Rev. S, Haughton on the Cells of the Wasp and Bee. 415 EXPLANATION OF PLATE XI, Fig. \. Acantholeberis curcirostris (Miiller), 5 . Fig. 2. Anterior antenna of the same species. Fig. 3. Portion of the terminal half of the seta attached to the first joint of the lower branch of the posterior antennas; greatly magnified. Fig. 4. Portion of the terminal half of the seta attached to the second joint of the lower branch of the posterior antennae, greatly en- larged. Fig. 5. Abdominal claws. Fig. 6. Acantholeberis sordida (Lievin), ? . Fig. 7. Abdomen of the same species. Fig. 8. Setae from the ventral margin of the carapace. Fig. 9. Setae from the posteroventod angle of the carapace. XLV. — On the Form of the Cells made by various Wasps and hy the Honey Bee ; with an Appendix on the Origin of Species. By the Rev. Samuel Haughton*, Fellow of Trinity College, Dublin. The geometrical form affected by the cells of various kinds of wasps and bees has attracted the attention, and called forth the speculations, of naturalists and geometers from the earliest periods. By one class of writers the geometrical properties of these cells have been used as proofs, not so much of the skill and instinct of the insects as of the wisdom and intelligence of their Creator ; whde, by the opposite class of writers, these same geometrical properties of the cells are alleged as a suflficient cause for the production of the insects that make them, from the advantages which these forms of cells are supposed to possess over other forms — advantages said to be so important as to de- cide the battle of life in favour of the insects that adopt the geometrical plan of making their cells. I have for a long time felt convinced that both parties in this controversy are in error, as men generally are when they attempt to speculate on the reasons for the existence of things ; and that the properties of the cells are only the necessary consequence of their geometrical form, which form itself is the necessary con- sequence of mechanical conditions totally unconnected with design, and incapable of rendering an account of the origin of the insects that make the cells. The geometrical cell of the wasps and bees that I have had an opportunity of examining may be dinded into three classes. 1st. Hexagonal cells formed of adjoining pyramidal figufes, with slightly curved axes, not terminating in a point, but in a rounded extremity. * Read before the Natural History Society of Dublin, November 21, 1863. [Reprinted from a separate pamphlet by permission of the author.] 416 Rev. S. Haughton on the Form of the Cells. The British tree-wasp forms its pupa-cells in this manner, and, in consequence of the pyramidal form of the hexagonal cells, the comb opens out on the lower side, so as to present a larger surface than on the upper side. 2nd. Hexagonal cells formed of adjoining prismatic figures, with rectilinear axes, terminated by a truncated plane, at right angles to the axes of the prisms. These cells are found in wasps' nests from St. Lucia, in the West Indies, and at Graham's Town, in South Africa, which were placed at my disposal for this investigation by Mr. Robert J. Montgomery. 3rd. Hexagonal cells formed of adjoining prismatic figures, with rectilinear axes, terminated by three faces of a rhombic dodecahedron, which three faces also form each one-third of the termination of a similar set of adjoining hexagonal prismatic cells, placed end to end behind the first set of prisms. This double comb is produced by the well-known form of the cells of the honey-bee. All these varieties of cells may be accounted for, simply by the mechanical pressure of the insects against each other during the formation of the cell. In consequence of the instinct that compels them to work with reference to a plane, and of the cylindrical form of the insects' bodies, the cells must be hexa- gons ; and in consequence of the instinct that induces the bees to form double combs, the mutual pressure of their heads against each other compels the bottom of the cell to assume the form of the rhombic dodecahedron. If we could imagine spherical insects endowed with the instinct of working from a point and not a plane, their cells would cease to affect the forms of the hexagon and rhombic dodecahedron, and would imitate the totally different form of the pentagonal dodecahedron — instances of which may be seen in the bubbles produced in the froth of an organic solution, and in the shapes of the elementary cells of vegetables, equally restricted in their growth in every direction — ■ and also in the pentagonal faces assumed by leaden bullets made to fill completely the inside of a hollow shell, and then dis- charged against a bank of earth, or a wall, from a mortar. On this subject, I cannot do better than quote the words of Ruffon, who was the first person that put forward a rational theory of the shape of the cells of bees. The passage which I quote may be found in his * Histoire Naturelle,' tom. iv. p. 99 : — "Dirai-je encore un mot; ces cellules des abeilles, ces hexa- gones, tant vantes, tant admires, me fournissent une preuve de plus centre I'enthousiasme et I'admiration : cette figure, toute geometrique et toute reguliere qu'elle nous paroit, et qu'elle est en effet dans la speculation, n'est ici qu'un resultat mecanique made by various Wasps and by the Honey-Bee. 417 et assez imparfait qui se trouve souvent dans la nature^ et que Pon remarque meme dans ses productions les plus brutes ; les cristaux et plusieurs autres pierres, quelques sels, &c., prennent constamment cette figure dans leur formation. Qu^on observe les petites ecailles de la peau d'une roussette, on verra qu^elles sont hexagones, parce que chaque ecaille croissant en meme temps se fait obstacle^ et tend a occuper le plus d'espace qu'il est possible dans un espace donne : on voit ces memes hexa- gones dans le second estomac des animaux ruminans, on les trouve dans les graines, dans leurs capsules, dans certaines fleurs, &c. Qu^on remplisse un vaisseau de pois, on plutot de quelque autre graine cylindrique, et qu^on le ferme exactement apres y avoir verse autant d'eau que les inter^^alles qui restent entre ces graines peuvent en recevoir ; qu^on fasse bouillir cette eau, tous ces cylindres deviendront de colonnes a six pans. On en voit clairement la raison, qui est purement mecanique; chaque graine, dont la figure est cylindrique, tend par son ren- flement a occuper le plus d'espace possible dans un espace donne, elles deviennent done toutes necessairement hexagones par la compression reciproque. Chaque abeille cherche a occuper de meme le plus d'espace possible dans un espace donne, il est done necessaire aussi puisque le coi'ps des abeilles est cyhndrique, que leurs cellules soient hexagones, — par la meme raison des obstacles reciproques. On donne plus d'esprit aux mouches dont les ouvrages sont les plus reguhers; les abeilles sont, dit-on, plus ingenieuses que les guepes, que les frelons, &c., qui savent aussi I'architecture, mais dont les con- structions sont plus grossieres et plus iiTcgulieres que celles des abeilles : on ne veut pas voii", ou Ton ne se doute pas que cette regularite, plus ou moins grande, depend uniquement du nombre et de la figure, et nullement de I'intelligence de ces petites betes ; plus elles sont nombreuses, plus il y a des forces qui agissent egalement et qui s'opposent de meme, plus il y a par consequent de contrainte mecanique, de regularite forcee, et de perfection apparente dans leurs productions." — Bvffon. The opinions of the older writers, especially of mathematicians, on this subject, differ widely from those advanced by Buffon. I shall here translate some of the most important of the pas- sages bearing on this point. The famous Pappus, of Alexandria, in the Introduction to the Fifth Book of his Mathematical Collections, says : — " God has imparted to men, indeed, the best and most perfect knowledge of wisdom and discipline ; and has assigned to some animals, devoid of reason, a certain portion. To men, there- fore, as making use of reason. He has permitted that they should do all things by reason and demonstration ; but to other animals 418 Rev. S. Haughton on the Form of the Cells . without reason, He has given the possession of what is useful and conducive to life, by a certain natural providence. "Any one may understand this to be so, as well in many other kinds of animals, and more especially in bees. For order, and a certain admirable deference to those who rule in their republic, ambition, moreover, and cleanliness, heap together an abundance of honey ; but their foresight and economy concern- ing its conservation are much more admirable : for holding it for certain, as is just, that they carry back some portion of ambrosia from the gods to choice men, they pour out this, not rashly on the ground, or into wood, or any other unformed and misshapen matter; but collecting from the sweetest flowers that grow in the earth, they form from them most excellent vases as a receptacle for the honey (which the Greeks call Krjpia, and the Latins favi), all indeed, equal, similar, and cohering among themselves, of the hexagon species. Now it is thus evident that they construct these by a certain geometrical fore- sight ; for they consider it fit that all the figures should cohere together and have common sides, lest anything, falling into the intervening spaces, should spoil and corrupt their work. " Hence, three rectilinear and ordinate figures can efiect what is proposed — I mean ordinate figures which are equilateral and equiangular, for ordinate and dissimilar figures did not please the bees themselves. Now, equilateral triangles, and squares, and hexagons (neglecting other dissimilar figures filling space) may be placed next each other, so as to have common sides — other ordinate figures cannot; for the space about the same point is filled, either by six equilateral triangles, or by four squares, or by three hexagons; but three pentagons are less than sufficient, and four are more than sufficient to fill the space round a point, neither can three heptagons be established, so as to fill the space round a point*. " The same reasoning will apply much more to figures having a greater number of sides. There being, then, three figures, w^hich, of themselves, can fill up the space round a point, viz. the triangle, the square, and the hexagon ; the bees have wisely selected for their structure that which contains most angles, suspecting, indeed, that it could hold more honey than either of the others. " The bees, forsooth, know only what is useful to themselves, viz. that the hexagon is greater than the square or triangle, and can hold more honey, an equal quantity of material being em- ployed in the construction of each ; but we, who profess to have more wisdom than the bees, will investigate something even * The proofs of these assertions are omitted in this translation. made by various Wasps and by the Honey -Bee. 419 more remarkable, viz. that, of plane figures, which are equi- lateral and equiangular, and have equal perimeters, that is always the greatest which consists of most angles, and the circle is the greatest of all, pro\'ided it be included in a perimeter equal to theirs." — Pappus. In 1712, Maraldi published, in the 'Memoires de FAcademie des Sciences, Paris,' 1712, p. 299, a remarkable paper, in which is investigated, for the first time, the terminal planes of the bees' cell, which are now well known to be formed of the faces of the rhombic dodecahedron. He appears to have believed that the object of having lozenges of the same form, as termi- nating planes, was to enable the bees to carry in their mind the idea of one geometrical form only, in addition to their original idea of the hexagon. The angles of the lozenge are found by him to be 110=^ and 70°, by observation; and 109^ 28' and 70° 32' by calculation. He gives, also, the following mean measurements of the cells : — In a foot long of comb there are from 60 to 66 cells, about two lines for each cell, and the depth of the cell is five lines. Reaumur appears to have been the first who introduced the fantastic idea of economy of wax, as the motive cause of the peculiar shape of the terminating planes, and, not being a geo- meter, he obtained the assistance of Konig to calculate the angle of the lozenge which should give the least surface with a given volume. Konig determined this angle at 109° 26', agree- ing with Maraldi within two minutes. MacLaurin published, in the 'Philosophical Transactions,' 1743, p. 565, an elaborate geometrical paper on the subject, in which he proves that the tangent of the angle in question is the square root of 2, and that it is therefore equal to 109° 28' 16"; and he computes the saving of wax as " almost one-fourth part of the pains and expense of wax they bestow, above what was necessary for completing the parallelogram side of the cells." L'HuUier, in 1781, published, in the ' Berlin Memoirs,' p. 277, an elaborate discussion of the entire problem, in which he arrived at the following results, already found by MacLamin's geometrical method : — a. That the economy of wax is less than one-fifth of what would make a flat base. b. That the economy of wax, referred to the total expenditure, is JfSt, so that the bees can make fifty-one cells, instead of fifty, by the adoption of the rhombic dodecahedron. He does not share, however, in the enthusiasm of the natu- ralists, but maintains and proves that mathematicians could make cells, of the same form as those of the bees, which, instead of using only a minimum of wax^ would use the minimum mini- 420 Rev. S. Haughton on the Origin of Species. morum, so that five cells could be made of less wax than that which now makes only four, instead of fifty-one out of fifty. Notwithstanding this conclusive decision in favour of the mathematicians, the advocates of final cause, and those who maintain that economy of wax can create a new species, have both persisted in using the bees' cell in illustration of their re- spective theories, with a pertinacity that proves the persistent vitality of an exploded theory. In illustration of this remark- able tendency of false theories to reproduce themselves, I shall here add, as an appendix to my account of the form of the wasps' and bees' cells, some remarks on the origin of species, the substance of which originally appeared in the ' Natural His- tory Review ' of 1860. Appendix on the Origin of Species. The active and restless mind of man has never been content with the knowledge of the present, but has always sought to know the future and the past. The guesses of the ancients as to the future of man are amongst the most interesting and, at the same time, the most puerile of their philosophical specula- tions. The reader of the Tusculan Disputations rises from his task, charmed by the style of the writer, but thankful that a certain revelation of the future renders him immeasurably su- perior in knowledge to the weavers of these pleasant webs of fiction ; and though he admires the skill of the ingenious sophists who live again and dispute in the pages of Cicero, he would not for an instant exchange his own position for theirs. The moderns have resolved, by their speculations on the past, to show that in ingenuity and oddness of conceit, and, probably, also in wideness from the truth, they are in no respect inferior to the ancients. The future being shut out from us, we are re- solved to try what we can effect, in proof of our versatility of imagination, by guessing at the history of the past. To establish a character for subtlety and skill, in drawing large conclusions on this subject from slender premises, the first requisite is ignorance of what other speculators have attempted before us in the same field ; and the second is, a firm confidence in our own special theory. Neither of these requisites can be considered wanting in those who are engaged in the task of reproducing Lamarck's theory of organic life, either as alto- gether new, or with but a tattered and threadbare cloak thrown over its original nakedness. The sciences of geology and political economy are mainly answerable for the revival of these exploded and forgotten fancies, — geology, in supplying the lost history of organic life, which could never be studied profoundly from the creatures Rev. S. Haughton on the Origin of Species. 421 living at any given time ; and political economy, in furnishing, from its mean and sordid motives, a ^lalthusiau force, supposed to be sufficient to supply the wants of previous theories. One of the earliest speculators on the origin of the diversified forms of life we see around us, and class as varieties, species, and genera, was Buffon, who published in 1766* his theory of the derivation of all mammal forms by degradation, from fifteen primary and perfect types, and nine special or isolated species. This theory of ^i,oyeve(Tt