to have been made at Nismes and Capri. APPENDIX. 67 by the grass, I came across a long line of ants, travelling four deep, some coming empty, and others laden each with one grass-seed, on their way home. "I followed up the procession to the nest, which was sub- terranean, and at the mouth of which on the level plain there was no trace of elevation caused by the soil brought up from below, owing to the habit of these ants of taking each grain of sand to some distance along their road, and depositing it on one side or the other. " There may have been five or six entrances to the nest, in and out of which a prodigious number of ants were passing, the species of which has been described by Dr. Jerdon. They were of a medium size, shortish bodies, and of a reddish-brown colour — Fseudomyrma rufo-nigra, Jerdon. Around the mouth of the nest, forming a circle of perhaps eighteen inches in diameter, was a space beaten flat, and kept clear by these said ants, from which radiated in every direction thirteen roads, each about four inches in width for about thirty to forty yards, when they branched off and became narrower, being ultimately lost amongst the grass roots. These paths were fairly straight; they did not cut through elevations, but went round them. " From a careful examination it appeared that they had been cleared of all obstacles, such as small stones, twigs, &c., but that their smoothness resulted only from the tread of countless feet. " The bearers of burdens took the seeds into the nest, which I did not dig up, and certainly stored them there, after having prepared them, probably by the removal of a portion of the outer husk. Of these husks there were large collections near the entrances to the nest, all carefully set aside by the ants. " In times of famine, I am told, not only are the nests rifled of their grass-seed stores, but these heaps of apparent husks are collected and ground with other grain to eke out a subsistence. "This kind of grain has a name, 'Jurroon,' derived from * Jharna,' to sweep, literally sweepings. I much regret that I have not preserved specimens of this ' Jurroon,' for it is very unlikely that the ants after taking it to their granary, should again throw it out, and yet, if grainless, what benefit F 2 68 HARVESTING ANTS. could there be in eating it? The season of the year when I observed them (November) is the beginning of the cold ■weather, and no rain would probably fall (excepting a little at Christmas) till next May or June. Later on seed would be rare ; and how the nest fares at a time when floods of water often pass over the plain I cannot conceive. " It is clear that some escape, and we know with what pro- digious rapidity these colonies increase. But these jottings have been recorded merely to show how this species of ant store grain against a time of scarcity, and fully bear out the statement in the text with which I commenced this paper." The following are Dr. Buchanan White's notes, alluded to above, published in the Transactions of the Entomological Society (London, 1872) part i.. Proceedings, p. v. : — "Capri, June 3, ]866. In the afternoon to the Punta Tragara, where a colony of ants afforded us much amusement. These little insects had a regular road, made by cutting away the grass and other plants in their way. This road was about one inch and a half wide, and several yards long, and led to a clump of plants in seed. Along this road a long train of ants were perpetually travelling to the nest {or formicarium), bearing with them pods of leguminous plants, seeds of grass and of Composites [Chrysanth. segetwni), &c. " The perseverance with which a single ant would tug and draw a pod four times his own length was very interesting ; sometimes three or four ants would unite in carrying one burden. Near the formicarium was a great mass of debris, consisting of empty pods, twigs, emptied snail shells, &c., cast out by the ants. The seeds appeared to be stored inside the nest, as in one that I opened the other day I found a large collection. The species was a black ant ; the formicarium was underground." D. 0)1 Collecting and Bxamining Ants. There are very few branches of natural history which might be more easily followed b}' a traveller, or one who fears to APPENDIX. 69 encumber himself with bulky collectious difficult to transport from place to place, than the study of ants. The whole European ant fauna might be adequately represented by specimens preserved in spirit of wine and packed in the compass of a hat-box. In taking specimens of ants it is important never to put the representatives of more than one nest in each bottle, but then in most cases a sufficient number may be placed in a single bottle of the size used for containing the smaller homoeopathic globules. If possible the winged male and female ants, as well as the wingless workers, should be secured. The ants die very quickly in pure spirit of wine, and they can afterwards, even after the lapse of months or more, be pinned out in the cabinet after having been washed in warm water. In examining the mouth organs of an ant in order to determine by the aid of books to what genus it belongs, it is best to relax the parts by first washing away the spirit of wine, and then leaving the specimen for a day or more in a stopper bottle partly filled with finely chopped laurel leaves. It is probable that a drop or two of prussic acid on a bit of sponge might act as efiectually in rendering the tissues pliable, A compound microscope is necessary for the final examina- tion of the joints of the labial and maxillary palpi (see Fig. D 2, Plate I., p. 21); but the neuration of the wing (D 1, Plate I.), another very important character, is easily detected with a good pocket-lens. The works which may most usefully be consulted are, for France, M. Ny lander's Forinicides de France et d'Algerie, published in vol. v. of the fourth series of the Zoological Division of the Annales des Sciences Naturelles ; for Eng- land, Mr. F. ^unih! ^Catalogue of British Fossorial Hymenop- tera (L856) ; and for a more general review of the species in the world at large, Mr. F. Smith's Catalogue of Hymenop- terous Insects in the Collection of the British Museum, Part vi., Formicidce (1858), and M. Mayr's Beitrdge zur Kenntniss der Ameisen, published in the Verhandlungen des Zoologisch-botanischen Vereines in Wien, iii. 1853. Ahhandlangeii (p. lOl). PART II. TRAP-DOOR SPIDERS. PART II. TRAP-DOOR SPIDERS. It is now one hundred and sixteen years since Patrick Browne gave an illustration in his Civil and Natural History of Jamaica"^ of the nest of a trap-door spider, the first record of the kind with which I am acquainted. Seven years later the careful observa- tions of the Abbe Sauvages appeared,! in which he gave a very good description of the nests of tlie " araignee mafonne " {Nemesia ccBmentaria) , which he discovered near Montpellier, likening them to little rabbit burrows lined with silk and closed by a tightly- fitting moveable door. In 1778 and 1794 Eossi | published an interesting account of the nest and habits of a trap-door spider which he had observed in Corsica and near Pisa ; and from that time ujd to the present day the curious dwellings of these creatures, many species of which have been discovered in warm climates, have continued to attract the attention of naturalists. Very little, however, has been added to our know- ledge of the life-history of these remarkable archi- * p. 420, tab. 44, fig. 3 a. This work was published in London in 175G. f In Histoire de I'Acad. Eoyale des Sciences (Paris 1763), p. 26-30. % Rossi (P.), Osservazione Insettologische (Memorie di Matematica e Fisica della Societa Italiana, vol. iv. (1778), and Fauna Etrusca, vol.ii. (17!'4). 74 TRAPDOOR SPIDERS. tects for several years past, and, indeed, I think it may be safely asserted that the study of the habits and interdependence of the members of the animate world has not, during the last fifty years, made any- thing like a corresponding progress to that which may be seen in classification and description. The microscope has led many who, a century ago, would have found their chief delight in observing those points in the habits and external characters of living creatures which the naked eye could readily seize upon, to look much closer, to anatomize and describe in detail every organism, great and small, and to examine every tissue and cell. It is, however, to the materials now being amassed by these modern " cabinet naturalists" that recourse must be had if we wish to form a true comprehension of the functions and habits of living things. They must tell us, for example, what instruments, tactile and visual, an animal possesses if we wish to under- stand how it constructs a particular fabric, so that the " field naturalist" will have to apply to his brother of the " cabinet " before he can turn his observations to good account. Still, the fact remains that the habits of plants and animals afibrd many openings for careful investiga- tion, and such as are especially within the reach of those lovers of nature who have ample time at their disposal, and the opportunity to spend it in a warm climate where life abounds, and is never wholly checked even in the depth of winter. It seems strange to think that collectors so frequently take creatures out of wonderfully constructed nests and yet never observe, or at any rate never describe, the TRAP-DOOR SPIDERS. 75 structure of these fabrics. Thus, for example, the dwellings of only eight out of the thirty-six species of trap-door spider stated by Prof. Ausserer* to be- long to the Mediterranean region are known in books, those of the remaining twenty-eight being, as far as I have been able to learn, yet to be discovered. This is the more strange as from the nocturnal habits of these creatures it is almost always necessary to dig them out of their nests ; indeed it is more than pro- bable that if all the dwellings which have been de- stroyed had been described, the following pages would never have appeared. Before proceeding to pass briefly in review what has been written on the subject of trap-door spiders, it will be w^ell to take one glance at the relation which these spiders bear to their fellows. The great order of spiders {Aranece) has recently! been divided into seven sub-orders, the fourth of which, Territelaria, includes all the trap-door spiders, and some others which do not construct trap-doors. This sub-order corresponds with that which was formerly called Mygalidce, but this name, as well as that of Mygahy originally given to all trap-door spiders, has been abandoned because this latter name had previously been applied to a genus of Mammals, and it was feared that confusion might arise. The Terriielarice [or underground weavers] are dis- tinguished from all other spiders by the position of * Prof. Ausserer (Anton.), Beitriige zurKenntniss der Arachniden Familie der Territelarise (Mygalidae), iu Verhandlungen der k.k. Zool. But. GescU- schaft in Wien. Jahrg. 1871, Band xxi. t Thorell, On European Spiders, in Nova Acta Reg. Soc. Scient. Upsa- liensis, ser. iii. vol. vii. fasc. 1 and 2 (1869-70). 76 TRAP-DOOR SPIDERS. their falces,* which have the fang directed downwards, and move vertically parallel to one another. Thus when a victim is seized by one of the Territelarits it receives a downward blow, while other sjjiders strike sideways, the falces moving in a horizontal or oblique direction. With very few excej^tions this sub-order may also be known by the presence of four blotches of paler colour at the base of the abdomen underneath, indicating the position of four air-sacs, almost all, or indeed perhaps all, other spiders having but two. Certain species of TerritelaricB are the only spiders known to construct nests closed with a door, and these creatures must be admitted to rank among the first of Nature's handicraftsmen and inventors. The geometrical webs of many common spiders are very beautiful structures, but these are for the most part only snares for prey, and not permanent dwellings, although the cocoons in which the eggs are placed are often most ingeniously contrived. Thus in the south we may sometimes find an inverted bal- loon of strong silk about an inch long attached to heath and other bushes, which, if examined during the winter, will be found to contain in its centre a case enclosing a mass of eggs about one-third the bulk of the entire cocoon. This inner case is shaped ex- actly like the outer, and both have a circular silk lid carefully closed, and the space between the two is filled with a dense mass of golden-brown silk, which acts no doubt as an excellent non-conductor. This cocoon is the work of Epeira fasclata, a species ap- parently only found in southern Europe. * Sometimes called mandibles. One of these is rei^resented, enlarged, at Fig, A 7. in Plate VII., p. 88. TRAP-DOOR SPIDERS. 77 Other spiders again, such as Theridion* suspend by a long and delicate cord of silk a minute balloon, scarcely larger than a seed-pearl, containing their eggs, which sways to and fro in the lightest breath of air. But admirable as these cocoons and geo- metrical snares are, the homes of these and of spiders generally do not exhibit much contrivance or ingenuity, or cannot at any rate be ranked in the same category as those of the trap-door spider. But it may be asked, why should we admire the one more than the other, since it is clear that the most squalid and mean- looking nest exactly serves the purpose of its occu- pant, whether for shelter or defence, and in many cases a spider might even say with truth that as for her home it would not be so safe if it were not so dirty. But the answer is simple : the trap-door spider's dwelling is to that of common spiders what the Mont Cenis tunnel is to other tunnels, and some- thing besides. What delights us is to see how by clever con- trivance and great perseverance new and multiplied difficulties have been overcome, and dangers avoided, and the interest aroused is exactly proportionate to the amount of these difficulties and dangers. It is hoped that the following pages and their ac- companying illustrations will vindicate the claims of these spiders to the marked attention and admiration which is here asserted to be their due as architects and engineers. There is but one British or North European re23re- sentative of the Territelarice — namely, Atypiis piceus (or * Theridion variegatum (Bl.). Ero tuberculata, Koch. 78 TRAP-DOOR SPIDERS. Suheri)* and this creature does not appear to deserve the name of trap -door spider, for in three nests which M. H. Lucas kindly showed me, pre- served at the Jardin des Plantes at Paris, the mouth of the tube was destitute of any covering. I gathered from what I saw, and from what M. Lucas told me, that these nests [wdiich he had taken in the neighbourhood of Paris], consist of a silk tube from eight to ten inches long, of which about one inch only at the lower extremity penetrates the earth, the remainder being carried upwards in an irregular and sinuous course among the stems and leaves of small plants and grasses to which it is attached. When the tube is removed from these supports it collapses, and appears like a rather coarsely woven ribbon-shaped strip of silk.f Four types of trap-door nest, properly so called, may now be distinguished in the world at large, and these are represented diagrammatically in the following "woodcut. \ * Unless it should prove, as Prof. Ausserer suggests, tliat the British .^ ^?/pMs is distinct from the Continental, when it would bear the name of Atypus Blachwallii. (Ausserer, 1. c. p. 17). 1 1 have never been able to meet with an English specimen of the nest of A typus ; but it would appear from the descriptions that the English differ from the French nests in being subterranean, and in having the mouth of the tube concealed by a loose flap of silk. Mr. Blackwall saj's : [Spiders of Great Britain and Ireland, part i. p. 14] "Dr. Leech has taken specimens of A typus Suheri in ths vicinity of London and Exeter. It excavates, in humid situations, a subterraneous gallery, which is at first horizontal, but inclines downwards towards its termination. In this gallery it spins a tube of white silk, of a compact texture, about half inch in diameter Part of the tube hangs outside of the aperture to jirotect the entrance." It would be interesting to learn whether these differences permanently distinguish the English from the French nests, and if so, whether the spiders which construct them are not, as Prof. Ausserer is inclined to believe, dis- tinct also. t Where all the figures, except C 1, and D 1, which are of the natural size, are reduced to about one-third of the actual size in large specimens. TRAP-DOOR SPIDERS. 79 Of these two only (A and B) were known up to the present time, the construction of which is much simpler than that of the two new types (C and D), which I have hitherto only found at Mentone and Cannes.* It will be seen at a glance that A and B have but * It must not be supposed that I have a sole or prior claim to what may prove to be new and of interest in the following observations on the Trap- door Spiders of the Riviera. This prioritj- belongs to the Hon. Mrs. Richard Boyle, to whom I owe it that I ever took np the subject. It was, thanks to her guidance, that I first became acquainted with these marvellously-con- cealed nests in their native haunts, and to her active help that I finally arrived at a comprehension of the different types of structure which they present. 80 TRAP-DOOR SPIDERS. one door, while C and D have two, these latter having a surface door, and also another door a short way under ground. All the nests consist of a tube excavated in the earth to a greater or less depth, unbranched in all but D, and in every case lined with silk, this lining being continuous with the lining of the door or doors of which it forms the hinge, I have found it convenient to distinguish these four types of nests by the following names : — A, the single door cork nest, or shortly tlie cork nest ; B, the single door wafer nest; C, the double door unbranched nest ; and D, the double door branched nest. The type B has only been found in the West India Islands, and is chiefly distinguished from A by hav- ing a thin and wafer-like door, wholly constructed of silk, Avithout admixture of earth, lying on rather than fitting into the aperture of the tube ; while in A the door is much thicker, made of layers of earth and silk, and so contrived that it tightly closes the mouth of the tube, which is bevelled to receive it, much as a cork closes the neck of a bottle.* The West Indian nests are of a much tougher and coarser texture than those which I have seen in Europe, and vary somewhat in the shape of their tube, which is curved or straight, and sometimes has near its lower extremity a short spur-shaped enlarge- ment, giving to the whole a ludicrous resemblance to a stocking, of which this spur is the heel. Mr. Gosse,t in his Naturalist' s Sojourn in Jamaica, * Nests belonging to the type A, are represented in Plates VII., p. 88, and VIII., p. 94. t Gosse (P. H.), Naturalist's Sojourn in Jamaica (1851), jx 115-118. See also for another description of the same nest Latreille's Vues Generales sur les Araueides, in the Nouv. Ann. du Museum (Paris, 1832), torn. i. p. 73-4. TRAP-DOOR SPIDERS. 81 lias given an admirable description of one of these single door wafer nests, the work of Cteniza nidulans, which I cannot do better than quote : — ■ The nest is " cylindrical, or nearly so, from four to ten inches deep, and about one inch in diameter ; the bottom is rounded ; and the top, which is at the surface of the soil, is closed very accurately with a circular lid. They are not all equally finished, some being much more compact, and having the lid more closely fitted than others. Some have irregular bulgings, and ragged laminated offsets on the outer surface ; but all are smooth and silky on the inside The mouth of the tube, and the parts near it, are very strong ; the walls here often having a thickness of from one- eighth to one-fourth of an inch ; but the lower parts are much thinner. The lid is continuous with the tube for about a third of its circumference, and this part may be called the hinge, though it presents no structure peculiar to itself ; it is simply bent at a right angle, as is manifest, if a nest be cut longitudinally through with scissors, the incision passing through the midst of the lid. The mode of construction, I judge, from examination of many nests, to be this. The spider digs a cylindrical hole in the moist earth, with her jointed fangs or mandibles, carrying out the fragments as they are dislodged. When the excavation has proceeded a little way, she begins to spin the lining which forms the dwelling. I conclude thus, because nests are occasionally found a few inches in length, with the lid and upper |)art perfect, but without any bottom, these being evidently in the course of formation. I suppose that she weaves her silk at first in unconnected patches, against the earthy sides. 82 TRAP- DOOR SPIDERS. perhaps where the mould is liable to fall in ; and thus I account for the loose rough laminae of silk tliat are always found projectin<^ from the outer surl'ace. These are overlaid with other patches more and more ex- tensive, until the whole interior walls are covered ; after which the silk is spun evenly and continuously all round the interior, in successive layers of very dense texture, though thin. Under the microscope, with a power of 220 diameters, these layers are resolved into threads laid across each other and intertwined in a very irregular manner ; some are simple, varying from y-oVo- to ^ToVo of an inch in diameter, and others are compound, several threads, in one part separate, being united into one of greater thickness which cannot then be resolved .... The mouth of the tube is com- monly dilated a little, so as to form a slightl}^ recurved brim or lip ; and the lid is sometimes a little convex internally, so as to fall more accurately into the mouth and close it. The thickening of the hinge by additional laj^ers is, I think, accidental only, as, out of the many specimens that I have examined, only one or two had such a structure. In the neatest examples, the lid is of equal thickness throughout its extent, agreeing also with the walls for the first few inches of their depth," Mr. Gosse says that he possesses one specimen of peculiar compactness, which differs from all the others that he has examined in having " a row of minute holes, such as might be made by a very fine needle, pierced around the free edge of the lid, and a double row of similar ones just within the margin of the tube. There are about fifteen or sixteen punctures in each series, and they penetrate through the whole TRAP-DOOR SPIDERS. 83 substance, the li^lit being clearly seen through each hole. I do not think, as I have somewhere seen suggested, that they are intended to afford a hold for the spider's claws when she would keep her door shut against the efforts of an enemy, for what would be the use of having them in the tube close to the lid, so close that not the eighth of an inch intervenes between the surface of the lid and that of the tube, when the former is tightly closed ? I would suggest whether they may not be air-holes, for so tight is the fitting of the lid, and so compact the texture of the material, that I should suppose the interior would be imper- meable to air but for this contrivance."* " The spider that inhabits this nest is black, with the thorax of an exceedingly lustrous polish, its abdomen is full and round, its legs very short." Another form of this single door wafer nest is de- scribed by j\Ir. Sells,! in wdiich there is a hinge-like thickening of the silk lining of the tube about an inch below the actual hinge of the door, which it is sug- gested may serve to give additional elasticity. This was not found, however, in all the nests examined, and Mr. Sells conjectures that in newly constructed nests the lid may close sufficiently firmly without this contrivance, and that it is only added in older nests. Patrick Browne's figure, to which reference has been made above, represents a nest with two doors, one applied against the other, at the mouth of the tube. * I cannot myself think this explanation probable, and should still be inclined to consider these punctures to be the claw marks of the spider, as is the case in some European nests. t ]Mr. W. Sells. Notes respecting the Nest of Cteniza nididans, in Trans. Ent. Soc. ii. 207-210. G 2 84 TRAP-DOOR SPIDERS. and it has often been asked what this could possibly mean. Some have thought that the drawing was fanciful, others that it was made from an abnormal or injured nest. However, I believe that the drawing, though rude, is, in fact, not ver}^ incorrect, and shows a case of repair or enlargement of the nest, a subject to be treated of more fully further on. There is a specimen exhibited in the British Museum which in this respect very nearly corresponds with Browne's figure ; it is labelled " Nest of Trap-door Spider with two doors, from the spider having enlarged its abode. — Jamaica." Here one sees that the spider has prolonged its tube about half an inch beyond the original mouth of the nest, where it has constructed a new mouth and door, the old door standing straight up at the back of and behind the new one. I imagine that the explanation of this curious piece of cobbling may be somewhat as follows : — When the nest was in its original state and had but one door, this door became by some accident covered over with earth to about the depth of half an inch, and the in- mate was thus imprisoned. Then the sj)ider, being, like most other members of its order, very unwilling to abandon its home, determined to clear away the entrance to its nest, and to lengthen the tube so that it should reach up to the new level of the surftice of the earth If I am right, this should rather be called a lengthening than an enlargement of the tube. The nests of the cork type (A, p. 7t)) may usually be distinguished at a glance from those of the wafer type by the greater thickness of the door, and by its manner of shutting, but a nest from Morocco has been figured TRAP- DOOR SPIDERS. 85 and described by Prof. Westwood,* whicli seems in- termediate between the two. The door in this case may perhaps be considered as of the cork type, thougli it is very thin, for it does fit into the mouth of the tube, which is bevelled to receive it. These nests were forwarded with their living occu- pants {Cteniza \_Actinopiis] {Bclifcatorius) from Tangiers to Prof. West wood, who describes the nests as being " about four inches deep, slightly curved within, about three quarters of an inch in diameter, the valve at the mouth not being circular, but rather of an oval form, one side where the hinge is i^laced being straighter than the other. The valve is formed of a number of layers of coarse silk, in the upper layers of which are imbedded particles of the earth, so as to give the cover the exact appearance of the surrounding soil, the several successive layers causing it, when more closely inspected, to resemble a small flattened oyster- shell.^ The resemblance between this nest and that of the West Indian species is the more interesting as Prof. Westwood says that both belong to the same genus, {Cteniza or Actinojjus of different authors,) and are so closely allied as to present scarcely any important distinction but that of size. We shall find, however, on comparing the nests of these trap-door spiders and their occupants, that we cannot as yet make any rule as to the kind of nest which we may expect from a given spider. * Observations on the Species of Trap-door Spiders, in Trans, of Entomo- logical Soc, Loudon, 1841-3, vol. iii. p. 175. I wish to take the present opportunity of thanking Prof. Westwood for having afforded me special facilities for examining this and other specimens forming part of the very valuable collections under his care at Oxford. 86 TRAP-DOOR SPIDERS. It will be seen tliat species belonging to the same genus, and closely resembling one another, sometimes build dissimilar nests ; wliile others, belonging to different genera, and unlike in many important respects, construct almost identical nests. This is the more strange, because, if we examine the structure of the claws and palpi, they often seem to be specially adapted to serve as carding instruments and to play a very important part in the weaving of the silk linings of the nest ; and yet nests of the same type are occasionally produced by spiders in which these appendages are quite unlike, and dissimilar nests where the claws and palpi are to all appearance identical. Thus, for example, if the reader will examine the drawings of part of the foremost right foot of Cteniza fodieus, figs. A, 9 and 10, Plate VII., p. 88, with that of Nemesia C(jem.enfaria,^^^s. A, 9 and 10, Plate VIII., p. 94, both of which make nests of the cork type, he will see that in the former the last joint of the tarsus is armed along the inner side, with many moveable spines, and that each of the two curved terminal claws has only one very strong tooth near the base ; while the same joint of the latter {N.ccementaria) has no spines, and the claws have three minute comb-like teeth near the base. On the other hand, in the reverse case, where the structure of the same joint is very similar, the nests may be wholly unlike, as in Nemesia Eleanora, Plate XII., p. 106, and N. ccBmentaria, Plate VIII., where the nest of the former is of the double-door unbranched type, and that of the latter of the single-door cork type. ^ It is probable however that a fuller and closer comparison of, and a more exact acquaintance with the TRAP-DOOR SPIDERS. 87 several parts and their functions might show us that all spiders wliich spin similar webs are furnished with equivalent instruments, so that what one leg lacks another may possess in some shape or another; brushes of stiff hairs in one place, compensating for a toothed claw, or for a row of moveable spines in another.* It would be interesting, from this point of view, to draw all the parts which may be supposed to aid in the act of weaving, and so to contrast the corresponding limbs of different spiders, the nests of which are known, that one might see at a glance in what they differed and agreed. I have done this for the falces and the last joint of the foremost right foot of the four spiders figured in Plates VII., VIII., IX., and XII., but to make the case complete all the limbs should be re- presented in the same way. * Tlie claws are probably of first-rate importance in tliis respect and should be most carefully studied. M. Lucas has stated that the claws of Myijale Blondii, and M. ni