: i at- REESE LIBRARY UNIVERSITY OF CALIFORNIA. BIOLOGY LIBRARY G Class UNIVERSITY EXTENSION MANUALS EDITED BY PROFESSOR KNIGHT THE STUDY OF ANIMAL LIFE EDITORS PREFACE This Series is primarily designed to aid the University Extension Movement throughout Great Britain, and to supply the need so widely felt by students, of Text-books for s-tudy and reference, in connection with the authorised Courses of Lectures. The Manuals differ from those already in existence in that they are not intended for School use, or for Examination purposes ; and that their aim is to educate, rather than to inform. The statement of details is meant to illustrate the working of general laws, and the development of principles ; while the historical evolution of the subject dealt with is kept in view, along with its philosophical significance. The remarkable success which has attended University Extension in Britain has been partly due to the combination of scientific treat- ment with popularity, and to the union of Duplicity with thorough- ness. This movement, however, can only reach those resident in the larger centres of population, while all over the country there are thoughtful persons who desire the same kind of teaching. It is for them also that this Series is designed. Its aim is to supply the general reader with the same kind of teaching as is given in the Lectures, and to reflect the spirit which has characterised the move- ment, viz. the combination of principles with facts, and of methods with results. The Manuals are also intended to be contributions to the Literature of the Subjects with which they respectively deal, quite apart from University Extension ; and some of them luill be found to meet a general rather than a special want. They will be issued simultaneously in England and America. Volumes dealing with separate sections of Literature, Science, Philosophy, History, and Art have been assigned to representative literary men, to University Professors, or to Extension Lecturers connected with Oxford, Cambridge, London, and the Universities of Scotland and Ireland. A list of the works in this Series will be found at the end of the volume. The Study of Animal Life BY J. ARTHUR THOMSON, M.A., F.R.S.E. LECTURER ON^ZOOLOGY, SCHOOL OF MEDICINE, EDINBURGH JOINT-AUTHOR OF ' THE EVOLUTION OF SEX ' AUTHOR OF 'OUTLINES OF ZOOLOGY' SECOND EDITION WITH ILLUSTRATIONS LONDON JOHN MURRAY, ALBEMARLE STREET i 892 OG T G "But, for my part, which write the English story, I acknowledge that no man must looke for that at my hands, which I have not received from some other : for I would bee unwilling to write anything untrue, or uncertaine out of mine own invention ; and truth on every part is so deare unto me, that I will not lie to bring any man in love and admiration with God and his works, for God needeth not the lies of men. ' , and S indicate the positions of Peripatus. Balanoglossus, and Sphenodon or Hatteria respectively. CHAP, i The Wealth of Life 13 There is of course no doubt as to the fact that some forms of life are more complex than others. It requires no faith to allow that the firstlings or Protozoa are simpler than all the rest ; that sponges, which are more or less loose colonies of unit masses imperfectly compacted together, are in that sense simpler than jellyfish, and so on. The animals most like ourselves are more intricate and more perfectly controlled organisms than those which are obviously more remote, and associated with this perfecting of structure there is an increasing fulness and freedom of life. We may arrange all the classes in series from low to high, from simple to complex, but this will express only our most generalised conceptions. For within each class there is great variety, each has its own masterpieces. Thus the simplest animals are often cased in shells of flint or lime whose crystalline architecture has great complexity. The simplest sponge is little more than a double-walled sack riddled by pores through which the water is lashed, but the Venus' Flower-Basket {Euplectella), one of the flinty sponges, has a complex system of water canals and a skeleton of flinty threads built up into a framework of marvellous intricacy and grace. The lowest insect is not much more intricate, centralised, or controlled than many a worm of the sea-shore, but the ant or the bee is a very complex self-controlled organism. More exact, therefore, than any linear series, is the image of a tree with branches springing from different levels, each branch again bearing twigs some of which rise higher than the base of the branch above. A perfect scheme of this sort might not only express the facts of structure, it might also express our notions of the blood-relationships of animals and the way in which we believe that different forms have arisen. But the wealth of form is less varied than at first sight appears. There is great wealth, but the coinage is very uniform. Our first impression is one of manifold variety ; but that gives place to one of marvellous plasticity when we see how structures apparently quite different are redu- cible to the same general plan. Thus, as the poet Goethe first clearly showed, the seed-leaves, root-leaves, stem-leaves, i4 The Study of Animal Life PART i and even the parts of the flower sepals, petals, stamens, and carpels, are in reality all leaves or appendages more or less modified for diverse work. The mouth-parts of a lobster are masticating legs, and a bird's wing is a modified arm. The old naturalists were so far right in insisting on i\ the fact of a few great types. Nature, Lamarck said, is \\ never brusque ; nor is she inventive so much as adaptive. 4. Wealth of Numbers. Large numbers are so unthink- able, and accuracy in census-taking is so difficult, that we need say little as to the number of different animals. The census includes far over a million living species a total so vast that, so far as our power of realising it is concerned, 'it is hardly affected when we admit that more than half are insects. To these recorded myriads, moreover, many newly-discovered forms are added every year now by the individual workers who with fresh eye or improved micro- scope find in wayside pond or shore pool some new thing, or again by great enterprises like the Challenger expedition. Exploring naturalists like Wallace and Semper return from tropical countries enriched with new animals from the dense forests or warm seas. Zoological Stations, notably that of Naples, are "register -houses" for the fauna of the neigh- bouring sea, not merely as to number and form, but in many cases taking account of life and history as well. Nor can we forget the stupendous roll of the extinct, to which the zoological historians continue to add as they disentomb primitive mammals, toothed birds, giant reptiles, huge amphibians, armoured fishes, gigantic cuttles, and a vast multitude of strange forms, the like of which no longer live. The length of the Zoological Record, in which the literature and discoveries of each year are chronicled, the portentous size of a volume which professes to discuss with some completeness even a single sub-class, the number of special departments into which the science of zoology is divided, suggest the vast wealth of numbers at first sight so bewildering. More than two thousand years ago Aristotle recorded a total of about 500 forms, but more new species may be described in a single volume of the Challenger Reports. We speak about the number of the stars, yet more than one CHAP, i The Wealth of Life 15 family of insects is credited with including as many different species as there are stars to count on a clear night. But far better than any literary attempt to estimate the numerical wealth of life is some practical observation, some attempted enumeration of the inmates of your aquarium, of the tenants of some pool, or of the visitors to some meadow. The naturalist as well as the poet spoke when Goethe celebrated Nature's wealth : " In floods of life, in a storm of activity, she moves and works above and beneath, working and weaving, an endless motion, birth and death, an infinite ocean, a changeful web, a glowing life ; she plies at the roaring loom of time and weaves a living garment for God." 5. Wealth of Beauty. To many, however, animal life is impressive not so much because of its amazing variety and numerical greatness, nor because of its intellectual suggestiveness and practical utility, but chiefly on account of its beauty. This is to be seen and felt rather than described or talked about. The beauty of animals, in which we all delight, is usually in form, or in colour, or in movement. Especially in the simplest animals, the beauty of form is often comparable to that of crystals ; witness the marvellous architecture in flint and lime exhibited by the marine Protozoa, whose empty shells form the ooze of the great depths. In higher animals also an almost crystalline exactness of symmetry is often apparent, but we find more frequent illustration of graceful curves in form and feature, resulting in part from strenuous and healthful exercise, which moulds the body into beauty. Not a little of the colour of animals is due to the physical nature of the skin, which is often iridescent ; much, on the other hand, is due to the possession of pig- ments, which may either be of the nature of reserve-products, and then equivalent, let us say, to jewels, or of the nature of waste-products, and thus a literal "beauty for ashes." It is often supposed that plants excel animals in colour, but alike in the number and variety of pigments the reverse is true. Then as to movement, how much there is to admire ; the birds soaring, hovering, gliding, and diving; the monkey's gymnastics ; the bat's arbitrary evolutions ; the grace of the i6 The Study of Animal Life PART fleet stag; the dolphin gamboling in the waves; the lithe lizards which flash across the path and are gone, and the snake flowing like a silver river ; the buoyant swimming of fishes and all manner of aquatic animals ; the lobster darting backwards with a powerful tail-stroke across the pool ; the butterflies flitting like sunbeams among the flowers. But FIG. 3. Humming-birds {Florisiiga mellivora) visiting flowers. (From Belt.) are not all the delights of form and colour and movement expressed in the songs of the birds in spring ? I am quite willing to allow that this beauty is in one sense a relative quality, varying with the surroundings and education, and even ancestral history, of those who appreciate it. A flower which seems beautiful to a bee may be unattractive to a bird, a bird may choose her mate for qualities by no means winsome to human eyes, and a CHAP, i The Wealth of Life 17 dog may howl painfully at our sweet music. We call the apple - blossom and the butterfly's wings beautiful, partly because the rays of light, borne from them to our eyes, cause a pleasantly harmonious activity in our brains, partly because this awakens reminiscences of past pleasant experi- ences, partly for subtler reasons. Still, all healthy organisms are harmonious in form, and seldom if ever are their colours out of tone with their surroundings or with each other, a fact which suggests the truth of the Platonic conception that a living creature is harmonious because it is possessed by a single soul, the realisation of a single idea. The plants which seem to many eyes to have least beauty are those which have been deformed or discoloured by cultivation, or taken altogether out of their natural set- ting ; the only ugly animals are the products of domestica- tion and human interference on the one hand, or of disease on the other ; and the ugliest things are what may be called the excretions of civilisation, which are certainly not beauty for ashes, but productions by which the hues and colours of nature have been destroyed or smothered, where the natural harmony has been forcibly put out of tune in short, where a vicious taste has insisted on becoming inventive. CHAPTER II THE WEB OF LIFE I. Dependence upon Surroundings 2. Inter-relations of Plants and Animals 3. Relation of Animals to the Earth 4. Nutritive Relations 5. More Complex Interactions IN the filmy web of the spider, threads delicate but firm bind part to part, so that the whole system is made one. The quivering fly entangled in a corner betrays itself throughout the web ; often, it is felt rather than seen by the lurking spinner. So in the substantial fabric of the world part is bound to part. In wind and weather, or in the business of our life, we are daily made aware of results whose first conditions are remote, and chains of influence not difficult to demonstrate link man to beast, and flower to insect. The more we know of our surroundings, the more we realise the fact that nature is a vast system of linkages, that isolation is impossible. i. Dependence upon Surroundings. Every living body is built up of various arrangements of at least twelve " elements," viz. Oxygen, Hydrogen, Carbon, Nitrogen, Chlorine, Phosphorus, Sulphur, Magnesium, Calcium, Pot- assium, Sodium, and Iron. All these elements are spread throughout the whole world. By the magic touch of life they are built up into substances of great complexity and instability, substances very sensitive to impulses from, or changes in, their surroundings. It may be that living matter differs from dead matter in no other way than this. The CHAP, ii The Web of Life 19 varied forms of life crystallise out of their amorphous beginnings in a manner that we conceive to be analogous to the growth of a crystal within its solution. Further, we do not believe in a " vital force." The movements of living things are, like the movements of all matter, the expression of the world's energy, and illustrate the same laws. But to these matters we shall return in another chapter. Interesting, because of its sharply defined and far-reaching significance, and because the essential mass is so nearly infinitesimal, is the part played by iron in the story of life. For food-supply we are dependent upon animals and plants, and ultimately upon plants. But these cannot produce their valuable food-stuffs without the green colouring-matter in their leaves, by help of which they are able to utilise the energy of sunshine and the carbonic acid gas of the air. But this important green pigment (though itself perhaps free from any iron) cannot be formed in the plant unless there be, as there almost always is, some iron in the soil. Thus our whole life is based on iron. And all our supplies of energy, our powers of doing work either with our own hands and brains, or by the use of animals, or through the application of steam, are traceable if we follow them far enough to the sun, which is thus the source of the energy in all creatures. 2. Inter-relations of Plants and Animals. We often hear of the "balance of nature," a phrase of wide appli- cation, but very generally used to describe the mutual dependence of plants and animals. Every one will allow that most animals are more active than most plants, that the life of the former is on an average more intense and rapid than that of the latter. For all typical plants the materials and conditions of nutrition are found in water and salts absorbed by the roots, in carbonic acid gas absorbed by the leaves from the air, and in the energy of the sunlight which shines on the living matter through a screen of green pigment. Plants feed on very simple sub- stances, at a low chemical level, and their most char- acteristic transformation of energy is that by which the kinetic energy of the sunlight is changed into the potential 20 The Study of Animal Life PART i energy of the complex stuffs which animals eat or which we use as fuel. But animals feed on plants or on creatures like themselves, and are thus saved the expense of build- ing up food - stuffs from crude materials. Their most characteristic transformation of energy is that by which the power of complex chemical substances is used in locomotion and work. In so working, and eventually in dying, they form waste-products water and carbonic acid, ammonia and nitrates, and so on which may be again utilised by plants. How often is the inaccurate statement repeated "that animals take in oxygen and give out carbonic acid, whereas plants take in carbonic acid and give out oxygen " ! This is most misleading. It contrasts two entirely dis- tinct processes a breathing process in the animal with a feeding process in the plant. The edge is at once taken off the contrast when the student realises that plants and animals being both (though not equally) alive, must alike breathe. As they live the living matter of both is oxi- dised, like the fat of a burning candle ; in plant, in animal, in candle, oxygen passes in, as a condition of life or com- bustion, and carbonic acid gas passes out as a waste-pro- duct. Herein there is no difference except in degree between plant and animal. Each lives, and must therefore breathe. But the living of plants is less intense, therefore the breath- ing process is less marked. Moreover, in sunlight the respiration is disguised by an exactly reverse process peculiar to plants the feeding already noticed, by which carbonic acid gas is absorbed, its carbon retained, and part of its oxygen liberated. There is an old-fashioned experiment which illustrates the "balance of nature." In a glass globe, half- filled with water, are placed some minute 'water-plants and water- animals. The vessel is then sealed. As both the plants and the animals are absorbing oxygen and liberating car- bonic acid gas, it seems as if the little living world enclosed in the globe would soon end in death. But, as we have seen, the plants are able in sunlight to absorb carbonic acid and liberate oxygen, and if present in sufficient numbers will CHAP, ii The Web of Life 21 compensate both for their own breathing and for that of animals. Thus the result within the globe need not be suffocation, but harmonious prosperity. If the minute animals ate up all the plants, they would themselves die for lack of oxygen before they had eaten up one another, while if the plants smothered all the animals they would also in turn die away. Some such contingency is apt to spoil the experiment, the end of which may be a vessel of putrid water tenanted for a long time by the very simple colourless plants known as Bacteria, and at last not even by them. Nevertheless the " vivarium " experiment is both theoretically and practically possible. Now in nature there is, indeed, no closed vivarium, for there is no isolation and there is open air, and it is an exaggeration to talk as if our life were dependent on there being a proportionate number of plants and animals in the neighbourhood. Yet the " balance of nature " is a general fact of much importance, though the economical relations of part to part over a wide area are neither rigid nor precise. We have just mentioned the very simple plants called .Bacteria. Like moulds or fungi, they depend upon other organisms for their food, being without the green colouring stuff so important in the life of most plants. These very minute Bacteria are almost omnipresent ; in weakly animals and sometimes in strong ones too they thrive and multiply and cause death. They are our deadliest foes, but we should get rid of them more easily if we had greater love of sunlight, for this is their most potent, as well as most economical antagonist. But it is not to point out the obvious fact that a Bacterium may kill a king that we have here spoken of this class of plants ; it is to acknowledge their beneficence. They are the great cleansers of the world. Animals die, and Bacteria convert their corpses into simple substances, restoring to the soil what the plants, on which the animals fed, originally absorbed through their roots. Bacteria thus complete a wide circle ; they unite dead animal and living plant. For though many a plant thrives quite independently of animals on the raw materials of earth and air, others are demonstrably raising 22 The Study of Animal Life PART i the ashes of animals into a new life. A strange partner- ship between Bacteria on the one hand and leguminous and cereal plants on the other has recently been discovered. There seems much likelihood that with some plants of the orders just named Bacteria live in normal partner- ship. The legumes and cereals in question do not thrive well without their guests, nay more, it seems as if the Bacteria are able to make the free nitrogen of the air available for their hosts. 3. Relation of Animals to the Earth. Bacteria are extremely minute organisms, however, and stories of their industry are apt to sound unreal. But this cannot be said of earthworms. For these can be readily seen and watched, and their trails across the damp footpath, or their castings on the grass of lawn and meadow, are familiar to us all. They are distributed, in some form or other, over most regions of the globe ; and an idea of their abundance may be gained by making a nocturnal expedition with a lantern to any convenient green plot, where they may be seen in great numbers, some crawling about, others, with their tails in their holes, making slow circuits in search of leaves and vegetable debris. Darwin estimated that there are on an average 53,000 earthworms in an acre of garden ground, that 10 tons of soil per acre pass annually through their bodies, and that they bring up mould to the surface at the rate of 3 inches thickness in fifteen years. Hensen found in his garden 64 large worm-holes in 14 J square feet, and estimated the weight of the daily castings at about 2 cwts. in two and a half acres. In the open fields, how- ever, it seems to be only about half as much. But whether we take Darwin's estimate that the earthworms of England pass annually through their bodies about 320,000,000 tons of earth, or the more moderate calculations of Hensen, or our own observations in the garden, we must allow that the soil-making and soil -improving work of these animals is momentous. In Yorubaland, on the West African coast, earthworms (Siphonogaster) somewhat different from the common Lum- bricus are exceedingly numerous. From two separate square CHAP, ii The Web of Life 23 feet of land chosen at random, Mr. Alvan Millson collected the worm-casts of a season and found that they weighed when dry lof Ibs. At this rate about 62,233 tons of sub- soil would be brought in a year to the surface of each square mile, and it is also calculated that every particle of earth to the depth of two feet is brought to the surface once in 27 years. We do. not wonder that the district is fertile and healthy. Devouring the earth as they make their holes, which are often 4 or even 6 feet deep ; bruising the particles in their gizzards, and thus liberating the minute elements of the soil ; burying leaves and devouring them at leisure ; preparing the way by their burrowing for plant roots and rain-drops, and gradually covering the surface with their castings, worms have, in the history of the habitable earth, been most important factors in progress. Ploughers before the plough, they have made the earth fruitful. It is fair, however, to acknowledge that vegetable mould sometimes forms inde- pendently of earthworms, that some other animals which burrow or which devour dead plants must also help in the process, and that the constant rain of atmospheric dust, as Richthofen has especially noted, must not be overlooked. In 1777, Gilbert White wrote thus of the earthworms "The most insignificant insects and reptiles are of much more consequence and have much more influence in the economy of nature than the incurious are aware of. ... Earthworms, though in appearance a small and despicable link in the chain of Nature, yet, if lost, would make a lamentable chasm. . . . Worms seem to be the great promoters of vegetation, which would proceed but lamely without them, by boring, perforating, and loosening the soil, and rendering it pervious to rains and the fibres of plants ; by drawing straws and stalks of leaves and twigs into it ; and, most of all, by throwing up such infinite numbers of lumps of earth called worm- casts, which, being their excrement, is a fine manure for grain and grass. Worms probably provide new soil for hills and slopes where the rain washes the earth away ; and they affect slopes probably to avoid being flooded. . . . The earth without worms would soon become cold, hard-bound, and void of fermentation, and con- sequently sterile. . . . These hints we think proper to throw out, in order to set the inquisitive and discerning to work. A good mono- 24 The Study of Animal Life PART i graph of worms would afford much entertainment and information at the same time, and would open a large and new field in natural history." After a while the discerning did go to work, and Hensen published an important memoir in 1877, while Darwin's "good monograph" on the formation of vegetable mould appeared after about thirty years' observation in 1881 ; and now we all say with him, "It may be doubted whether there are many other animals which have played so important a part in the history of the world as have these lowly-organised creatures." Prof. Drummond, while admitting the supreme import- ance of the work of earthworms, eloquently pleads the claims of the Termite or White Ant as an agricultural agent. This insect, which dwelt upon the earth long before the true ants, is abundant in many countries, and notably in Tropical Africa. It ravages dead wood with great rapidity. "If a man lay down to sleep with a wooden leg, it would be a heap of sawdust in the morning," while houses and decaying forest trees, furniture and fences, fall under the jaws of the hungry Termites. These fell workers are blind and live underground ; for fear of their enemies they dare not show face, and yet without coming out of their ground they cannot live. " How do they solve the difficulty? They take the ground out along with them. I have seen white ants working on the top of a high tree, and yet they were underground. They took up some of the ground with them to the tree-top. They construct tunnels which run from beneath the soil up the sides of trees and posts ; grain after grain is carried from beneath and mortared with a sticky secretion into a reddish sandpaper-like tube ; this is rapidly ex- tended to a great height even of 30 feet from the ground till some dead branch is reached. Now as many trees in a forest are thus plastered with tunnels, and as there are besides elaborate subterranean galleries and huge obelisk-like ant-hills, sometimes 10-15 feet high, it must be granted that the Termites, like the earthworms, keep the soil circulating. The earth-tubes crumble to dust, which is scattered by the wind ; the rains lash the forests and soils with fury and wash off the loosened grains to swell the alluvium of a distant valley." CHAP, ii The Web of Life 25 The influences of plants and animals on the earth are manifold. The sea- weeds cling around the shores and lessen the shock of the breakers. The lichens eat slowly into the stones, sending their fine threads beneath the sur- face as thickly sometimes " as grass-roots in a meadow-land," so that the skin of the rock is gradually weathered away. On the moor the mosses form huge sponges, which mitigate floods and keep the "streams flowing in days of drought. Many little plants smooth away the wrinkles on the earth's face, and adorn her with jewels ; others have caught and stored the sunshine, hidden its power in strange guise in the earth, and our hearths with their smouldering peat or glowing coal are warmed by the sunlight of ancient summers. The grass which began to grow in comparatively modern (i.e. Tertiary) times has made the earth a fit home for flocks and herds, and protects it like a garment ; the forests affect the rainfall and temper the climate, besides sheltering multi- tudes of living things, to some of whom every blow of the axe is a death-knell. Indeed, no plant from Bacterium to oak tree either lives or dies to itself, or is without its influence on earth and beast and man. There are many animals besides worms which influence the earth by no means slightly. Thus, to take the minus side of the account first, we see the crayfish and their enemies the water-voles burrowing by the river banks and doing no little damage to the land, assisting in that process by which the surface of continents tends gradually to diminish. So along the shores in the harder substance of the rocks there are numerous borers, like the Pholad bivalves, whose work of disintegration is individually slight, but in sum-total great. More conspicuous, however, is the work of the beavers, who, by cutting down trees, building dams, digging canals, have cleared away forests, flooded low grounds, and changed the aspect of even large tracts of country. Then, as every one knows, there are injuri- ous insects innumerable, whose influence on vegetation, on other animals, and on the prosperity of nations, is often disastrously great. But, on the other hand, animals cease not to pay their 26 The Study of Animal Life PART i filial debts to mother earth. We see life rising like a mist in the sea, lowly creatures living in shells that are like mosques of lime and flint, dying in due season, and sinking gently to find a grave in the ooze. We see the submarine volcano top, which did not reach the surface of the ocean, slowly raised by the rainfall of countless small shells. Inch by inch for myriads of years, the snow-drift of dead shells forms a patient preparation for the coral island. The tiniest, hardly bigger than the wind-blown dust, form when added together the strongest foundation in the world. The vast whale skeleton falls, but melts away till only the ear- bones are left. Of the ruthless gristly shark nothing stays but teeth. The sea-butterflies (Pteropods), with their frail shells, are mightier than these, and perhaps the microscopic atomies are strongest of all. The pile slowly rises, and the exquisite fragments are cemented into a stable foundation for the future city of corals. At length, when the height at which they can live is reached, coral germs moor themselves to the sides of the raised mound, and begin a new life on the shoulders of death. They spread in brightly coloured festoons, and have often been likened to flowers. The waste salts of their living perhaps unite with the gypsum of the sea-water, at any rate in some way the originally soft young corals acquire strong shells of carbonate of lime. Sluggish creatures they, living in calcareous castles of indolence ! In silence they spread, and crowd and smother one another in a struggle for stand- ing-room. The dead forms, partly dissolved and cemented, become a broad and solid base for higher and higher growth. At a certain height the action of the breakers begins, great severed masses are piled up or roll down the sloping sides. Clear daylight at last is reached, the mound rises above the water. The foundations are ever broadened, as vigorously out-growing masses succumb to the brunt of the waves and tumble downwards. Within the surface -circle weathering makes a soil, and birds resting there with weary wings, or perhaps dying, leave many seeds of plants the begin- nings of another life. The waves cast up forms of dormant life which have floated from afar, and a ter- CHAP, ii The Web of Life 27 restrial fauna and flora begin. It is a strange and beautiful story, dead shells of the tenderest beauty on the rugged shoulders of the volcano ; corals like meadow flowers on the graveyard of the ooze ; at last plants and trees, the hum of insects and the song of birds, over the coral island. 4. Nutritive Relations. What we may call " nutritive chains " connect many forms of life higher animals feed- ing upon lower through long series, the records of which sound like the story of "The House that Jack built." On land and on the shore these series are usually short, for plants are abundant, and the carnivores feed on the vegetarians. In the open sea, where there is less vegeta- tion, and in the great depths, where there is none, carni- vore preys upon carnivore throughout long series fish feeds upon fish, fish upon crustacean, crustacean upon worm, worm on debris. Disease or disaster in one link affects the whole chain. A parasitic insect, we are told, has killed off the wild horses and cattle in Paraguay, thereby influencing the vegetation, thereby the insects, thereby the birds. Birds of prey and small mammals so-called "vermin" are killed off in order to preserve the grouse, yet this interference seems in part to defeat itself by making the survival of weak and diseased birds unnaturally easy, and epidemics of grouse- disease on this account the more prevalent. A craze of vanity or gluttony leads men to slaughter small insect-eating birds, but the punishment falls unluckily on the wrong shoulders when the insects which the birds would have kept down increase in unchecked numbers, and destroy the crops of grain and fruit. In a fuel -famine men have sometimes been forced to cut down the woods which clothe the sides of a valley, an action repented of when the rain-storms wash the hills to skeletons, when the valley is flooded and the local climate altered, and when the birds robbed of their shelter leave the district to be ravaged by caterpillar and fly. American entomologists have proved that the ravages of destructive insects may be checked by importing and fostering their natural enemies, and on the other hand, the sparrows which have established themselves in the States 28 The Study of Animal Life PART i have in some districts driven away the titmice and thus favoured the survival of injurious caterpillars. 5. More Complex Interactions. The flowering plants and the higher insects have grown up throughout long ages together, in alternate influence and mutual per- fecting. They now exhibit a notable degree of mutual dependence ; the insects are adapted for sipping the nectar from the blossoms ; the flowers are fitted for giving or receiving the fertilising golden dust or pollen which their visitors, often quite unconsciously, carry from plant to plant. The mouth organs of the insects have to be interpreted in relation to the flowers which they visit ; while the latter show structures which may be spoken of as the " footprints " of the insects. So exact is the mutual adaptation that Darwin ventured to prophesy from the existence of a Madagascar orchid with a nectar- spur 1 1 inches long, that a butterfly would be found in the same locality with a suctorial proboscis long enough to drain the cup ; and Forbes confirmed the prediction by discovering the insect. As information on the relations of flowers and insects is readily attainable, and as the subject will be discussed in the volume on Botany, it is sufficient here to notice that, so far as we can infer from the history half hidden in the rocks, the floral world must have received a marked impulse when bees and other flower-visiting insects appeared ; that for the successful propagation of flowering plants it is advantageous that pollen should be carried from one indi- vidual to another, in other words, that cross -fertilisation should be effected ; and that, for the great majority of flowering plants, this is clone through the agency of insects. How plants became bright in colour, fragrant in scent, rich in nectar, we cannot here discuss ; the fact that they are so is evident, while it is also certain that insects are attracted by the colour, the scent, and the sweets. Nor can there be any hesitation in drawing the inference that the flowers which attracted insects with most success, and insects which got most out of the flowers, would, ipso facto, succeed better in life. CHAP, ii The Web of Life 29 No illustration of the web of life can be better than the most familiar one, in which Darwin traced the links of influence between cats and clover. If the possible seeds in the flowers of the purple clover are to become real seeds, they must be fertilised by the golden dust or pollen from some adjacent clover plants. But as this pollen is uncon- sciously carried from flower to flower by the humble-bees, the proposition must t>e granted that the more humble-bees, the better next year's clover crop. The humble-bees, how- ever, have their enemies in the field-mice, which lose no opportunity of destroying the combs ; so that the fewer field-mice, the -more humble-bees, and the better next year's clover crop. In the neighbourhood of villages, however, it is well known that the cats make as effective war on the field-mice as the latter do on the bees. So that next year's crop of purple clover is influenced by the number of humble- bees, which varies with the number of field-mice, that is to say, with the abundance of cats ; or, to go a step farther, with the number of lonely ladies in the village. It should be noted, however, that according to Mr. James Sime there were abundant fertile clover crops in New Zealand before there were any humble-bees in that island. Indeed, many think that the necessity of cross-fertilisation has been exaggerated. Not all insects, however, are welcome visitors to plants ; there are unbidden guests who do harm. To their visits, however, there are often obstacles. Stiff hairs, impassably slippery or viscid stems, moats in which the intruders drown, and other structural peculiarities, whose origin may have had no reference to insects, often justify themselves by saving the plant. Even more interesting, however, is the preservation of some acacias and other shrubs by a bodyguard of ants, which, innocent themselves, ward off the attacks of the deadly leaf-cutters. In some cases the bodyguard has become almost hereditarily accustomed to the plants, and the plants to them, for they are found in constant companionship, and the plants exhibit structures which look almost as if they had been made as shelters for the ants. On some of our European trees similar little homes or domatia constantly occur, and shelter small 30 The Study of Animal Life TART i insects which do no harm to the trees, but cleanse them from injurious fungi. In many ways plants are saved from the appetite of animals. The nettle has poisonous hairs ; thistles, furze, and holly are covered with spines ; the hawthorn has its thorns and the rose its prickles ; some have repulsive odours ; others contain oils, acids, fer- ments, and poisons which many animals dislike ; the cuckoo-pint {Arum) is full of little crystals which make our lips smart if we nibble a leaf. In our studies of plants we endeavour to find out what these qualities primarily mean to their possessors ; here we think rather of their secondary significance as protections against animals. For though snails ravage all the plants in a district ex- cept those which are repulsive, the snails are at most only the second- FIG. 4. Acacia (A. spharocephala), with hoi- ary factors in the CVOlu- (After h SchLperJ hlch *"*' find shGher ' tion of the repulsive qualities. The strange inter-relations between plants and animals are again illustrated by the carnivorous, generally insecti- vorous, plants. It is not our business to discuss the original or primary import of the pitchers of pitcher-plants CHAP, ii The Web of Life 31 or of the mobile and sensitive leaves of Venus' Fly-Trap ; nowadays, at any rate, insects are attracted to them, captured by them, and used. Let us take only one case, that of the common Bladderwort (Utricularia). Many of the leaflets of this plant, which floats in summer in the marsh pond, are modified into little bladders, so fashioned that minute " water-fleas " which swarm in every corner of the pool can readily enter them, but can in no wise get out again. The small entrance is guarded by a valve or door, which opens inwards, but allows no egress. The little crusta- ceans are attracted by some mucilage made by the leaves, or sometimes perhaps by sheer curiosity ; they enter and cannot return ; they die, and their ddbris is absorbed by the leaf. Again, in regard to distribution, there are numerous relations between organisms. Spiny fruits like those of Jack-run-the-hedge adhere to animals, and are borne from place to place ; and minute water-plants and animals are carried from one watercourse to another on the muddy feet of birds. Darwin removed a ball of mud from the leg of a bird, and from it fourscore seeds germinated. Not a bird can fall to the ground and die without sending a throb through a wide circle. A conception of these chains or circles of influence is important, not only for the sake of knowledge, but also as a guide in action. Thus, to take only one instance among a hundred, it may seem a far cry from a lady's toilet-table to the African slave-trade, but when we remember the ivory backs of the brushes, and how the slaves are mainly used for transporting the tusks of elephants a doomed race from the interior to the coast, the riddle is read, and the respon- sibility is obvious. Over a ploughed field in the summer morning we see the spider-webs in thousands glistening with mist-drops, and this is an emblem of the intricacy of the threads in the web of life to be seen more and more as our eyes grow clear. Or, is not the face of nature like the surface of a gentle stream, where hundreds of dimpling circles touch and influence one another in an infinite com- plexity of action and reaction beyond the ken of the wisest ? CHAPTER III THE STRUGGLE OF LIFE I. Nature and Extent of the Struggle 2. Armour and Weapons 3. Different Forms of Struggle 4. Cruelty of the Struggle i. Nature and Extent of the Struggle. If we realise what is meant by the " web of life," the recognition of the " struggle for existence " cannot be difficult. Animals do not live in isolation, neither do they always pursue paths of peace. Nature is not like a menagerie where beast is separated from beast by iron bars, neither is it a melee such as would result if the bars of all the cages were at once removed. It is not a continuous Waterloo, nor yet an amiable compromise between weaklings. The truth lies between these extremes. In most places where animals abound there is struggle. This may be silent and yet decisive, real without being very cruel, or it may be full of both noise and bloodshed. This struggle is very old ; it is older than the conflicts of men, older than the ravin of tooth and claw, it is as old as life. The struggle is often very keen often for life or death. But though few animals escape experience of the battlefield and for some there seems no discharge from this war we must not misinterpret nature as "a continual free-fight." One naturalist says that all nature breathes a hymn of love, but he is an optimist under sunny southern skies ; another compares nature to a huge gladiatorial show with a plethora of fighters, but he speaks as a pes- CHAP, in The Struggle of Life 33 simist from amid the din of individualistic competition. Nature is full of struggle and fear, but the struggle is sometimes outdone by sacrifice, and the fear is sometimes cast out by love. We must be careful to remember Darwin's proviso that he used the phrase "struggle for existence " " in a large and metaphorical sense, including the dependence of one being on another, and including (which is more important) not only the life of the individual, but success in leaving progeny." He also acknowledged the importance of mutual aid, sociability, and sympathy among animals, though he did not carefully estimate the relative importance of competition on the one hand and sociability on the other. Discussing sympathy, Darwin wrote, "In however complex a manner this feeling may have originated, as it is one of high importance to all those animals which aid and defend one another, it will have been increased through natural selection ; for those communities which included the greatest number of the most sympathetic members would flourish best, and rear the greatest number of offspring." I should be sorry to misrepresent the opinions of any man, but after considerable study of modern Darwinian literature, I feel bound to join in the protest which others have raised against a tendency to narrow Darwin's conception of " the struggle for existence," by exaggerating the occurrence of internecine competitive struggle. Thus Huxley says, " Life was a continuous free- fight, and beyond the limited and temporary relations of the family, the Hobbesian war of each against all was the normal state of existence." Against which Kropotkine maintains that this "view of nature has as little claim to be taken as a scientific deduction as the opposite view of Rousseau, who saw in nature but love, peace, and harmony destroyed by the accession of man." . . . " Rousseau has committed the error of excluding the beak -and- claw fight from his thoughts, and Huxley is committing the opposite error ; but neither Rousseau's optimism nor Huxley's pessi- mism can be accepted as an impartial interpretation of nature." 2. Armour and Weapons. If you doubt the reality D 34 The Stiidy of Animal Life FART i of the struggle, take a survey of the different classes ot animals. Everywhere they brandish weapons or are forti- fied with armour. " The world," Diderot said, " is the abode of the strong." Even some of the simplest animals have offensive threads, prophetic of the poison- ous lassoes with which jellyfish and sea -anemones are equipped. Many worms have horny jaws; crustaceans have strong pincers ; many insects have stings, not to speak of mouth organs like surgical instruments ; spiders give poisonous bites ; snails have burglars' files; the cuttle- fish have strangling suckers and parrots' beaks. Among backboned animals we recall the teeth of the shark and the sword of the swordfish, the venomous fangs of serpents, the jaws of crocodiles, the beaks and talons of birds, the horns and hoofs and canines of mammals. Now we do not say that these and a hundred other weapons were from their first appearance weapons, indeed we know that most of them were not. But they are weapons now, and just as we would conclude that there was considerable struggle in a community where every man bore a revolver, we must draw a similar inference from the offensive equipment of animals. As to armoured beasts, we remember that shells of lime or flint occur in many of the simplest animals, that most sponges are so rich in spicules that they are too gritty to be pleasant eating, that corals are polypes withfn shells of lime, that many worms live in tubes, that the members of the starfish class are in varying degrees lime-clad, that crustaceans and insects are emphatically armoured animals, and that the majority of molluscs live in shells. So among backboned animals, how thoroughly bucklered were the fishes of the old red sandstone against hardly less effect- ive teeth, how the scales of modern fishes glitter, how securely the sturgeon swims with its coat of bony mail ! Amphibians are mostly weaponless and armourless, but reptiles are scaly animals par excellence, and the tortoise, for instance, lives in an almost impregnable citadel. Birds soar above pursuit, and mammals are swift and strong, but among the latter the armadillos have bony shields of CHAP, in The Struggle of Life 35 marvellous strength, and hedgehog and porcupine have their hair hardened into spines and quills. Now we do not say that all these structures were from the first of the nature of armour, indeed they admit of other explanations, but that they serve as armour now there can be no doubt. And just as we conclude that a man would not wear a chain shirt without due reason, so we argue from the prevalence of animal" armour to the reality of struggle. For a moment let me delay to explain the two saving- clauses which I have inserted. The pincers of a crab are modified legs, the sting of a bee has probably the same origin, and it is likely that most weapons originally served some other than offensive purpose. We hear of spears becoming pruning-hooks ; the reverse has sometimes been true alike of animals and of men. By sheer use a structure not originally a weapon became strong to slay ; for there is a profound biological truth in the French proverb : "A force de forger on dement for geron? And again as to armour, it is, or was, well known that a boy's hand often smitten by the " tawse " became callous as to its epidermis. Now that callousness was not a device providential or otherwise to save the youth from the pains of chastisement, and yet it had that effect. By bearing blows one naturally and necessarily becomes thick-skinned. Moreover, the epidermic callousness referred to might be acquired by work or play altogether apart from school discipline, though it might also be the effect of the blows. In the same way many structures which are most useful as armour may be the " mechanical " or natural results of what they afterwards help to obviate, or they may arise quite apart from their future significance. 3. Different Forms of Struggle. If you ask why animals do not live at peace, I answer, more Scottico^ Why do not we ? The desires of animals conflict with those of their neighbours, hence the struggle for bread and the competition for mates. Hunger and love solve the world's problems. Mouths have to be filled, but population tends locally and temporarily to outrun the means of subsistence, and the question "which mouths" CHAP, in The Struggle of Life 37 has to be decided sometimes by peaceful endeavour, as in migration, sometimes with teeth clenched or ravenous. Many animals are carnivorous, and must prey upon weaker forms, which do their best to resist. Mates also have to be won, and lover may fight with lover till death is stronger than both. But these struggles for food and for mates are often strivings rather than strife, nor is a recognition of the frequent keenness and fierceness of the competition incon- sistent with the recognition of mutual aid, sociability, and love. There is a third form of the struggle, that between an animal and its changeful surroundings. This also is a struggle without strife. Fellow competitors strive for their share of the limited means of subsistence ; between foes there is incessant thrust and parry ; in the courtship of mates there are many disappointed and worsted suitors ; over all are the shears of fate a changeful physical environment which has no mercy. An analysis of the various forms of struggle may be attempted as follows : (a) Between animals of the same kind which compete for similar food and other necessaries of life Struggle between For Food fellows-. (b) Between animals of different kinds, the one set striving to devour, the other set endeavouring to escape their foes, e.g. between carnivores and herbivores Struggle between foes. ( (c) Between the rival suitors for desired mates Struggle between rivals in [ love. For \ Foot- I ^ Between animals and changeful surround- hold j ings Struggle with fate. In most cases, besides the egoism or individualism, one must recognise the existence of altruism, parental love and sacrifice, mutual aid, care for others, and sociality. 38 The Study of Animal Life PART i Before we consider these different forms of struggle, let us notice the rapid multiplication of individuals which furnishes the material for what in " a wide and meta- phorical sense" may be called a "battlefield." A single Infusorian may be the ancestor of millions by the end of a week. A female aphis, often producing one offspring per hour for days together, might in a season be the ancestor of a progeny of atomies which would weigh down five hundred millions of stout men. " The roe of a cod contains sometimes nearly ten million eggs, and sup- posing each of these produced a young fish which arrived at maturity, the whole sea would immediately become a solid mass of closely packed codfish." The unchecked multiplication of a few mice or rabbits would soon leave no standing-room on earth. But fortunately, with the exception of the Infusorians, these multiplications do not occur. We have to thank the struggle in nature, and especially the physical environment, that they do not. The fable of Mirza's bridge is continually true, few get across. (a) It is often said that the struggle between fellows of the same kind and with the same needs is keenest of all, but this is rather an assumption than an induction from facts. The widespread opinion is partly due to an a priori con- sideration of the problem, partly to that anthropomorphism which so easily besets us. We transfer to the animal world our own experience of keen competition with fellows of the same caste, and in so doing are probably unjust. Thus Mr. Grant Allen says " The baker does not fear the competition of the butcher in the struggle for life ; it is the competition of the other bakers that sometimes inexorably crushes him out of existence. ... In this way the great enemies of the individual herbivores are not the carnivores, but the other herbivores. . . . It is not so much the battle between the tiger and the antelope, between the wolf and the bison, between the snake and the bird, that ultimately results in natural selection or survival of the fittest, as the struggle between tiger and tiger, between bison and bison, between snake and snake, between antelope and antelope. . . . Homo homini lupus, says the old proverb, and so, we may add, in a wider sense, lupus lupo CHAP, in The Struggle of Life 39 hiptis, also. . . . The struggle is fierce between allied kinds, and fiercest of all between individual members of the same species." I have quoted these sentences because they are clearly and cleverly expressed, after the manner of Grant Allen, but I do not believe that they are true statements of facts. The evidence is very unsatisfactory. In his paragraph sum- marised as " struggle, for life most severe between indi- viduals and varieties of the same species ; often severe between species of the same genus," Darwin gave five illustrations : one species of swallow is said to have ousted another in North America, the missel-thrush has increased in Scotland at the expense of the song-thrush, the brown rat displaces the black rat, the small Asiatic cockroach drives its great congener before it, the hive -bee imported to Australia is rapidly exterminating- the small, stingless native bee. But the cogency of these instances may be disputed : thus what is said about the thrushes is denied by Professor Newton. And on the other hand, we know that reindeer, beavers, lemming, buffaloes and many other animals migrate when the means of subsistence are unequal to the demands of the population, and there are other peaceful devices by which animals have discovered a way out of a situation in which a life-and-death struggle might seem inevitable. Very instructive is the fact that beavers, when too numerous in one locality, divide into two parties and migrate up and down stream. The old proverb which Grant Allen quotes, Homo homini lupus, appears to me a libellous inaccuracy ; the extension of the libel to the animal world has certainly not been justified by careful induction. For a discussion of the alleged competition between fellows, I refer, and that with pleasure and grati- tude, to Kropotkine's articles on " Mutual Aid among Animals," Nineteenth Century ', September and November 1890. (/;) Of the struggle between foes differing widely in kind little need be said. It is very apparent, especially in wild countries. Carnivores prey upon herbivores, which some- times unite in successful resistance. Birds of prey devour 40 TJie Study of Animal Life PART i small mammals, and sometimes have to fight hard for their booty. Reptiles also have their battles witness the combats between snake and mongoose. In many cases, however, carnivorous animals depend upon small fry ; thus many birds feed on fishes, insects, and worms, and many fishes live on minute crustaceans. In such cases the term FIG. 6. Weasel attacking a grouse. (From St. John's Wild Sports.} struggle must again be used " in a wide and metaphorical sense." (c) In a great number of cases there is between rival males a contest for the possession of the females, a competition in which beauty and winsomeness are sometimes as im- portant as strength. Contrast the musical competition between rival songsters with the fierce combats of the stags. CHAP, in The Struggle of Life 41 Many animals are not monogamous, and this causes strife ; a male seal, for instance, guards his harem with ferocity. (d) Finally, physical nature is quite careless of life. Changes of medium, temperature, and moisture, continually occur, and the animals flee for their lives, adapt themselves to new conditions, or perish. Cataclysms are rare, but changes are common, and especially in such schools of experience as the sea'-shore we may study how vicissitude has its victims or its victors. The struggle with Fate, that is to say, with changeful surroundings, is more pleasant to contemplate than the other kinds of struggle, for at the rigid mercilessness of physical nature we shudder less than at the cruel competi- tion between living things, and we are pleased with the devices by which animals keep their foothold against wind and weather, storm and tide, drought and cold. One illus- tration must suffice : drought is common, pools are dried up, the inhabitants are left to perish. But often the organism draws itself together, sweats off a protective sheath, which is not a shroud, and waits until the rain refreshes the pools. Not the simplest animals only, but some of comparatively high degree, are thus able to survive desiccation. The simplest animals encyst, and may be blown about by the wind, but they rest where moisture moors them, and are soon as lively as ever. Leaping a long way upwards, we find that the mud- fish (Protopterus) can be transported from Africa to Northern Europe, dormant, yet alive, within its ball of clay. We do not believe in toads appear- ing out of marble mantelpieces, and a palaeontologist will but smile if you tell him of a frog which emerged from an intact piece of old red sandstone, but amphibians may remain for a long time dormant either in the mud of their native pools or in some out-of-the-way chink whither they had wandered in their fearsome youth. A shop which had once been used in the preparation of bone-dust was after prolonged emptiness reinstated in a new capacity. But it was soon fearfully infested with mites (Glytiphagus), which had been harboured in crevices in a strange state of dry dormancy. Every mite had in a sense 42 The Study of Animal Life PART i died, but remnant cells in the body of each had clubbed together in a life-preserving union so effective that a return of prosperity was followed by a reconstitution of mites and by a plague of them. Of course great caution must be exercised with regard to all such stories, as well as in regard to the toads within stones. Of common little animals known as Rotifers, it is often said, and sometimes rightly, that they can survive prolonged desiccation. In a small pool on the top of a granite block, there flourished a family of these Rotifers. Now this little pool was period- ically swept dry by the wind, and in the hollow there remained only a scum of dust. But when the rain returned and filled the pool, there were the Rotifers as lively as ever. What inference was more natural than that the Rotifers survived the desiccation, and lay dormant till moisture returned ? But Professor Zacharias thought he would like to observe the actual revivification, and taking some of the dusty scum home, placed it under his micro- scope on a moist slide, and waited results. There were the corpses of the Rotifers plain enough, but they did not revive even in abundant moisture. What was the explanation ? The eggs of these Rotifers survived, they developed rapidly, they reinstated the family. And of course it is much easier to understand how single cells, as eggs are, could survive being dried up, while their much more complex parents perished. I do not suggest that no Rotifers can survive desiccation, it is certain that some do ; but the story I have told shows the need of caution. There is no doubt, moreover, that certain simple "worms," known as "paste- eels," " vinegar-eels," etc., from their frequent occurrence in such substances, can survive desiccation for many years. Repeated experiments have shown that they can lie dormant for as long as, but not longer than, fourteen years ! and it is interesting to notice that the more prolonged the period of desiccation has been, the longer do these threadworms take to revive after moisture has been supplied. It seems as if the life retreated further and further, till at length it may retreat beyond recall. In regard to plants there are many similar facts, for though accounts of the germination CHAP, in The Struggle of Life 43 of seeds from the mummies of the pyramids, or from the graves of the Incas, are far from satisfactory, there is no doubt that seeds of cereals and leguminous plants may retain their life in a dormant state for years, or even for tens of years. But desiccation is only one illustration out of a score of the manner in which animals keep their foothold against fate. I need hardly say that they are often unsuccessful ; the individual has often fearful odds against it. How many winged seeds out of a thousand reach a fit resting-place where they may germinate ? Professor Mobius says that out of a million oyster embryos only one individual grows up, a mortality due to untoward currents and surroundings, as well as to hungry mouths. Yet the average number of thistles and oysters tends to continue, " So careful of the type she seems, so careless of the single life." Yet though the average usually remains constant, there is no use trying to ignore, what Richard Jefferies sometimes exaggerated, that the physical fates are cruel to life. But how much wisdom have they drilled into us ? " For life is not as idle ore, But iron dug from central gloom, And heated hot with burning fears, And dipt in baths of hissing tears, And battered by the shocks of doom To shape and use." 4. Cruelty of the Struggle. Opinions differ much as to the cruelty of the " struggle for existence," and the question is one of interest and importance. Alfred Russel Wallace and others try to persuade us that our conception of the " cruelty of nature" is an anthropomorphism; that, like Balbus, animals do not fear death ; that the rabbit rather enjoys a run before the fox ; that thrilling pain soon brings its own anaesthetic ; that violent death has its pleasures, and starvation its excitement. Mr. Wallace, who speaks with the authority of long and wide ex- perience, enters a vigorous protest against Professor Huxley's description of the myriads of generations of 44 The Study of Animal Life PART i herbivorous animals " which have been tormented and devoured by carnivores " ; of both alike " subject to all the miseries incidental to old age, disease, and over-multiplica- tion " ; of the " more or less enduring suffering " which is the meed of both vanquished and victor ; of the whole creation groaning in pain. " There is good reason to believe," says Mr. Wallace, " that the supposed torments and miseries of animals have little real existence, but are the reflection of the imagined sensations of cultivated men and women in similar circumstances ; and that the amount of actual suffering caused by the struggle for existence among animals is altogether insignificant." " Animals are spared from the pain of anticipating death ; violent deaths, if not too prolonged, are painless and easy ; neither do those which die of cold or hunger suffer much ; the popular idea of the struggle for existence entailing misery and pain on the animal world is the very reverse of the truth." He concludes by quoting the conclusion of Darwin's chapter on the struggle for existence : " When we reflect on this struggle, we may console ourselves with the full belief that the war of nature is not incessant, that no fear is felt, that death is generally prompt, and that the vigorous, the healthy, and the happy survive and multiply." Yet it was Darwin who confessed that he found in the world "too much misery." We have so little security in appreciating the real life the mental and physical pain or happiness of animals, that there is apt to be exaggeration on both sides, according as a pessimistic or an optimistic mood predominates. I there- fore leave it to be settled by your own observation whether hunted and captured, dying and starving, maimed and half- frozen animals have to endure " an altogether insignificant amount of actual suffering in the struggle for existence." But I think we must admit that there is much truth in what Mr. Wallace urges. Moreover, the term cruelty can hardly be used with accuracy when the involved infliction of pain is necessary. In many cases the carnivores are less " cruel " to their victims than we are to our domesti- cated animals. We must also remember that the " struggle CHAP, in The Struggle of Life 45 for existence " is often applicable only in its " wide and metaphorical sense." And it is fair to balance the happiness and mutual helpfulness of animals against the pain and deathful competition which undoubtedly exist. What we must protest against is that one-sided inter- pretation according to which individualistic competition is nature's sole method of progress. We are told that animals have got on by their struggle for individual ends ; that they have made progress on the corpses of their fellows, by a " blood and iron " competition in which each looks out for himself, and extinction besets the hindmost. To those who accept this interpretation the means employed seem justified by the results attained. But it is only in after-dinner talk that we can slur over whatever there is of pain and cruelty, overcrowding and starvation, hate and individualism, by saying complacently that they are justified in us their children; that we can rest satisfied that what has been called "a scheme of salvation for the elect by the damnation of the vast majority " is a true statement of the facts ; that we can seriously accept a one-sided account of nature's regime as a justification of our own ethical and economic practice. The conclusions, which I shall afterwards seek to substantiate, are, that the struggle for existence, with its associated natural selection, often involves cruelty, but certainly does not always do so ; that joy and happiness, helpfulness and co-operation, love and sacrifice, are also facts of nature, that they also are justified by natural selection ; that the precise nature of the means employed and ends attained must be carefully considered when we seek from the records of animal evolution support or justification for human conduct ; and that the tragic chapters in the history of animals (and of men) must be philosophically considered in such light as we can gather from what we know of the whole book. CHAPTER IV SHIFTS FOR A LIVING I. Insulation 2. Concealment 3. Parasitism 4. General Re- semblance to Surroundings 5. Variable Colouring 6. Rapid Change of Colour 7. Special Protective Resemblance 8. Warning Colours 9. Mimicry 10. Masking n. Com- bination of Advantageous Qtialities 12. Surrender of Paris GRANTING the struggle with fellows, foes, and fate, we are led by force of sympathy as well as of logic to think of the shifts for a living which tend to be evolved in such con- ditions, and also of some other ways by which animals escape from the intensity of the struggle. i. Insulation. Some animals have got out of the struggle through no merit of their own, but as the result of geological changes which have insulated them from their enemies. Thus, in Cretaceous times probably, the marsupials which inhabited the Australasian region were insulated. In that region they were then the only re- presentatives of Mammalia, and so, excepting the " native dog," some rodents and bats, and more modern imports, they still continue to be. By their insulation they were saved from that contest with stronger mammals in which all the marsupials left on the other continents were exterminated, with the exception of the opossums, which hide in American forests. A similar geological insulation accounts for the large number of lemurs in the island of Madagascar. CHAP, iv Shifts for a Living 47 2. Concealment. A change of habitat and mode of life is often as significant for animals as it is for men. It is easy to understand how mammals which passed from terrestrial to more or less aquatic life, for instance beaver and polar bear, seals, and perhaps whales, would enjoy a period of relative immunity after the awkward time of transition was over. So, too, many must have passed from the battlefield of the sea -shore to relative peace on land or in the deep-sea. In a change from open air to underground life, illustrated for instance in the mole, many animals have sought and found safety, and the change seems even now in progress, as in the New Zealand parrot Stringops, which, having lost the power of flight, has taken to burrowing. Similarly the power of flight must have helped insects, some ancient saurians, and birds out of many a scrape, though it cannot be doubted that this transition, and also that from diurnal to nocturnal habits, often brought only a temporary relief. 3. Parasitism. From the simple Protozoa up to the beginning of the backboned series, we find illustrations of animals which have taken to a thievish existence as unbidden guests in or on other organisms. Flukes, tapeworms, and some other "worms," many crustaceans, insects, and mites, are the most notable. Few animals are free from some kind of parasite. There are various grades of parasitism ; there are temporary and permanent, external and internal, very degenerate, and very slightly affected parasites. Some- times the adults are parasitic while the young are free -liv- ing, sometimes the reverse is true ; sometimes the parasite completes its life in one host, often it reaches maturity only after the host in which its youth has been passed is de- voured by another. In many cases the habit was probably begun by the females, which seek shelter during the period of egg-laying; in not a few crustaceans and insects the females alone are parasitic. Most often, in all probability, hunger and the search for shelter led to the establishment of the thievish habit. Now, the advantages gained by a thoroughgoing parasite are great safety, warmth, abund- ant food, in short, " complete material well-being." But 48 The Study of Animal Life PART i there is another aspect of the case. Parasitism tends to be followed by degeneration of appendages, food - canal, sense-organs, nervous system, and other structures, the possession and use of which make life worth living. More- over, though the reproductive system never degenerates, the odds are often many against an embryo reaching a fit host or attaining maturity. Thus Leuckart calculates that a tapeworm embryo has only about I chance in 83,000,000 of becoming a tapeworm, and one cannot be sorry that its chance is not greater. In illustration of the degenera- tion which is often associated with parasitism, and varies as the habit is more or less predominant, take the case of Sacculina a crustacean usually ranked along with bar- nacles and acorn-shells. It begins its life as a minute free "nauplius," with three pairs of appendages, a short food- canal, an eye, a small brain, and some other structures characteristic of many young crustaceans. In spite of this promiseful beginning, the young Sacculina becomes a para- site, first within the body, and finally under the tail, of a crab. Attached by absorptive suckers to its host, and often doing no slight damage, it degenerates into an oval sac, almost without trace of its former structure, with reproductive system alone well developed. Yet the degeneration is seldom so great as this, and it is fair to state that many parasites, especially those which remain as external hangers-on, seem to be but slightly affected by their lazy thievish habit ; nor can it be denied that most are well adapted to the conditions of their life. But on the whole the parasitic life tends to degeneration, and is unprogress- ive. Meredith writes of Nature's sifting " Behold the life of ease, it drifts. The sharpened life commands its course : She winnows, winnows roughly, sifts, To dip her chosen in her source. Contention is the vital force Whence pluck they brain, her prize of gifts. " 4. General Resemblance to Surroundings. Many transparent and translucent blue animals are hardly CHAP, iv Shifts for a Living 49 visible in the sea ; white animals, such as the polar bear, the arctic fox, and the ptarmigan in its winter plumage, are inconspicuous upon the snow ; green animals, such as insects, tree-frogs, lizards, and snakes, hide among the leaves and herbage ;(. tawny animals harmonise with sandy soil ; and the hare escapes detection among the clods. So do spotted animals such as snakes and leopards live unseen in the interrupted light" of the forest, and the striped tiger is lost in the jungle. Even the eggs of birds are often well suited to the surroundings in which they are laid. There can be no doubt that this resemblance between the colour of an animal and that of its surroundings is sometimes of protective and also aggressive value in the struggle for existence, and where this is the case, natural selection would foster it, favouring with success those variations which were best adapted, and eliminating those which were conspicuous. But there are many instances of resemblance to sur- roundings which are hard to explain. Thus Dr. A. Seitz describes a restricted area of woodland in South Brazil, where the great majority of the insects were blue, although but a few miles off a red colour was dominant. He maintains that the facts cannot in this case be explained as due either to general protective resemblance or to mimicry. I have reduced what I had written in illustration of advantageous colouring of various kinds, because this exceedingly interesting subject has been treated in a readily available volume by one who has devoted much time and skill to its elucidation. Mr. E. B. Poulton's Colours of Animals (International Science Series, London, 1890) is a fascinating volume, for which all interested in these aspects of natural history must be grateful. With this a forth- coming work (Animal Coloration^ London, 1892) by Mr. F. E. Beddard should be compared. 5. Variable Colouring. Some animals, such as the ptarmigan and the mountain-hare, become white in winter, and are thereby safer and warmer. In some cases it is certain that the pigmented feathers and hairs become white, in other cases the old feathers and hairs drop E 50 TJie Study of Animal Life PART i off and are replaced by white ones ; sometimes the whiteness is the result of both these processes. It is directly due to the formation of gas bubbles inside the hairs or feathers in sufficient quantity to antagonise the effect of any pigment that may be present, but in the case of new growths it is not likely that any pig- ment is formed. In some cases, e.g. Ross's lemming and the American hare (Lepus americamis\ it has been clearly shown that the change is due to the cold. It is likely that this acts somewhat indirectly upon the skin through the nervous system. We may therefore regard the change as a variation due to the environment, and it is at least possible that the permanent whiteness of some northern animals, e.g. the polar bear, is an acquired character of similar origin. There are many objections to the theory that the winter whiteness of arctic animals arose by the accumulation of small variations in individuals which, being slightly whiter than their neighbours, became dominant by natural selection, though there can be no doubt that the whiteness, however it arose, would be conserved like other advantageous variations. To several naturalists, but above all to Mr. Poulton, we are indebted for much precise information in regard to the variable colouring of many caterpillars and chrysalides. These adjust their colours to ithose of the surroundings, and even the cocoons are sometimes harmoniously coloured. There is no doubt that the variable colouring often has protective value. Mr. Poulton experimented with the caterpillars of the peacock butterfly (Vanessa to\ small tortoise-shell ( Vanessa urticce), garden whites (Pieris brassica and Pieris rapce), and many others. Caterpillars of the small tortoise-shell in black surroundings tend to be- come darker as pupae ; in a white environment the pupae are lighter ; in gilded boxes they tend to become golden. The surrounding colour seems to influence the caterpillar " during the twenty hours immediately preceding the last twelve hours of the larval state," " and this is probably the true meaning of the hours during which the caterpillar rests motionless on the surface upon which it will pupate." CHAP, iv Shifts for a Living 51 "It appears to be certain that it is the skin of the larva which is influenced by surrounding colours during the sensitive period, and it is probable that the effects are wrought through the medium of the nervous system." Accepting the facts that caterpillars are subtly affected by surrounding colours, so that the quiescent pupae har- monise with their environment, and that the adjustment has often protective value, we are led to inquire into the origin of this sensitiveness. That the change of colour is not a direct result of external influence is certain, but of the physiological nature of the changes we know little more than that it must be complex. It may be main- tained, that " the existing colours and markings are at any rate in part due to the accumulation through heredity of the indirect influence of the environment, working by means of the nervous system;" "to which it may be replied," Poulton continues, " that the whole use and meaning of the power of adjustment depends upon its freedom during the life of the individual ; any hereditary bias towards the colours of ancestors would at once destroy the utility of the power, which is essentially an adaptation to the fact that different individuals will probably meet with different environments. As long ago as 1873 Professor Meldola argued that this power of adjustment is adaptive, and to be explained by the operation of natural selection." Poulton's opinion seems to be, that the power of producing Variable colouring arose as a constitutional variation apart from the influence of the environment, that the power was fostered in the course of natural selection, and that its limits were in the same way more or less defined in adapta- tion to the most frequent habitat of the larvae before and during pupation. The other theory is that the power arose as the result of environmental influence, was accumu- lated by inheritance throughout generations, and was fostered like other profitable variations by natural selection. The question is whether the power arose in direct relation to environmental influence or not, whether external influence was or was not a primary factor in evolving the power of adapting colour, and in defining it within certain limits. 52 The Study of Animal Life PART i 6. Rapid Change of Colour. For ages the chamaeleon has been famous for its rapid and sometimes striking changes of colour. The members of the Old World genus Chamceleo quickly change from green to brown or other tints, but rather in response to physical irrita- tion and varying moods than in relation to change of situation and surrounding colours. So the American " chamaeleons " (Anolis) change, for instance, from emerald to bronze under the influence of excitement and various kinds of light. Their sensitiveness is exquisite ; " a pass- ing cloud may cause the bright emerald to fade." Some- times they may be thus protected, for " when on the broad green leaves of the palmetto, they are with difficulty per- ceived, so exactly is the colour of the leaf counterfeited. But their dark shadow is very distinct from beneath." Most of the lizards have more or less of this colour-changing power, which depends on the contraction and expansion of the pigmented living matter of cells which lie in layers in the under-skin, and are controlled by nerves. In a widely different set of animals the cuttle-fishes the power of rapid colour-change is well illustrated. When a .cuttle-fish in a tank is provoked, or when one almost stranded on the beach struggles to free itself, or, most beautifully, when a number swim together in strange unison, flushes of colour spread over the body. The sight suggests the blushing of higher animals, in which nervous excitement passing from the centre along the peripheral nerves influ- ences the blood-supply in the skin ; but in colour-change the nervous thrills affect the pigment-containing cells or chroma- tophores, the living matter of which contracts or expands in response to stimulus. It must be allowed that the colour- change of cuttle-fish is oftenest an expression of nervous excitement, but in some cases it helps to conceal the animals. More interesting to us at present are those cases of colour -change in which animals respond to the hues of their surroundings. This has been observed in some Amphibians, such as tree-frogs ; in many fishes, such as plaice, stickleback, minnow, trout, Gobius rutkensparri, v Shifts for a Living 53 Serranus ; and in not a few crustaceans. The researches of Briicke, Lister, and Pouchet have thrown much light on the subject. Thus we know that the colour of surround- ings affects the animals through the eyes, for blind plaice, trout, and frogs do not change their tint. The nervous thrill passes from eye to brain, and thence extends, not down the main path of impulse the spinal cord but down the sympathetic chain. If this be cut, the colour-change does not take place. The sympathetic system is connected with nerves passing from the spinal cord to the skin, and it is along these that the impulse is further transmitted. The result is the contraction or expansion of the pigment in the skin-cells. Though the path by which the nervous influence passes from the eye to the skin is somewhat circuitous, "the change is often very rapid. As the resulting resemblance to surroundings is often precise, there can be no doubt that the peculiarity sometimes profits its possessors. 7. Special Protective Resemblance. The likeness between animals and their surroundings is often very precise, and includes form as well as colour. Thus some bright butter- flies, e.g. Kallima, are conspicuous in flight, but become precisely like brown withered leaves when they settle upon a branch and expose the under sides of their raised wings ; the leaf-insects (Phylliuin) have leaf-like wings and legs ; the " walking-sticks " (Phasmid