Chapter XIX: Part 19

The brilliant warning colours of many caterpillars attracted the attention of Darwin when he was thinking over his hypothesis of sexual selection, and he wrote to A. R. Wallace on the subject (C. Darwin, _Life and Letters_, London, 1887, iii. 93). Wallace, in reply, suggested their interpretation as warning colours, a suggestion since verified by experiment (_Proc. Ent. Soc. Lond._, 1867, p. lxxx; _Trans. Ent. Soc. Lond._, 1869, pp. 21 and 27). Although animals with warning colours are probably but little attacked by the ordinary enemies of their class, they have special enemies which keep the numbers down to the average. Thus the cuckoo appears to be an insectivorous bird which will freely devour conspicuously coloured unpalatable larvae. The effect of the warning colours of caterpillars is often intensified by gregarious habits. Another aposematic use of colours and structures is to divert attention from the vital parts, and thus give the animal attacked an extra chance of escape. The large, conspicuous, easily torn wings of butterflies and moths act in this way, as is found by the abundance of individuals which may be captured with notches bitten symmetrically out of both wings when they were in contact. The eye-spots and "tails" so common on the hinder part of the hind wing, and the conspicuous apex so frequently seen on the fore wing, probably have this meaning. Their position corresponds to the parts which are most offen found to be notched. In some cases (e.g. many _Lycaenidae_) the "tail" and eye-spot combine to suggest the appearance of a head with antennae at the posterior end of the butterfly, the deception being aided by movements of the hind wings. The flat-topped "tussocks" of hair on many caterpillars look like conspicuous fleshy projections of the body, and they are held prominently when the larva is attacked. If seized, the "tussock" comes out, and the enemy is greatly inconvenienced by the fine branched hairs. The tails of lizards, which easily break off, are to be similarly explained, the attention of the pursuer being probably still further diverted by the extremely active movements of the amputated member. Certain crabs similarly throw off their claws when attacked, and the claws continue to snap most actively. The tail of the dormouse, which easily comes off, and the extremely bushy tail of the squirrel, are probably of use in the same manner. Animals with warning colours often tend to resemble each other superficially. This fact was first pointed out by H. W. Bates in his paper on the theory of mimicry (_Trans. Linn. Soc._ vol. xxiii., 1862, p. 495). He showed that the conspicuous, presumably unpalatable, tropical American butterflies, belonging to very different groups, which are mimicked by others, also tend to resemble each other, the likeness being often remarkably exact. These resemblances were not explained by his theory of mimicry, and he could only suppose that they had been produced by the direct influence of a common environment. The problem was solved in 1879 by Fritz Muller (see _Proc. Ent. Soc. Lond._, 1879, p. xx.), who suggested that life is saved by this resemblance between warning colours, inasmuch as the education of young inexperienced enemies is facilitated. Each species which falls into a group with common warning (_synaposematic_) colours contributes to save the lives of the other members. It is sufficiently obvious that the amount of learning and remembering, and consequently of injury and loss of life involved in the process, are reduced when many species in one place possess the same aposematic colouring, instead of each exhibiting a different "danger-signal." These resemblances are often described as "Mullerian mimicry," as distinguished from true or "Batesian mimicry" described in the next section. Similar synaposematic resemblances between the specially protected groups of butterflies were afterwards shown to exist in tropical Asia, the East Indian Islands and Polynesia by F. Moore (_Proc. Zool. Soc._, 1883, p. 201), and in Africa by E. B. Poulton (_Report Brit. Assoc._, 1897, p. 688). R. Meldola (_Ann. and Mag. Nat. Hist._ x., 1882, p. 417) first pointed out and explained in the same manner the remarkable general uniformity of colour and pattern which runs through so many species of each of the distasteful groups of butterflies; while, still later, Poulton (_Proc. Zool. Soc._, 1887, p. 191) similarly extended the interpretation to the synaposematic resemblances between animals of all kinds in the same country. Thus, for example, longitudinal or circular bands of the same strongly contrasted colours are found in species of many groups with distant affinities.

Certain animals, especially the Crustacea, make use of the special defence and warning colours of other animals. Thus the English hermit-crab, _Pagurus Bernhardus_, commonly carries the sea-anemone, _Sagartia parasitica_, on its shell; while another English species, _Pagurus Prideauxii_, inhabits a shell which is invariably clothed by the flattened _Adamsia palliata_.

The white patch near the tail which is frequently seen in the gregarious Ungulates, and is often rendered conspicuous by adjacent black markings, probably assists the individuals in keeping together; and appearances with probably the same interpretation are found in many birds. The white upturned tail of the rabbit is probably of use in enabling the individuals to follow each other readily. The difference between a typical aposematic character appealing to enemies, and episematic intended for other individuals of the same species, is well seen when we compare such examples as (1) the huge banner-like white tail, conspicuously contrasted with the black or black and white body, by which the slow-moving skunk warns enemies of its power of emitting an intolerably offensive odour; (2) the small upturned white tail of the rabbit, only seen when it is likely to be of use and when the owner is moving, and, if pursued, very rapidly moving, towards safety.

_Mimicry_ (see also MIMICRY) or _Pseudo-sematic Colours_.--The fact that animals with distant affinities may more or less closely resemble each other was observed long before the existing explanation was possible. Its recognition is implied in a number of insect names with the termination -_formis_, usually given to species of various orders which more or less closely resemble the stinging Hymenoptera. The usefulness of the resemblance was suggested in Kirby and Spence's _Introduction to Entomology_, London, 1817, ii. 223. H. W. Bates (_Trans. Linn. Soc._ vol. xxiii., 1862, p. 495) first proposed an explanation of mimicry based on the theory of natural selection. He supposed that every step in the formation and gradual improvement of the likeness occurred in consequence of its usefulness in the struggle for life. The subject is of additional interest, inasmuch as it was one of the first attempts to apply the theory of natural selection to a large class of phenomena up to that time well known but unexplained. Numerous examples of mimicry among tropical American butterflies were discussed by Bates in his paper; and in 1866 A. R. Wallace extended the hypothesis to the butterflies of the tropical East (_Trans. Linn. Soc._ vol. xxv., 1866, p. 19); Roland Trimen (_Trans. Linn. Soc._ vol. xxvi., 1870, p. 497) to those of Africa in 1870. The term mimicry is used in various senses. It is often extended, as indeed it was by Bates, to include all the superficial resemblances between animals and any part of their environment. Wallace, however, separated the cryptic resemblances already described, and the majority of naturalists have followed this convenient arrangement. In cryptic resemblance an animal resembles some object of no interest to its enemy (or prey), and in so doing is concealed; in mimicry an animal resembles some other animal which is specially disliked by its enemy, or some object which is specially attractive to its prey, and in so doing becomes conspicuous. Some naturalists have considered mimicry to include all superficial likenesses between animals, but such a classification would group together resemblances which have widely different uses. (1) The resemblance of a mollusc to the coral on which it lives, or an external parasite to the hair or skin of its host, would be _procryptic_; (2) that between moths which resemble lichen, _syncryptic_; (3) between distasteful insects, _synaposematic_; (4) between the Insectivor mole and the Rodent mole-rat, _syntechnic_; (5) the essential element in mimicry is that it is a false warning (pseud-aposematic) or false recognition (pseud-episematic) character. Some have considered that mimicry indicates resemblance to a moving object; but apart from the non-mimetic likenesses between animals classified above, there are ordinary cryptic resemblances to drifting leaves, swaying bits of twig, &c., while truly mimetic resemblances are often specially adapted for the attitude of rest. Many use the term mimicry to include synaposematic as well as pseudo-sematic resemblances, calling the former "Mullerian," the latter "Batesian," mimicry. The objection to this grouping is that it takes little account of the deceptive element which is essential in mimicry. In synaposematic colouring the warning is genuine, in pseud-aposematic it is a sham. The term mimicry has led to much misunderstanding from the fact that in ordinary speech it implies deliberate imitation. The production of mimicry in an individual animal has no more to do with consciousness or "taking thought" than any of the other processes of growth. Protective mimicry is here defined as an advantageous and superficial resemblance of one animal to another, which latter is specially defended so as to be disliked or feared by the majority of enemies of the groups to which both belong--a resemblance which appeals to the sense of sight, sometimes to that of hearing, and rarely to smell, but does not extend to deep-seated characters except when the superficial likeness is affected by them. _Mutatis mutandis_ this definition will apply to aggressive (pseud-episematic) resemblance. The conditions under which mimicry occurs have been stated by Wallace:--"(1) that the imitative species occur in the same area and occupy the same station as the imitated; (2) that the imitators are always the more defenceless; (3) that the imitators are always less numerous in individuals; (4) that the imitators differ from the bulk of their allies; (5) that the imitation, however minute, is _external_ and _visible_ only, never extending to internal characters or to such as do not affect the external appearance." It is obvious that conditions 2 and 3 do not hold in the case of Mullerian mimicry. Mimicry has been explained, independently of natural selection, by the supposition that it is the common expression of the direct action of common causes, such as climate, food, &c.; also by the supposition of independent lines of evolution leading to the same result without any selective action in consequence of advantage in the struggle; also by the operation of sexual selection.

It is proposed, in conclusion, to give an account of the broad aspects of mimicry, and attempt a brief discussion of the theories of origin of each class of facts (see Poulton, _Linn. Soc. Journ. Zool._, 1898, p. 558). It will be found that in many cases the argument here made use of applies equally to the origin of cryptic and sematic colours. The relationship between these classes has been explained: mimicry is, as Wallace has stated (_Darwinism_, London, 1889), merely "an exceptional form of protective resemblance. "Now, protective (cryptic) resemblance cannot be explained on any of the lines suggested above, except natural selection; even sexual selection fails, because cryptic resemblance is especially common in the immature stages of insect life. But it would be unreasonable to explain mimetic resemblance by one set of principles and cryptic by another and totally different set. Again, it may be plausible to explain the mimicry of one butterfly for another on one of the suggested lines, but the resemblance of a fly or moth to a wasp is by no means so easy, and here selection would be generally conceded; yet the appeal to antagonistic principles to explain such closely related cases would only be justified by much direct evidence. Furthermore, the mimetic resemblances between butterflies are not haphazard, but the models almost invariably belong only to certain sub-families, the _Danainae_ and _Acraeinae_ in all the warmer parts of the world, and, in tropical America, the _Ithomiinae_ and _Heliconinae_ as well. These groups have the characteristics of aposematic species, and no theory but natural selection explains their invariable occurrence as models wherever they exist. It is impossible to suggest, except by natural selection, any explanation of the fact that mimetic resemblances are confined to changes which produce or strengthen a superficial likeness. Very deep-seated changes are generally involved, inasmuch as the appropriate instincts as to attitude, &c., are as important as colour and marking. The same conclusion is reached when we analyse the nature of mimetic resemblance and realize how complex it really is, being made up of _colours_, both pigmentary and structural, _pattern_, _form_, _attitude_ and _movement_. A plausible interpretation of colour may be wildly improbable when applied to some other element, and there is _no_ explanation except natural selection which can explain all these elements. The appeal to the direct action of local conditions in common often breaks down upon the slightest investigation, the difference in habits between mimic and model in the same locality causing the most complete divergence in their conditions of life. Thus many insects produced from burrowing larvae mimic those whose larvae live in the open. Mimetic resemblance is far commoner in the female than in the male, a fact readily explicable by selection, as suggested by Wallace, for the female is compelled to fly more slowly and to expose itself while laying eggs, and hence a resemblance to the slow-flying freely exposed models is especially advantageous. The facts that mimetic species occur in the same locality, fly at the same time of the year as their models, and are day-flying species even though they may belong to nocturnal groups, are also more or less difficult to explain except on the theory of natural selection, and so also is the fact that mimetic resemblance is produced in the most varied manner. A spider resembles its model, an ant, by a modification of its body-form into a superficial resemblance, and by holding one pair of legs to represent antennae; certain bugs (Hemiptera) and beetles have also gained a shape unusual in their respective groups, a shape which superficially resembles an ant; a Locustid (_Myrmecophana_) has the shape of an ant painted, as it were, on its body, all other parts resembling the background and invisible; a Membracid (Homoptera) is entirely unlike an ant, but is concealed by an ant-like shield. When we further realize that in this and other examples of mimicry "the likeness is almost always detailed and remarkable, however it is attained, while the methods differ absolutely," we recognize that natural selection is the only possible explanation hitherto suggested. In the cases of aggressive mimicry an animal resembles some object which is attractive to its prey. Examples are found in the flower-like species of _Mantis_, which attract the insects on which they feed. Such cases are generally described as possessing "alluring colours," and are regarded as examples of aggressive (anticryptic) resemblance, but their logical position is here.

_Colours displayed in Courtship, Secondary Sexual Characters, Epigamic Colours._--Darwin suggested the explanation of these appearances in his theory of _sexual selection_ (_The Descent of Man_, London, 1874). The rivalry of the males for the possession of the females he believed to be decided by the preference of the latter for those individuals with especially bright colours, highly developed plumes, beautiful song, &c. Wallace does not accept the theory, but believes that natural selection, either directly or indirectly, accounts for all the facts. Probably the majority of naturalists follow Darwin in this respect. The subject is most difficult, and the interpretation of a great proportion of the examples in a high degree uncertain, so that a very brief account is here expedient. That selection of some kind has been operative is indicated by the diversity of the elements into which the effects can be analysed. The most complete set of observations on epigamic display was made by George W. and Elizabeth G. Peckham upon spiders of the family _Attidae_ (_Nat. Hist. Soc. of Wisconsin_, vol. i., 1889). These observations afforded the authors "conclusive evidence that the females pay close attention to the love-dances of the males, and also that they have not only the power, but the will, to exercise a choice among the suitors for their favour." Epigamic characters are often concealed except during courtship; they are found almost exclusively in species which are diurnal or semi-diurnal in their habits, and are excluded from those parts of the body which move too rapidly to be seen. They are very commonly directly associated with the nervous system; and in certain fish, and probably in other animals, an analogous heightening of effect accompanies nervous excitement other than sexual, such as that due to fighting or feeding. Although there is epigamic display in species with sexes alike, it is usually most marked in those with secondary sexual characters specially developed in the male. These are an exception to the rule in heredity, in that their appearance is normally restricted to a single sex, although in many of the higher animals they have been proved to be latent in the other, and may appear after the essential organs of sex have been removed or become functionless. This is also the case in the Aculeate Hymenoptera when the reproductive organs have been destroyed by the parasite _Stylops_. J. T. Cunningham has argued (_Sexual Dimorphism in the Animal Kingdom_, London, 1900) that secondary sexual characters have been produced by direct stimulation due to contests, &c., in the breeding period, and have gradually become hereditary, a hypothesis involving the assumption that acquired characters are transmitted. Wallace suggests that they are in part to be explained as "recognition characters," in part as an indication of surplus vital activity in the male.

AUTHORITIES.--The following works may also be consulted:--T. Eimer,
_Orthogenesis der Schmetterlinge_ (Leipzig, 1898); E. B. Poulton, _The
Colours of Animals_ (London, 1890); F. E. Beddard, _Animal Coloration_
(London, 1892); E. Haase, _Researches on Mimicry_ (translation,
London, 1896); A. R. Wallace, _Natural Selection and Tropical Nature_
(London, 1895); _Darwinism_ (London, 1897); A. H. Thayer and G. H.
Thayer, _Concealing-Coloration in the Animal Kingdom_ (New York,
1910). (E. B. P.)

2. CHEMISTRY

The coloration of the _surface_ of animals is caused either by _pigments_, or by a certain _structure_ of the surface by means of which the light falling on it, or reflected through its superficial transparent layers, undergoes diffraction or other optical change. Or it may be the result of a combination of these two causes. It plays an important part in the relationship of the animal to its environment, in concealment, in mimicry, and so on; the presence of a pigment in the integument may also serve a more direct physiological purpose, such as a respiratory function. The coloration of birds' feathers, of the skin of many fishes, of many insects, is partially at least due to structure and the action of the peculiar pigmented cells known as "chromatophores" (which W. Garstang defines as pigmented cells specialized for the discharge of the chromatic function), and is much better marked when these have for their background a "reflecting layer" such as is provided by guanin, a substance closely related to uric acid. Such a mechanism is seen to greatest advantage in fishes. Among these, guanin may be present in a finely granular form, causing the light falling on it to be scattered, thus producing a white effect; or it may be present in a peculiar crystalline form, the crystals being known as "iridocytes"; or in a layer of closely apposed needles forming a silvery sheet or mirror. In the iris of some fishes the golden red colour is produced by the light reflected from such a layer of guanin needles having to pass through a thin layer of a reddish pigment, known as a "lipochrome." Again, in some lepidopterous insects a white or a yellow appearance is produced by the deposition of uric acid or a nearly allied substance on the surface of the wings. In many animals, but especially among invertebrates, colouring matters or pigments play an important role in surface coloration; in some cases such coloration may be of benefit to the animal, but in others the integument simply serves as an organ for the excretion of waste pigmentary substances. Pigments (1) may be of direct physiological importance; (2) they may be excretory; or (3) they may be introduced into the body of the animal with the food.

Of the many pigments which have been described up to the present time, very few have been subjected to elementary chemical analysis, owing to the great difficulties attending their isolation. An extremely small amount of pigment will give rise to a great amount of coloration, and the pigments are generally accompanied by impurities of various kinds which cling to them with great tenacity, so that when one has been thoroughly cleansed very little of it remains for ultimate analysis. Most of these substances have been detected by means of the spectroscope, their absorption bands serving for their recognition, but mere identity of spectrum does not necessarily mean chemical identity, and a few chemical tests have also to be applied before a conclusion can be drawn. The absorption bands are referred to certain definite parts of the spectrum, such as the Fraunhofer lines, or they may be given in wave-lengths. For this purpose the readings of the spectroscope are reduced to wave-lengths by means of interpolation curves; or if Zeiss's microspectroscope be used, the position of bands in wave-lengths (denoted by the Greek letter [lambda]) may be read directly.

Haemoglobin, the red colouring matter of vertebrate blood, C758H1203N195S3FeO218, and its derivatives haematin, C32H30N4FeO3, and haematoporphyrin, C16H18N2O3, are colouring matters about which we possess definite chemical knowledge, as they have been isolated, purified and analysed. Most of the bile pigments of mammals have likewise been isolated and studied chemically, and all of these are fully described in the text-books of physiology and physiological chemistry. Haemoglobin, though physiologically of great importance in the respiratory process of vertebrate animals, is yet seldom used for surface pigmentation, except in the face of white races of man or in other parts in monkeys, &c. In some worms the transparent skin allows the haemoglobin of the blood to be seen through the integument, and in certain fishes also the haemoglobin is visible through the integument. It is a curious and noteworthy fact that in some invertebrate animals in which no haemoglobin occurs, we meet with its derivatives. Thus haematin is found in the so-called bile of slugs, snails, the limpet and the crayfish. In sea-anemones there is a pigment which yields some of the decomposition-products of haemoglobin, and associated with this is a green pigment apparently identical with biliverdin (C16H18N2O4), a green bile pigment. Again, haematoporphyrin is found in the integuments of star-fishes and slugs, and occurs in the "dorsal streak" of the earth-worm _Lumbricus terrestris_, and perhaps in other species. Haematoporphyrin and biliverdin also occur in the egg-shells of certain birds, but in this case they are derived from haemoglobin. Haemoglobin is said to be found as low down in the animal kingdom as the Echinoderms, e.g. in _Ophiactis virens_ and _Thyonella gemmata_. It also occurs in the blood of _Planorbis corneus_ and in the pharyngeal muscles of other mollusca.

A great number of other pigments have been described; for example, in the muscles and tissues of animals, both vertebrate and invertebrate, are the histohaematins, of which a special muscle pigment, myohaematin, is one. In vertebrates the latter is generally accompanied by haemoglobin, but in invertebrates--with the exception of the pharyngeal muscles of the mollusca--it occurs alone. Although closely related to haemoglobin or its derivative haemochromogen, the histohaematins are yet totally distinct, and they are found in animals where not a trace of haemoglobin can be detected. Another interesting pigment is turacin, which contains about 7% of nitrogen, found by Professor A. H. Church in the feathers of the Cape lory and other plantain-eaters, from which it can be extracted by water containing a trace of ammonia. It has been isolated, purified and analysed by Professor Church. From it may be obtained turacoporphyrin, which is identical with haematoporphyrin, and gives the band in the ultra-violet which J. L. Soret and subsequently A. Gamgee have found to be characteristic of haemoglobin and its compounds. Turacin itself gives a peculiar two-banded spectrum, and contains about 7% of copper in its molecule. Another copper-containing pigment is haemocyanin, which in the oxidized state gives a blue colour to the blood of various Mollusca and Arthropoda. Like haemoglobin, it acts as an oxygen-carrier in respiration, but it takes no part in surface coloration.

A class of pigments widely distributed among plants and animals are the lipochromes. As their name denotes, they are allied to fat and generally accompany it, being soluble in fat solvents. They play an important part in surface coloration, and may be greenish, yellow or red in colour. They contain no nitrogen. As an example of a lipochrome which has been isolated, crystallized and purified, we may mention carotin, which has recently been found in green leaves. Chlorophyll, which is so often associated with a lipochrome, has been found in some Infusoria, and in _Hydra_ and _Spongilla_, &c. In some cases it is probably formed by the animal; in other cases it may be due to symbiotic algae, while in the gastric gland of many Mollusca, Crustacea and Echinodermata it is derived from food-chlorophyll. Here it is known as entero-chlorophyll. The black pigments which occur among both vertebrate and invertebrate animals often have only one attribute in common, viz. blackness, for among the discordant results of analysis one thing is certain, viz. that the melanins from vertebrate animals are not identical with those from invertebrate animals. The melanosis or blackening of insect blood, for instance, is due to the oxidation of a chromogen, the pigment produced being known as a uranidine. In some sponges a somewhat similar pigment has been noticed. Other pigments have been described, such as actiniochrome, echinochrome, pentacrinin, antedonin, polyperythrin (which appears to be a haematoporphyrin), the floridines, spongioporphyrin, &c., which need no mention here; all these pigments can only be distinguished by means of the spectroscope.

Most of the pigments are preceded by colourless substances known as "chromogens," which by the action of the oxygen of the air and by other agencies become changed into the corresponding pigments. In some cases the pigments are built up in the tissues of an animal, in others they appear to be derived more or less directly from the food. Derivatives of chlorophyll and lipochromes especially, seem to be taken up from the intestine, probably by the agency of leucocytes, in which they may occur in combination with, or dissolved by, fatty matters and excreted by the integument. In worms especially, the skin seems to excrete many effete substances, pigments included. No direct connexion has been traced between the chlorophyll eaten with the food and the haemoglobin of blood and muscle. Attention may, however, be drawn to the work of Dr E. Schunck, who has shown that a substance closely resembling haematoporphyrin can be prepared from chlorophyll; this is known as phylloporphyrin. Not only does the _visible_ spectrum of this substance resemble that of haematoporphyrin, but the _invisible_ ultra-violet also, as shown by C. A. Schunck.

The reader may refer to E. A. Schafer's _Text-Book of Physiology_
(1898) for A. Gamgee's article "On Haemoglobin, and its Compounds"; to
the writer's papers in the _Phil. Trans._ and _Proc. Roy. Soc._ from
1881 onwards, and also _Quart. Journ. Micros. Science_ and _Journ. of
Physiol._; to C. F. W. Krukenberg's _Vergleichende physiologische
Studien_ from 1879 onwards, and to his _Vortrage_. Miss M. I. Newbigin
collected in _Colour in Nature_ (1898) most of the recent literature
of this subject. Dr E. Schunck's papers will be found under the
heading "Contribution to the Chemistry of Chlorophyll" in _Proc. Roy.
Soc._ from 1885 onwards; and Mr C. A. Schunck's paper in _Proc. Roy.
Soc._ vol. lxiii. (C. A. MacM.)

COLSTON, EDWARD (1636-1721), English philanthropist, the son of William Colston, a Bristol merchant of good position, was born at Bristol on the 2nd of November 1636. He is generally understood to have spent some years of his youth and manhood as a factor in Spain, with which country his family was long connected commercially, and whence, by means of a trade in wines and oil, great part of his own vast fortune was to come. On his return he seems to have settled in London, and to have bent himself resolutely to the task of making money. In 1681, the date of his father's decease, he appears as a governor of Christ's hospital, to which noble foundation he afterwards gave frequently and largely. In the same year he probably began to take an active interest in the affairs of Bristol, where he is found about this time embarked in a sugar refinery; and during the remainder of his life he seems to have divided his attention pretty equally between the city of his birth and that of his adoption. In 1682 he appears in the records of the great western port as advancing a sum of L1800 to its needy corporation; in 1683 as "a free burgess and _meire_ (St Kitts) merchant" he was made a member of the Merchant's Hall; and in 1684 he was appointed one of a committee for managing the affairs of Clifton. In 1685 he again appears as the city's creditor for about L2000, repayment of which he is found insisting on in 1686. In 1689 he was chosen auditor by the vestry at Mortlake, where he was residing in an old house once the abode of Ireton and Cromwell. In 1691, on St Michael's Hill, Bristol, at a cost of L8000, he founded an almshouse for the reception of 24 poor men and women, and endowed with accommodation for "Six Saylors," at a cost of L600, the merchant's almshouses in King Street. In 1696, at a cost of L8000, he endowed a foundation for clothing and teaching 40 boys (the books employed were to have in them "no tincture of Whiggism"); and six years afterwards he expended a further sum of L1500 in rebuilding the school-house. In 1708; at a cost of L41,200, he built and endowed his great foundation on Saint Augustine's Back, for the instruction, clothing, maintaining and apprenticing of 100 boys; and in time of scarcity, during this and next year, he transmitted "by a private hand" some L20,000 to the London committee. In 1710, after a poll of four days, he was sent to parliament, to represent, on strictest Tory principles, his native city of Bristol; and in 1713, after three years of silent political life, he resigned this charge. He died at Mortlake in 1721, having nearly completed his eighty-fifth year; and was buried in All Saints' church, Bristol.

Colston, who was in the habit of bestowing large sums yearly for the release of poor debtors and the relief of indigent age and sickness, and who gave (1711) no less than L6000 to increase Queen Anne's Bounty Fund for the augmentation of small livings, was always keenly interested in the organization and management of his foundations; the rules and regulations were all drawn up by his hand, and the minutest details of their constitution and economy were dictated by him. A high churchman and Tory, with a genuine intolerance of dissent and dissenters, his name and example have served as excuses for the formation of two political benevolent societies--the "Anchor" (founded 1769) and the "Dolphin" (founded 1749),--and also the "Grateful" (founded 1758), whose rivalry has been perhaps as instrumental in keeping their patron's memory green as have the splendid charities with which he enriched his native city (see BRISTOL).

See Garrard, _Edward Colston, the Philanthropist_ (4to, Bristol,
1852); Pryce, _A Popular History of Bristol_ (1861); Manchee, _Bristol
Charities_.

COLT, SAMUEL (1814-1862), American inventor, was born on the 19th of July 1814 at Hartford, Connecticut, where his father had a manufactory of silks and woollens. At the age of ten he left school for the factory, and at fourteen, then being in a boarding school at Amherst, Massachusetts, he made a runaway voyage to India, during which (in 1829) he constructed a wooden model, still existing, of what was afterwards to be the revolver (see PISTOL). On his return he learned chemistry from his father's bleaching and dyeing manager, and under the assumed name "Dr Coult" travelled over the United States and Canada lecturing on that science. The profits of two years of this work enabled him to continue his researches and experiments. In 1835, having perfected a six-barrelled rotating breech, he visited Europe, and patented his inventions in London and Paris, securing the American right on his return; and the same year he founded at Paterson, New Jersey, the Patent Arms Company, for the manufacture of his revolvers only. As early as 1837 revolvers were successfully used by United States troops, under Lieut.-Colonel William S. Harney, in fighting against the Seminole Indians in Florida. Colt's scheme, however, did not succeed; the arms were not generally appreciated; and in 1842 the company became insolvent. No revolvers were made for five years, and none were to be had when General Zachary Taylor wrote for a supply from the seat of war in Mexico. In 1847 the United States government ordered 1000 from the inventor; but before these could be produced he had to construct a new model, for a pistol of the company's make could nowhere be found. This commission was the beginning of an immense business. The little armoury at Whitneyville (New Haven, Connecticut), where the order for Mexico was executed, was soon exchanged for larger workshops at Hartford. These in their turn gave place (1852) to the enormous factory of the Colt's Patent Fire-Arms Manufacturing Company, doubled in 1861, on the banks of the Connecticut river, within the city limits of Hartford, where so many millions of revolvers with all their appendages have been manufactured. Thence was sent, for the Russian and English governments, to Tula and Enfield, the whole of the elaborate machinery devised by Colt for the manufacture of his pistols. Colt introduced and patented a number of improvements in his revolver, and also invented a submarine battery for harbour defence. He died at Hartford on the 10th of January 1862.

COLT'S-FOOT, the popular name of a small herb, _Tussilago Farfara_, a member of the natural order Compositae, which is common in Britain in damp, heavy soils. It has a stout branching underground stem, which sends up in March and April scapes about 6 in. high, each bearing a head of bright yellow flowers, the male in the centre surrounded by a much larger number of female. The flowers are succeeded by the fruits, which bear a soft snow-white woolly pappus. The leaves, which appear later, are broadly cordate with an angular or lobed outline, and are covered on the under-face with a dense white felt. The botanical name, _Tussilago_, recalls its use as a medicine for cough (_tussis_). The leaves are smoked in cases of asthma.

COLUGO, or COBEGO, either of two species of the zoological genus _Galeopithecus_. These animals live in the forests of the Malay Peninsula, Sumatra, Borneo and the Philippine Islands, where they feed chiefly on leaves, and probably also on insects. In size they may be compared with cats; the long slender limbs are connected by a broad fold of skin extending outwards from the sides of the neck and body, the fingers and toes are webbed, and the hind-limbs joined by an outer membrane as in bats. Their habits are nocturnal, and during the daytime they cling to the trunks or limbs of trees head downwards in a state of repose. With the approach of night their season of activity commences, when they may be occasionally seen gliding from tree to tree supported on their cutaneous parachute, and they have been noticed as capable of traversing in this way a space of 70 yds. with a descent of only about one in five. Europeans in the East know these animals as "flying lemurs." (See GALEOPITHECUS.)

COLUMBA, SAINT (Irish, _Colum_), Irish saint, was born on the 7th of December 521, in all probability at Gartan in Co. Donegal. His father Feidlimid was a member of the reigning family in Ireland and was closely allied to that of Dalriada (Argyll). His mother Eithne was of Leinster extraction and was descended from an illustrious provincial king. To these powerful connexions as much as to his piety and ability, he owed the immense influence he possessed. Later lives state that the saint was also called Crimthann (fox), and Reeves suggests that he may have had two names, the one baptismal, the other secular. He was afterwards known as Columkille, or Columba of the Church, to distinguish him from others of the same name. During his early years the Irish Church was reformed by Gildas and Finian of Clonard, and numerous monasteries were founded which made Ireland renowned as a centre of learning. Columba himself studied under two of the most distinguished Irishmen of his day, Finian of Moville (at the head of Strangford Lough) and Finian of Clonard. Almost as a matter of course, under such circumstances, he embraced the monastic life. He was ordained deacon while at Moville, and afterwards, when about thirty years of age, was raised to the priesthood. During his residence in Ireland he founded, in addition to a number of churches, two famous monasteries, one named Daire Calgaich (Derry) on the banks of Lough Foyle, the other Dair-magh (Durrow) in King's county.

In 563 he left his native land, accompanied by twelve disciples, and went on a mission to northern Britain, perhaps on the invitation of his kinsman Conall, king of Dalriada. Irish accounts represent Columba as undertaking this mission in consequence of the censure expressed against him by the clergy after the battle of Cooldrevny; but this is probably a fabrication. The saint's labours in Scotland must be regarded as a manifestation of the same spirit of missionary enterprise with which so many of his countrymen were imbued. Columba established himself on the island of Hy or Iona, where he erected a church and a monastery. About the year 565 he applied himself to the task of converting the heathen kingdom of the northern Picts. Crossing over to the mainland he proceeded to the residence, on the banks of the Ness, of Brude, king of the Picts. By his preaching, his holy life, and, as his earliest biographers assert, by the performance of miracles, he converted the king and many of his subjects. The precise details, except in a few cases, are unknown, or obscured by exaggeration and fiction; but it is certain that the whole of northern Scotland was converted by the labours of Columba, and his disciples and the religious instruction of the people provided for by the erection of numerous monasteries. The monastery of Iona was reverenced as the mother house of all these foundations, and its abbots were obeyed as the chief ecclesiastical rulers of the whole nation of the northern Picts. There were then neither dioceses nor parishes in Ireland and Celtic Scotland; and by the Columbite rule the bishops themselves, although they ordained the clergy, were subject to the jurisdiction of the abbots of Iona, who, like the founder of the order, were only presbyters. In matters of ritual they agreed with the Western Church on the continent, save in a few particulars such as the precise time of keeping Easter and manner of tonsure.

Columba was honoured by his countrymen, the Scots of Britain and Ireland, as much as by his Pictish converts, and in his character of chief ecclesiastical ruler he gave formal benediction and inauguration to Aidan, the successor of Conall, as king of the Scots. He accompanied that prince to Ireland in 575, and took a leading part in a council held at Drumceat in Ulster, which determined once and for all the position of the ruler of Dalriada with regard to the king of Ireland. The last years of Columba's life appear to have been mainly spent at Iona. There he was already revered as a saint, and whatever credit may be given to some portions of the narratives of his biographers, there can be no doubt as to the wonderful influence which he exercised, as to the holiness of his life, and as to the love which he uniformly manifested to God and to his neighbour.

In the summer of 597 he knew that his end was approaching. On Saturday the 8th of June he was able, with the help of one of his monks, to ascend a little hill above the monastery and to give it his farewell blessing. Returning to his cell he continued a labour in which he had been engaged, the transcription of the Psalter. Having finished the verse of the 34th Psalm where it is written, "They who seek the Lord shall want no manner of thing that is good," he said, "Here I must stop:--what follows let Baithen write"; indicating, as was believed, his wish that his cousin Baithen should succeed him as abbot. He was present at evening in the church, and when the midnight bell sounded for the nocturnal office early on Sunday morning he again went thither unsupported, but sank down before the altar and passed away as in a gentle sleep.

Several Irish poems are ascribed to Columba, but they are manifestly compositions of a later age. Three Latin hymns may, however, be attributed to the saint with some degree of certainty.

The original materials for a life of St Columba are unusually full.
The earliest biography was written by one of his successors, Cuminius,
who became abbot of Iona in 657. Much more important is the
enlargement of that work by Adamnan, who became abbot of Iona in 679.
These narratives are supplemented by the brief but most valuable
notices given by the Venerable Bede. See W. Reeves, _Life of St
Columba, written by Adamnan_ (Dublin, 1857); W. F. Skene, _Celtic
Scotland_, vol. ii. "Church and Culture" (Edinburgh, 1877).
(E. C. Q.)

COLUMBAN (543-615), Irish saint and writer, was born in Leinster in 543, and was educated in the monastery of Bangor, Co. Down. About the year 585 he left Ireland together with twelve other monks, and established himself in the Vosges, among the ruins of an ancient fortification called Anagrates, the present Anegray in the department of Haute-Saone. His enemies accused him before a synod of French bishops (602) for keeping Easter according to the old British and now unorthodox way, and a more powerful conspiracy was organized against him at the court of Burgundy for boldly rebuking the crimes of King Theuderich II. and the queen-mother Brunhilda. He was banished and forcibly removed from his monastery, and with St Gall and others of the monks he withdrew into Switzerland, where he preached with no great success to the Suebi and Alamanni. Being again compelled to flee, he retired to Italy, and founded the monastery of Bobbio in the Apennines, where he remained till his death, which took place on the 21st of November 615. His writings, which include some Latin poems, prove him a man of learning, and he appears to have been acquainted not only with the Latin classics, but also with Greek, and even Hebrew.

The collected edition of St Columban's writings was published by
Patrick Fleming in his _Collectanea sacra Hiberni_ (Louvain, 1667),
and reproduced by Migne, p. 4, vol. lxxxvi. (Paris, 1844). See
further, Wright's _Biographia Literaria_. Columban's _Regula
Coenobitalis cum Poenitenliali_ is to be found in the _Codex
Regularum_ (Paris, 1638). A complete bibliography is given in U.
Chevallier, _Repertoire des sources hist_. (Bio. Bibliogr.), vol. i.
990 (Paris, 1905).

COLUMBANI, PLACIDO, Italian architectural designer, who worked chiefly in England in the latter part of the 18th century. He belonged to the school of the Adams and Pergolesi, and like them frequently designed the enrichments of furniture. He was a prolific producer of chimney-pieces, which are often mistaken for Adam work, of moulded friezes, and painted plaques for cabinets and the like. There can be no question that the English furniture designers of the end of the 18th century, and especially the Adams, Hepplewhite and Sheraton, owed much to his graceful, flowing and classical conceptions, although they are often inferior to those of Pergolesi. His books are still a valuable store-house of sketches for internal architectural decoration. His principal works are:--_Vases and Tripods_ (1770); _A New Book of Ornaments, containing a variety of elegant designs for Modern Panels, commonly executed in Stucco, Wood or Painting, and used in decorating Principal Rooms_ (1775); _A variety of Capitals, Friezes and Corniches, and how to increase and decrease them, still retaining their proportions_ (1776). He also assisted John Crunden in the production of _The Chimneypiece Makers' Daily Assistant_ (1776).

COLUMBARIUM (Lat. _columba_, a dove), a pigeon-house. The term is applied in architecture to those sepulchral chambers in and near Rome, the walls of which were sunk with small niches (_columbaria_) to receive the cinerary urns. Vitruvius (iv. 2) employs the term to signify the holes made in a wall to receive the ends of the timbers of a floor or roof.

COLUMBIA, a city and the county-seat of Boone county, Missouri, U.S.A., situated in the central part of the state, about 145 m. (by rail) W.N.W. of St Louis. Pop. (1890) 4000; (1900) 5651 (1916 negroes); (1910) 9662. Columbia is served by the Wabash and the Missouri, Kansas & Texas railways. It is primarily an educational centre, is a market for grain and farm products, and has grain elevators, a packing house, a shoe factory and brick works. Columbia is the seat of the University of Missouri, a coeducational state institution, established in 1839 and opened in 1841; it received no direct financial support from the state until 1867, and its founding was due to the self-sacrifice of the people of the county. It is now liberally supported by the state; in 1908 its annual income was about $650,000. In 1908 the university had (at Columbia) 200 instructors and 2419 students, including 680 women; included in its library is the collection of the State Historical Society. The School of Mines of the university is at Rolla, Mo.; all other departments are at Columbia. A normal department was established in 1867 and opened in 1868; and women were admitted to it in 1869. The College of Agriculture and Mechanic Arts became a department of the university in 1870. The law department was opened in 1872, the medical in 1873, and the engineering in 1877. The graduate department was established in 1896, and in 1908 a department of journalism was organized. On the university campus in the quadrangle is the monument of grey granite erected over the grave of Thomas Jefferson, designed after his own plans, and bearing the famous inscription written by him. It was given to the university by descendants of Jefferson when Congress appropriated money for the monument now standing over his grave. Near the city is the farm of the agricultural college and the experiment station. At Columbia, also, are the Parker Memorial hospital, the Teachers College high school, the University Military Academy, the Columbia Business College, Christian College (Disciples) for women, established in 1851, its charter being the first granted by Missouri for the collegiate education of Protestant women; the Bible College of the Disciples of Christ in Missouri; and Stephens College (under Baptist control) for women, established in 1856. The municipality owns the water-works and the electric lighting plant. Columbia was first settled about 1821.

COLUMBIA, a borough of Lancaster county, Pennsylvania, U.S.A., on the W. bank of the Susquehanna river (here crossed by a long steel bridge), opposite Wrightsville and about 81 m. W. by N. of Philadelphia. Pop. (1890) 10,599; (1900) 12,316, of whom 772 were foreign-born; (1910) 11,454. It is served by the Pennsylvania, the Philadelphia, Baltimore & Washington, the Philadelphia & Reading, and the Northern Central railways, and by interurban electric railways. The river here is about a mile wide, and a considerable portion of the borough is built on the slope of a hill which rises gently from the river-bank and overlooks beautiful scenery. The Pennsylvania railway has repair shops here, and among Columbia's manufactures are silk goods, embroidery and laces, iron and steel pipe, engines, laundry machinery, brushes, stoves, iron toys, umbrellas, flour, lumber and wagons; the city is also a busy shipping and trading centre. Columbia was first settled, by Quakers, in 1726; it was laid out as a town in 1787; and in 1814 it was incorporated. In 1790 it was one of several places considered in Congress for a permanent site of the national capital.

COLUMBIA, the capital city of South Carolina, U.S.A., and the county-seat of Richland county, on the E. bank of the Congaree river, a short distance below the confluence of the Saluda and the Broad rivers, about 130 m. N.W. of Charleston. Pop. (1890) 15,353; (19O0) 21,108, of whom 9858 were negroes; and (1910) 26,319. It is served by the Atlantic Coast Line, the Southern, the Seaboard Air Line, and the Columbia, Newberry & Laurens railways. Columbia is picturesquely situated on the level top of a bluff overlooking the Congaree, which falls about 36 ft. in passing by, but is navigable for the remainder of its course. The surrounding country is devoted chiefly to cotton culture. The state house, United States government building and city hall are fine structures. Some of the new business houses are ten or more storeys in height. The state penitentiary and the state insane asylum are located here, and Columbia is an important educational centre, being the seat of the university of South Carolina, the Columbia College for women (Methodist Episcopal South, 1854), the College for women (Presbyterian, 1890), and the Presbyterian Theological Seminary (1828); and the Allen University (African Methodist Episcopal; coeducational, 1880), and the Benedict College (Baptist) for negroes. The University of South Carolina, organized in 1801 and opened in 1805, was known as South Carolina College in 1805-1863, 1878-1887 and 1891-1906, and as the university of South Carolina in 1866-1877, 1888-1891 and after 1906; in 1907-1908 it had departments of arts, science, pedagogy and law, an enrolment of 285 students, and a faculty of 25 instructors. By means of a canal abundant water power is furnished by the Congaree, and the city has some of the largest cotton mills in the world; it has, besides, foundries and machine shops and manufactories of fertilizers and hosiery. The manufactures under the factory system were valued at $3,133,903 in 1900 and at $4,676,944 in 1905--a gain, greater than that of any other city in the state, of 49.2% in five years. In the neighbourhood are several valuable granite quarries. The municipality owns and operates its water-works.

While much of the site was still a forest the legislature, in 1786, chose it for the new capital. It was laid out in the same year, and in 1790 the legislature first met here. Until 1805, when it was incorporated as a village, Columbia was under the direct government of the legislature; in 1854 it was chartered as a city. On the morning of the 17th of February 1865 General W. T. Sherman, on his march through the Carolinas, entered Columbia, and on the ensuing night a fire broke out which was not extinguished until most of the city was destroyed. The responsibility for this fire was charged by the Confederates upon the Federals and by the Federals upon the Confederates.

COLUMBIA, a city and the county-seat of Maury county, Tennessee, U.S.A., situated on the Duck river, in the central part of the state, 46 m. S. of Nashville. Pop. (1890) 5370; (1900) 6052 (2716 negroes); (1910) 5754. Columbia is served by the Louisville & Nashville, and the Nashville, Chattanooga & St Louis railways. It is the seat of the Columbia Institute for girls (under Protestant Episcopal control), founded in 1836, and of the Columbia Military Academy. Columbia is in a fine farming region; is engaged extensively in the mining and shipping of phosphates; has an important trade in live-stock, especially mules; manufactures cotton, lumber, flour, bricks, pumps and woollen goods; and has marble and stone works. Columbia was settled about 1807 and was incorporated in 1822. During the Civil War it was the base from which General N. B. Forrest operated in 1862-1863, and was alternately occupied by Confederate and Federal forces during General Hood's Nashville campaign (November-December 1864).