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If you are not located in the United States, you'll have -to check the laws of the country where you are located before using this ebook. - -Title: The Courtship of Animals - -Author: William Plane Pycraft - -Release Date: October 18, 2019 [EBook #60517] - -Language: English - -Character set encoding: UTF-8 - -*** START OF THIS PROJECT GUTENBERG EBOOK THE COURTSHIP OF ANIMALS *** - - - - -Produced by Turgut Dincer (This file was produced from -images made available by The Internet Archive) - - - - - - - - Hutchinson’s - - Nature - - Library - - - THE COURTSHIP OF ANIMALS - - [Illustration: Plate 1. - - LOVE-MAKING. - - Frontispiece.] - - - - - The - - Courtship of Animals - - BY - - W. P. PYCRAFT - - OF THE - - ZOOLOGICAL DEPARTMENT OF THE BRITISH MUSEUM: FELLOW OF THE ZOOLOGICAL - SOCIETY OF LONDON; ASSOCIATE OF THE LINNEAN SOCIETY: MEMBER OF THE - ROYAL ANTHROPOLOGICAL INSTITUTE; MEMBER OF THE BRITISH ORNITHOLOGISTS’ - UNION; HON. MEMBER OF THE AMERICAN ORNITHOLOGISTS’ UNION; ETC., ETC. - - Author of “A History of Birds,” “The Natural History Museum,” “Pads, - Paws and Claws,” “The Infancy of Animals,” etc., etc., etc. - - _With 40 Plates on art paper Containing over 80 Illustrations_ - - _THIRD EDITION_ - - LONDON - - HUTCHINSON & CO. - - PATERNOSTER ROW - - - I DEDICATE THIS VOLUME - - TO - - H. ELIOT HOWARD - - WHOSE OBSERVATIONS OF THE COURTSHIP OF BIRDS RECORDED IN HIS “HISTORY - OF THE BRITISH WARBLERS” CONSTITUTE A BEACON FOR ALL ENGAGED IN THE - STUDY OF ANIMAL BEHAVIOUR - - - - -PREFACE - - -That “one touch of Nature which makes the whole World kin” is surely -nowhere more obvious than in the “Courtship” of Animals. For the -“Beasts that Perish,” no less than Man himself, are stirred by the -same emotions; the Fever of Love runs as high in them as in ourselves; -and its modes of expression are not so different, though they may -superficially appear to be so. The nature of these differences and -their interpretation, it is the purpose of this book to set forth. - -Charles Darwin laid the foundation for the study of this phase of -Animal behaviour in his masterly work on the “Descent of Man,” a work -which has been much criticized and much misunderstood since Carlyle’s -crude abuse of it as the “Gospel of Dirt.” Darwin was the first to show -us that the fierce battles, and strange antics, which characterize so -many of the “Lower Orders of Creation” under the exaltation of the -Sexual emotions are manifestations fraught with tremendous consequences -to the race. - -The facts which he brought to light, and the discussions to which -they have given rise, have, however, unfortunately been too commonly -regarded as merely interesting to those who have a liking for Natural -History. - -This is a most unfortunate mistake. For such facts have a vitally -important bearing on the very problems of social well-being which now -loom so largely among us. “Reform” is in the air. Its protagonists -are busy amongst us with schemes for our regeneration, among which -“Sex-problems” are made to occupy a very conspicuous place. But no -good can come of their cogitations so long as they fail to realize the -springs of behaviour in this regard. The facts herein set down will, it -is hoped, help much towards this end. - -My labours in the preparation of these pages have been materially -lightened by the help and counsel of many friends. To them I desire now -to record my very grateful thanks. More especially am I indebted to my -friends Mr. H. Eliot Howard, Professor Lloyd Morgan and Mr. John Cooke. - -I must also thank those who have contributed towards the illustrations -which enliven these pages. The delightful Frontispiece, and many of the -plates scattered through this work, I owe to the generosity of Messrs. -Rowland Ward, Limited. The excellent rendering of the Birds of Paradise -adapted in part from the work of Mr. G. E. Lodge and the late J. G. -Keulemans, and partly drawn from specimens in the British Museum, is -the work of Mr. Roland Green. The very difficult, and less fascinating, -technical figures I owe to the skill of Mr. Philip Whelpley. The -wonderful photographs illustrating the “Display” of the Sun-bittern and -the Kagu were taken by my friend Mr. D. Seth-Smith. - -Finally I have to thank Mr. Roger Ingpen for the immense amount of -trouble which he has taken in seeing these pages through the press. - -W. P. Pycraft. - -October, 1913. - - - - -CONTENTS - - CHAPTER - - INTRODUCTION - - The nature of Life and its power of reproduction—The - stuff of which Life is made—The Emotions—The simplest - living things—Where is neither Birth nor Death yet the - Population increases—The First Marriage—The beginning - of sex—The two dominating instincts—The conditions - of survival—The Oyster’s narrow world—“Fiddling - work”—Amorousness—The superior Male—Where Death - begins—“Germ-plasm” and what it means—Sex and “Secondary - sexual Characters”—Some theories—“Hormones” what are - they? 1 - - - CHAPTER II - - “MANKIND IN THE MAKING” - - The use of the term “Courtship”—Primitive Man and - the Foundations of Society—“Amorousness” as a motive - force—Polygamy—Our half human ancestors—Standards of - Beauty—Disquieting signs 21 - - - CHAPTER III - - MAN’S COUSINS THE APES - - The Man-like Apes and their mode of Life—Their - “Courtships”—Musical Chimpanzees—How the Orang-utan - improves his voice—His likeness to Caliban—The truculent - visage of the Gorilla—“Ornament” in the lower Apes—The - Concerts of the Howler Monkeys 40 - - - CHAPTER IV - - AT DAGGERS DRAWN - - The Birth of Weapons—All Flesh is Grass—Utility and - Ornament—The Fever of Love—The “Challenge” of the - Deer—What it means-More about “Hormones”—“Hummel” - Stags—The Age of Deer—The “Courtship” of the Moose—Types - of Antlers—Antlered Females—Fighting Topi—The Lance - of the Oryx in the Lion’s Flanks—Happiness and - Hartebeestes—Odoriferous Suitors—The Bloody Sweat - of the Hippopotamus—The Elephant in Love—Concerning - Tusks—Polygamy 49 - - - CHAPTER V - - THE LION AND HIS KIN - - A Surprising Relationship—The Lion’s Mane—The - Sabre-toothed Tiger—Some Theories about Origins—Sea-lions - in Love—Some Strange Ornaments—Whales and Weapons 77 - - - CHAPTER VI - - COURTSHIP AMONG BIRDS - - Generalities—Darwin v. Wallace—The Peacock in his - Pride—The “Display” of the Peacock Pheasant—The - Splendour of the Argus Pheasant and the Marvel of - its Eyes—The Frill of the Amherst Pheasant—Birds of - Paradise in the Toils of Love—Inflated Suitors-Ruffs - and Reeves—Fearsome Weapons and their Uses—Birds which - dance-Musical Birds—The Bird’s Voice-box—The “Lek” of - the Capercaillie—Instruments of Percussion—The Curious - Performance of the Woodpecker 92 - - - CHAPTER VII - - THE SEXUAL SELECTION THEORY AS APPLIED TO BIRDS - - Where the Rôle of the Sexes is reversed—Polygamy and how - it is brought about—Coloration and Courtship—Instinctive - Actions—The Importance of Landed Possessions—The Meaning - of “Display”—The Springs of “Behaviour”—A New Light - on the Wild-duck—The “Display” of the Great-crested - Grebe—Some Neglected Factors 134 - - - CHAPTER VIII - - SOME “COLD-BLOODED” LOVERS - - The Courtship of the Crocodile—Amorous Lizards—Horned - Chameleons—A Flagellating Terrapin—The Frog that would - a-wooing go—Some Musical Frogs—Some marvellous instincts - in Newts 161 - - - CHAPTER IX - - LOVE-MAKING AMONG THE FISHES - - Germinal variations—Fishes and Mate-hunting—Some - Remarkable Sexual Differences displayed by the Teeth - of “Rays”—The Double-eyed Fish—The Coloration of the - Dragonet—Some Curious Facts about Salmon—The Strange Use - of the Kidney in the Stickle-back—The Stickle-back and - Parental Duties—Siamese Fighting-fish 175 - - - CHAPTER X - - SOME OF THE “LOWER ORDERS” - - Butterflies and Moths, and the Coloration of their - Wings—Female Choice and “Fine Feathers”—When Male - Butterflies are Dominant—Sexual Selection among - Butterflies—Abortive Experiments—Wallace and the Sexual - Selection Theory—The Sense of Smell in Butterflies and - Moths—Fragrant Butterflies—Wingless Moths and their - Lures to Lovers—Methods of Pairing among Butterflies and - Moths—More Experiments 185 - - - CHAPTER XI - - BEETLES THAT “BLUFF” - - The Coloration, and other Forms of Ornament in Beetles, - and the Significance thereof in regard to the Sexual - Selection Theory—The Courtship of Grasshoppers and their - Kin—The Remarkable Ears of Locusts and Grasshoppers—The - Field-cricket and the Katydid as Troubadours—The - Wonderful Performances of the Cicadas—The Duels of - Long-horned Locusts—Dragon flies—The May-flies’ “Dance of - Death”—The Jaws of the Giant Alder-fly and their Strange - Use—Some Curious Facts about Stone-flies 208 - - - CHAPTER XII - - SCORPIONS, SPIDERS AND CRABS - - Musical Lovers among Spiders and Scorpions—Colour - among Spiders, and its uses—The Spiders’ Dance of - Death—Spiders and Conjugal Bliss—How Pairing is - accomplished—Scorpions in Love—Musical Crabs—Quarrelsome - Fiddler-crabs—Crabs and Courtship in the Deep Sea-Amazons - among Prawns—Brine-shrimps and Water-fleas—“Natural” v. - “Sexual” Selection 236 - - - CHAPTER XIII - - SOME STRANGE MARRIAGE-CUSTOMS: AND VIRGIN BIRTHS - - The Courtship of the Cuttle-fish—The Sumptuous Cradle of - the Argonaut—The Love-darts of the Snail—Hermaphrodites - and the Dangers of Self-fertilization—Oysters and - Beauty—Sex reduced to its Lowest Terms—Parthenogenesis - and Virgin Birth—The Story of the Hive-Bee—The Departure - of the Queen—The New Queen and her Marriage-flight—The - Celebration of the Nuptials and its Surprising Sequel—The - Widowed Queen turns Executioner—The Queen as Mother—The - Queen’s Daughters—Nursemaids’ Duties—Change of Work—The - Drones and their Career—Food and Sex—The Bumble-bee and - its Life-story 265 - - - CHAPTER XIV - - PARTHENOGENESIS AND ITS SEQUEL - - Courtship among the Ants—The Great Renunciation—Maternity - carried to Extremes—Where Males are - Superfluous—Degenerate Males—Keeping Death at Bay—Where - Females are Unknown 296 - - - - - LIST OF ILLUSTRATIONS - - Love-making Frontispiece - - Facing page - - The Gorilla preparing for hostilities 42 - - The barometer of maleness—among the Apes 44 - - Weapons of offence 52 - - Manchurian Wapiti “calling” 54 - - Group of Beisa Oryx 60 - - Eland Cows 64 - - American Bison 64 - - Elephants 70 - - Head of male Wart-hog 72 - - Male and female Babirusa 72 - - Somali Zebras 72 - - Giraffe 72 - - Californian Sea-lions, or Eared Seals 82 - - Elephant Seal 88 - - Northern Elephant Seal 88 - - “The Peacock in his pride” 96 - - Peacock Pheasant 96 - - Patterns which puzzled Darwin 98 - - The “Strutting Turkey” 100 - - The display of the Great Bustard 100 - - Some of Fortune’s favourites 104 - - The love-making of the Prairie Hen 110 - - Grades of evolution in the syrinx or organ of voice in - the males of Surface-feeding and Diving-ducks 126 - - Fighting for territory 140 - - The display of the Grasshopper Warbler 142 - - The display of the Sun-bittern 142 - - The Kagu in display 142 - - A male-Savi’s Warbler 152 - - Another aspect of the Kagu’s “display” 154 - - Some strange accompaniments of courtship: - - The White-headed Bell-bird 156 - - The Umbrella-bird 156 - - Skull of the American white-beaked Pelican 156 - - Head of a Puffin, showing the moulting of the beak - sheath 156 - - The Satin Bower-bird and its bower 158 - - The “bower” of the Bower-bird 158 - - The Bearded Lizard 166 - - Bright colours which cannot be attributed to “sexual - selection” 200 - - Stridulating organs, etc. 218 - - Crickets and May-flies 220 - - Male Astia displaying before the less brilliant female 242 - - Male Icius displaying 242 - - Scorpions 252 - - Death of the male Scorpion 254 - - The female Mantis devouring her mate 254 - - The “Fiddler-crab” among mangrove roots 258 - - The “Fiddler-crab” 258 - - Some remarkable devices 262 - - Some remarkable methods of “courtship” 268 - - -THE COURTSHIP OF ANIMALS - - - - -CHAPTER I - -INTRODUCTION - - -The nature of Life and its power of reproduction—The stuff of which -Life is made—The Emotions—The simplest living things—Where is neither -Birth nor Death yet the Population increases—The First Marriage—The -beginning of sex—The two dominating instincts—The conditions of -survival—The Oyster’s narrow world—“Fiddling work”—Amorousness—The -superior Male—Where Death begins—“Germ-plasm” and what it means—Sex and -“Secondary sexual Characters”—Some theories—“Hormones” what are they? - -The nature of life is generally regarded as affording a theme which -possesses no more than an academic interest: but there is one aspect -of this great subject which must attract us all, and that is its power -of reproducing itself. Life begets Life, as Love is said to beget -Love. The nature of this mysterious power we can only dimly realize, -and the forces which underlie its manifestations few even suspect, -save perhaps in a vague way. Yet the tree of Knowledge bears no fruit -more vitally important to our well-being, than that which will make us -“as Gods, knowing good and evil” in all that concerns the processes -of reproduction. But curiously enough, this is a forbidden fruit, and -those who eat thereof are expected to maintain a discreet silence on -the subject. These enlightened ones, however, cannot remain altogether -dumb. But they speak, in the veiled language, of Art and Poetry, -Literature and the Drama. They talk round the subject rather than of -it. Love, Hate, Jealousy, and Envy, are but attributes thereof. We -profess to believe that “Knowledge is Power” and to desire to increase -its force among us by raising the standard of our system of education. -But education which does not, of set purpose, reveal the sources of our -being and of our emotions, good and evil, is no more than a travesty -of education; and they who seek to foist upon the community Knowledge -thus emasculated, are unworthy to wield the power which has been placed -in their hands. If social well-being be the aim of the high-priests -of Education, then something more than copybook maxims like “Be good -and you will be happy” must henceforth be preached. Of what avail is -it to exalt the name of Knowledge, while the straightest road thereto -is barred across and marked “No thoroughfare!” These blind leaders of -the blind seem to imagine that the social well-being they profess to -desire can only be attained by side roads, leading anywhere, save in -the direction of this Pool of Siloam. - -The stuff of which living things are made is called “Protoplasm.” -Text-books of Physiology give its chemical constituents with fearsome -accuracy, and each of these constituents can be isolated in the -laboratory, but “all the king’s horses and all the king’s men” cannot -build these up again into living matter. Its consistent inconsistency -defies us; every statement we make of it has to be qualified by -reservations and saving clauses. Its permanency is attested by the -fact that it has endured through millions of years, yet we are daily -reminded of its evanescent nature. Its power of reproducing itself -according to type, none can doubt, yet no two individuals are exactly -alike. - -The purely physical phenomena of life, to be rightly appreciated, must -always be considered in relation to the psychical phenomena which -are the soul of life. These subtle and intangible forces cannot be -experimented with in the laboratory, or expressed in formulæ; we -cannot denote their strength in horse-power. Just as the physical -manifestations of life begin with lowly types, so the psychical begin, -and they gather strength and complexity with the bodies they pervade. -These manifestations we call behaviour, and in their more intense -developments, “emotions.” - -These emotions present an infinite range of variety in the higher -animals, and they attain their maximum of intensity wherever the -reproductive activities are concerned. The part which these activities -play in controlling behaviour is by no means always apparent, and -is commonly not even suspected. Even man himself is subject to this -control. And it is this fact which lifts the “Courtship” of the lower -animals out of the category of merely curious phenomena. For the -springs of his conduct, his behaviour and “emotions” under varying -circumstances, can only be understood, and even then but imperfectly, -by comparison with other creatures lower in the scale, so far, of -course, as comparison is possible. - -This line of inquiry, then, takes one back to the simplest living -things, among which there is neither marrying nor giving in marriage, -neither birth nor death. Life is reduced to its simplest terms—a speck -of animated jelly is all that confronts one, and this is only to be -seen under a high power of the microscope. It has neither mouth nor -organs of digestion, no visible means of locomotion are traceable, -and the special senses of sight and hearing are wanting; but taste -and smell, of a nebulous kind, are there. Shape it cannot be said to -have, for its bodily outline is constantly changing, thereby it moves. -A long tongue of its jelly-like substance, or “protoplasm” as it is -called, is thrust forwards, and the rest of the body is, as it were, -dragged after it. Whatever animal, or vegetable, matter it passes -over, in the course of its wanderings, is drawn up into the semi-fluid -substance of this diaphanous body, and its juices extracted, the -undigestible residue is left behind in the course of the morning’s -walk! In due time it becomes adult; further growth is impossible. When -this stage is attained a strange thing happens. A certain minute, more -solid portion of this body, which lies in the very centre of the mass -and is known as the “nucleus,” begins to assume an hour-glass shape. -Speedily the constriction becomes apparent across the whole body and -rapidly increasing, cuts it in two, as if by the tightening of some -invisible thread. Here Death is cheated, and records of births are -unknown! And just as there are no parents so there are no children. -But a foreshadowing of what is to be occurs even here. For every now -and then two individuals, to all appearances identical, meet and -promptly begin to merge the one into the other till they twain become -one flesh in very truth. Here is the most primitive form of marriage -in Nature. And here, in this union, or fusion, of separate entities of -Germ-plasm, we have the beginning of sex. Such unions are common among -these primeval forms of life. In many cases this “marriage” takes place -between two particles of Protoplasm of which one is rather larger than -the other. In such case the smaller is regarded as male, the larger as -female. Here we have the first sign of “sexual differentiation” or the -evolution of “male” and “female” individuals. - -Some such union, some such process of “rejuvenation” by the importation -of “fresh blood” seems to be imperative for the continuance of -existence throughout the whole animal world, even though it may take -place at rare intervals of time. Why should this be? Is this strange -meeting and commingling a matter of chance, or is the one seeking the -other possessed by a ravenous mate-hunger? - -As we ascend higher in the scale it becomes apparent that life has -gathered force. That primitive speck of jelly, the Amœba, with which -we started, gave but two signs of animation—the power of movement, -and hunger. Whether these responses to internal stimuli can be called -instinctive is open to argument. But there can be no question about -the instinctive nature of the behaviour of these higher animals. -After the instinct to feed the two most powerful are the desire for -self-preservation—the avoidance of danger—and the desire to mate. These -two are the dominating instincts throughout the rest of the animal -world, not even excepting man himself. - -The tremendous power of “mate-hunger” has been overlooked by a strange -confusion between cause and effect. Almost universally its sequel, the -production of offspring, has been regarded as the dominant instinct in -the higher animals. This view has no foundation in fact. “Desire” for -the sake of the pleasure it affords, and not its consequences, is the -only hold on life which any race possesses. And this is true both in -the case of man himself and of the beasts that perish. Wherever this -instinct becomes weak, or defective, extinction speedily and inevitably -follows. This “Amorousness” is the motive power of “Courtship” wherever -it is met with; manifesting itself in the eccentric, and often -grotesque posturings, or in the loud and often musical cries which -constitute the study of courtship. Intensity of desire is indispensable -to survival. - -Only the lowly and sedentary types, of which the Oyster may be taken -as an example, lack this fire; and here because it is unnecessary. For -the reproductive germs of this animal are discharged into the water, -to take their chance of attaining their object. They are liberated -unconsciously, discharged like the undigested residue of the food, -without effort, and without cognizance of the act. This must be so, -for the Oyster merely lives—vegetates. Sightless, and without power -of movement, after its larval wanderings are over, it lives merely -to eat. And even in this, choice is denied it. The currents of water -mechanically brought to afford the necessary oxygen for the maintenance -of life, bring with them the food which is to restore the slowly -wasting tissues. To such a creature there can be no “outer-world,” no -consciousness of the existence of individuality other than its own. - -The desire for sexual intercourse is met with only where the -co-operation of two individuals is necessary to ensure the production -of offspring. Such individuals being free to roam, must have some -incentive to seek one another at the time when their germ-cells have -attained maturity. And this incentive is furnished by the glands in -which these elements are produced: supplemented by the secretions -of certain ancillary glands. These stimulating juices, known as the -“Hormones,” will be presently described. - -But if we owe our existence to the gratification of what may be called -our lower instincts, it is no less certain that all that is best in us -we owe to our offspring. We meet with the beginnings of altruism, which -the begetting of offspring entails, far down in the animal kingdom, and -it attains to its full perfection in the human race. Here only, in its -best and truest sense, Love begins: though affection may be found, and -in a high degree, in many of the lower animals. - -Living things are as clay in the hands of the potter. But it is as if -they made themselves, for the designer and the guiding hand are alike -invisible. No vessel is exactly like its neighbour, either in the -quality of its substance or in the details of its construction. And -this because the clay of which it is made possesses that mysterious -property we call life. A property which endows each new feature as -it appears, with an individuality of its own, whose survival, or -suppression, depends entirely on its relationship to surrounding parts; -on its harmony with its environment, in short. Colour, size, shape, -temperament, behaviour, may each be regarded as so many entities -depending for survival on whether or not they can exist in harmony with -their environment—the several parts which make up what we call the -individual. - -In like manner the individual—the complex bundle of parts and -qualities—must attain, and maintain, a certain harmony with its -environment—the outer world. The process of change, both in quality -and quantity, which is for ever going on among the several parts of -every separate individual, brings about the elimination of unfavourable -variations; and “selects” those which vary in the right direction: -that is to say, which serve to maintain a place in the sun for the -individual in which these momentous changes are going on. But it is not -enough that the individual should be in “working order”; it must be in -harmony with all the conditions on which existence depends. And the -standard of this harmony is set by that very exacting arbiter of life -and death, “Natural Selection.” It is not enough that the instincts -in regard to this or that habit should be keen, or that this or that -particular organ of the body should be efficient—a certain minimum, -all-round, standard of efficiency is demanded, or elimination follows. -It is through this instability of “temperament,” this tendency to vary -in infinite directions, that the balance between the individual and -the environment is maintained. Evolution follows the line of least -resistance. - -The little boy who remarked that it must be “fiddling work, making -flies,” was more sage than he knew. The complex web of factors which -even a fly represents are beyond the grasp of human understanding. But -it is clear that the reproductive instincts, and the emotions they -beget, have played, and play, a tremendous part in the evolution of the -higher animals. - -Those whose business it is, for one reason or another, to study -these emotions know well that “mate-hunger” may be as ravenous as -food-hunger, and that, exceptions apart, it is immensely more insistent -in the males than in the females. But for this, reproduction in many -species could not take place: for the sexes often live far apart, and -mates are only to be won after desperate conflict with powerful rivals -no less inflamed. Thus it is idle to speak of an equality between the -sexes in this matter, in regard to the human race. Dogmatism, and the -frequent repetition of pretty platitudes, will not alter what Nature -has ordained. The failure to realize this is painfully obvious in -the utterances of many who speak in the name of the newly-founded -“Eugenics” society, which seeks the means to ensure the well-being -of the race by the spread of a more intimate knowledge of this -all-important subject. The existence of what Mr. Heape has recently -called a “sex-antagonism” is beyond dispute, for the instincts of the -male and female are fundamentally different. The male is dominated -by the desire to gratify the sexual appetite; in the female this is -counteracted by the stimulation of other instincts concerned with the -cares of offspring. - -Amorousness, then, is the dominant feature of the males among all -animals: and this sex presents yet another characteristic which is to -be borne in mind. In all that concerns the evolution of ornamental -characters the male leads. In him we can trace the trend which -evolution is taking; the female and young afford us the measure of the -advance along the new line which has been taken. Why this should be is -inexplicable. But sooner or later the females assume, or will assume, -all the features originally possessed by their lords; and finally the -young also follow suit. That is to say, the females and young tend to -retain the ancestral characters. In the course of time the ability to -develop new features by the male loses its impetus, and not till then, -apparently, do the females, and still later, the young, begin to share -his glory. These remarkable features are strikingly illustrated among -the birds, as these pages will show. - -Nature is nothing, if not perverse. And hence it happens that there -are many exceptions to every rule which one formulates. Among the -birds, for example, there are species wherein the rule that the female -follows her mate in the acquisition of new characters is, so to -speak, set aside. She follows a line of her own. This is true, at any -rate, of superficial characters, such as coloration. By some curious -change in her “metabolism,” as the conversion into living tissue of -the substances taken as food is called, this coloration may attain -a brilliance in no way inferior to that of the male, but strikingly -different. The beautiful Orange Fruit-pigeon (_Chrysoenas victor_) -furnishes a case in point, the male being of a gorgeous orange-yellow, -the female of a no less vivid green. But the differences are not so -great as they appear at first sight. For the male was originally green, -and the female has thus but intensified the ancestral livery. Green, -it should be remarked, of a more or less olive shade, always precedes -yellow in development; and yellow may yield to red, but this order is -never reversed. A no less striking case is that of the Upland Goose -(_Cloephaga magellanica_), the male of which is pure white, while the -female wears a livery of chestnut and brown. But so sharply are the -colours defined that it would be difficult to say that one was of a -higher order of coloration than the other. To what causes or factors -are these departures due? - -Reproduction in the simplest living things takes place by a simple -division of the body into two as soon as its maximum size or adult -condition has been attained. In such simple types the body consists -only of a single “blob,” or particle, of jelly. But a new era began -when large numbers of such particles, or “cells,” began to form -coherent masses, different parts of the mass performing different work -for the mutual benefit of the community. Some have come to form what -we call the body, which is born, and in due course dies. Others are -alone concerned with the task of reproduction. They are nourished by -the body, and on attaining maturity, give rise to new bodies. These -reproductive cells are excessively small. The male, or “sperm” cell, -can only be distinguished under the highest powers of the microscope. -The female cell, or “Ovum,” is always larger than the male, because, in -addition to the germinal matter which it contains, it is furnished with -a store of food in the shape of yolk. This accounts for the relatively -enormous size of the egg of the hen. Within the hardened shell the -germ develops into the chick, deriving food for its growth from this -generous store. Where this yolk is limited in quantity the growing -body is hastily fashioned, and launched forth into the world in the -form of a “larva,” when it must forage for itself till it has attained -its adult form. Or it is retained within the body of the mother until -development is complete. - -The reproductive cells are the bearers of the Germ-plasm, the stuff -of which man and the beasts of the field alike are fashioned. Only a -portion of this germ-plasm gives rise to a new body; the rest is, as it -were, held over and stored within the new body to give rise to another -in due course. That which produces the body we call the “Somatoplasm,” -because it is the “plasm” or stuff of which the “Soma,” or body, -is made. As to the nature of this Germ-plasm and its mysterious -properties, a wide divergency of opinion exists among _savants_. But the -views which find most favour to-day are those of the veteran Professor -August Weissmann, as set forth in his work on the “Germ-Plasm, a Theory -of Heredity.” - -The excessively minute quantity of this germ-plasm which suffices -to form a new body is incredible. By what miracle of miracles is -the essence of a man distilled? His body arises from the union or -commingling of two particles of living matter so minute as to be -invisible to the naked eye. One of these particles is the “sperm”—cell -furnished by the male parent; the other, the “ovum,” furnished by -the mother. True the ovum may measure as much as the one-hundred and -fiftieth part of an inch, but the bulk of this is yolk-food necessary -to furnish the tender germ with life and energy till it shall have -attached itself to the walls of the womb, whence all its future -nourishment is derived. - -By no process of analysis known to us could the germ-plasm of man be -distinguished from that of, say, a jelly-fish; and in the matter of -quantity there is no more difference. Yet, identical to our senses, in -potentiality how amazingly different are these two particles of jelly! -In the lowliest animals, such as jelly-fish, one cannot distinguish -male and female at sight. The appearance of separate male and female -individuals begins somewhat high in the scale marking an epoch in -the history of animal life. For the birth of sex inaugurated not -merely individuals producing distinctive “male” and “female” germs, -but individuals which, by virtue of their sex, developed differences -of behaviour and mentality which were to be followed by tremendous -consequences. Certain aspects of this behaviour are to furnish the -theme of these pages; others, and no less important, those who will may -discover in Professor Arthur Thomson’s “Evolution of Sex.” - -We are far, indeed, from being able to explain the attributes of sex. -At most, we can but endeavour to interpret the behaviour associated -therewith. This was the task which Darwin set himself to achieve in -his theory of sexual selection. He was influenced in the train of -thought which he followed up with such brilliant success by what he had -observed in the behaviour of highly-ornamented species, such as the -Peacock and the Birds of Paradise. The strange antics of these birds -when under the influence of sexual excitement persuaded him that they -were at least dimly conscious of their splendour, and of its power -to fascinate. The female, on the other hand, was supposed to be coy, -and to bestow her person on the finest performer. In this way the -dullest birds and the poorest performers were gradually eliminated. -Here, indeed, was sexual selection. The frills thus begotten he called -“Secondary Sexual Characters,” a term which is also used, and was used, -by him, to include any feature whereby the sexes can be distinguished -apart from the character of the genital organs. - -Horns, tusks, and spurs are other forms of secondary sexual characters. -And these stand for another form of sexual selection—that of selection -by battle. Herein victory falls to the strongest and most pugnacious -male who, as the spoils of victory, annexes the females which formed -the subject of the duel. This theory, which must be discussed at -greater length in the course of these pages, has had many critics, -and among them men of mark. But whatever modifications may be deemed -necessary, they will be such as are demanded by the results of later -discoveries rather than to the force and subtlety of the arguments of -his opponents. - -One of the most formidable of the opponents of the Sexual Selection -theory was Wallace. But his arguments were far from convincing, and -often inconsistent. He attributed the more frequent occurrence in male -animals of brilliant coloration and exaggerations of growth such as -give rise to manes, beards, long plumes, and so on, to a “surplus of -strength, vitality and growth-power which is able to expend itself -in this way without injury,” or, as he sometimes expresses it, to -superabundant vitality. He was evidently striving to find words for -the faith that was in him, and he was nearer the truth than he knew -or than his critics supposed. He was seeking facts which only the -physiologist could furnish. And these made their appearance long years -after with Professor Starling’s discovery of Hormones. We are far from -understanding the origin of these mysterious juices which must be so -frequently alluded to in these pages, but they are evidently intimately -associated with the expenditure of energy. This may sometimes find an -outlet in increased stature, sometimes in pure luxuries of growth. The -force of Wallace’s arguments was crushed out by the weight of detail -they were made to bear. - -Mr. J. T. Cunningham a few years ago entered the lists and failed to -achieve his purpose no less completely. His was a theory which assumed -too much. In the first place it was based on the transmissibility of -acquired characters, of the truth of which there is at present no -evidence. - -He contends, for example, that the vivid hues of scarlet, blue, yellow -and violet which colour the naked skin of the neck of the cassowaries -and of both sexes, and the curious horny casque which surmounts the -head, are the outcome of the constant laceration of the skin inflicted -by the males during their conflicts for the possession of the females. -He assumes that such conflicts take place, and he assumes that such -“acquired characters” are transmitted. Now, as a matter of fact, these -birds do not fight with their beaks, but with their feet. And to this -end the claw of the inner toe is enlarged to form a great spur. But -there is no evidence that the skin of the neck is ever damaged in such -conflicts as they may engage in. No scars are ever found, at any rate, -to lend support to this theory. The casque, which is similarly supposed -to be a mark of honourable conflict, is an “ornament” of great frailty, -for it is composed of a delicate filigree-work of bone covered with a -thin sheath of horn. In like manner, the long plumes which surmount the -heads of birds like the Peacock, and many Birds of Paradise, and the -wattle which surmounts the beak of the Turkey, are supposed to have -had their origin in similar pugilistic encounters in the past. Mr. -Cunningham is surely pushing the theory of the transmission of acquired -characters a little far. For what has been transmitted in these cases -is not a number of scarred surfaces, but a series of hypertrophied -structures. An amazing array of ornamental characters, symmetrically -disposed, and often vividly coloured, in short, has been produced from -lacerated tissues which in kind and extent can have varied but little. - -Evidence has been accumulating during the last few years which would -have rejoiced the heart of Darwin. Had he known that birds of sober -hues “display” with the same animation and with as much elaboration -of posture as the Peacock and the Pheasant, his theory of “Sexual -Selection “would probably have left little for those who came after him -to criticize. Since his time it has been discovered that both permanent -and recurrent secondary sexual characters, such as the antlers of -deer and the temporary nuptial plumage of birds, such as the Ruff for -example, are controlled as to their growth by the stimulating action -of the “secretions or juices formed by certain of the ductless glands -“; that is to say, of glands having no apparent connection with their -surrounding tissues. We owe much of our knowledge of this subject to -Professor Starling, who has called these secretions “Hormones.” - -Darwin knew that the essential sexual glands, the testes and the -ovaries, in some mysterious way controlled, in a large degree, the -development of these “hall-marks” of sex, for it was known in his -time that castrated stags failed to produce antlers, and that hen -pheasants, for example, in extreme old age, or when the ovaries were -damaged by disease or injury, at once assumed the plumage of the cock; -but the part played by these ductless glands was quite unsuspected. -They are the Thyroid, and the Thymus glands, which are attached to -the outer walls of the trachea or windpipe. The Pituitary body, which -forms part of the brain, and the Suprarenal bodies, attached to the -kidneys. It would be foreign to the purpose of these pages to enter -into the functions of these glands; suffice it to say, that the juices -formed therein are taken up by the blood, and distributed over the -system. Their action is only very imperfectly understood. We know -that any derangement in their efficiency results in disease, and that -they play a very important part in the reproductive system, as will -become abundantly evident in the course of these pages. Much hitherto -attributed to the action of “Sexual Selection” alone, it is now evident -is largely due to their action. - -The all-sufficiency of the “Sexual Selection” theory to account for -the development of armature, such as horns, antlers, and the huge -spine-like outgrowths which form so conspicuous a feature of many -of the extinct Land-dragons, or Dinosaurs, has been by no means -universally accepted. Some authorities like Dr. A. Smith Woodward and -Professor Osborne interpret these after another fashion. They hold -that these are the “expression points” of inherent growth forces, a -process of concentration marking the final stages of evolution prior -to extinction. From which it may be inferred that there is a term to -the life of a species as there is to the life of the individual. In -many cases it is suggested the very exuberance of growth has been the -exterminating factor, as in the case of the huge antlers of the Irish -“elk,” whose enormous weapons hampered his endeavours to escape his -enemies. This is the theory of “Orthogenesis,” or direct development. -According to this, new structures, arising in the germ-plasm as -“variations,” will of their own inherent vitality go on increasing in -each generation unless, and until, checked by “Natural Selection.” -Changes in the character of the “Hormones” might very well bring -about these excesses of growth. It is well known that the exuberance -of growth which produces giants among the human race is due to a -derangement of the secretions or hormones of the pituitary body which -largely control growth. - -Another factor of Sexual Selection which is commonly ignored, but which -is of profound importance, is to be found in the part played by the -emotions in regard to sexual relationships; the part which the “mind” -has played, and plays, in the mating of animals, at any rate of the -higher types. - -Darwin touched but lightly on this theme. Later writers have almost -entirely ignored it. Almost all that is worth knowing on the subject -we owe to Professor Lloyd Morgan, who was one of the first to take up -this difficult line of investigation, and to Professor Groos. Their -researches have shown that there can be no doubt but that the emotions -have played and are playing an important part in the phenomena we are -striving to analyse. Sexual selection, in short, is concerned not -merely with the evolution of the physical characters of the body, but -also, and no less, with the psychological attributes thereof. Many new -and extremely valuable facts in this regard have been brought to light -by Mr. H. Eliot Howard in the course of his remarkable studies on our -native warblers. Not until the psychology of sex in the lower orders -of creation has been further investigated shall we have a properly -balanced account of the part played by sexual selection in the scheme -of evolution. - -By now it will have become apparent that the study of the “Courtship” -of animals is one of alluring interest and full of pitfalls for the -unwary. And this because of the apparent difficulty in drawing any -hard-and-fast line between the part played by “Natural” and the part -played by “Sexual” Selection, at any rate in some cases. - -To this aspect of the theme Professor Lloyd Morgan has drawn particular -attention. “It is difficult,” he remarks, to accept the view that -individual choice has played no part where the sexual instincts are -concerned. But supposing that it has played its part ... the effects -will be wrought into the congenital tissue of the race if, and only -if, there are certain individuals which, through failure to elicit the -pairing response, die unmated. Is preferential mating, supposing it to -occur, carried to such a degree that some individuals fail to secure -a mate? That is the question. If so, sexual selection is a factor in -race progress; if not, though it may occur in nature, it is inoperative -as a means of evolutionary development. The whole question, in itself a -difficult one, is further complicated by the fact that the males which -are possessed of the most exuberant vitality, and are therefore by -hypothesis rendered the most acceptable through emotional suggestion, -are likely to compete with other males of less exuberant vitality by -direct combat. Such competition, by which the weakest are excluded from -mating through no choice on the part of the female, falls under the -head of natural selection, and not of sexual selection, if by that term -we understand preferential mating. - -“This serves to bring out the difference ... between natural selection -through elimination and conscious selection through choice.... -Sexual selection by preferential mating begins by selecting the most -successful in stimulating the pairing instinct.... The process is -determined by conscious choice. It is in and through such choice that -consciousness has been a factor in evolution.” - -Herein Lloyd Morgan, like Darwin, recognizes the existence of a -dual machinery in determining survival, where this depends on the -co-operation of two individuals leading separate existences—Natural, -and Sexual, Selection—sometimes the one and sometimes the other -prevailing. In the former, the females are seized by force; in the -latter, won by displays. - -But is this really so? In these pages it is contended that a sharp -line must be drawn between all those attributes and characters which -are necessary to achieve individual survival, the survival of the -Ego, and all those which, on the other hand, are necessary to achieve -reproduction and the survival of the race. The former are governed, or -determined, by Natural, the latter by Sexual, Selection. - -The sphere of influence of these two factors may be delimited, if -we regard natural selection as the factor accountable only for the -qualities necessary for the survival of the individual—necessary to -ensure success in the struggle for existence. Then it will become -apparent that the qualities and attributes necessary to achieve the -survival of the _race_ are of a different kind, and these are the -factors which are embraced under the term “Sexual Selection.” - -It is a mistake to regard animals in relation to the selection theory -as if they were so many tailors’ “mannikins.” Yet a large number of the -critics of the selection theory seem to fall into this error, ignoring -all but the most superficial characters. - -The peculiarities of colour, structure and behaviour, that is to say, -the characters and qualities which distinguish the individuals of any -given race, are due to inherent qualities of the germ-plasm. Each of -such qualities, therefore, may be regarded as entities. Selection -determines their survival. Intracellular selection is the first sieve -through which they have to pass, natural and sexual selection are -others, as circumstances may determine. - -As a rule the sex of an individual is attested by more or less -conspicuous external features. These are known as the “Secondary Sexual -Characters.” But no hard-and-fast line can be established for these, -at any rate, so far as colour and ornament are concerned, for such, as -will become apparent in the course of these pages, tend to appear first -in the male, and then, later, to be acquired by the female, until in -many cases the two sexes become again indistinguishable. - - - - -CHAPTER II - -“MANKIND IN THE MAKING” - -The use of the term “Courtship”—Primitive Man and the Foundations -of Society—“Amorousness” as a motive force—Polygamy—Our half human -ancestors—Standards of Beauty—Disquieting signs. - -Our ideas on the subject of the “Courtship” of animals are of necessity -largely framed on what has been observed by each of us in regard to our -own race; and without any very careful analysis of motives, or thought -of what lies behind. But no real insight into this most tremendous -subject can be gained which does not strive to penetrate beyond what is -actually seen; which does not endeavour to get at the source of conduct -in this regard. - -“Courtship” is the word we commonly employ to describe the act of -wooing; and in civilized human society at any rate, the intensity of -the emotions which inspire the desire to woo are held in restraint by -a variety of causes—and hence the “Courtship.” In the lower animals it -is a moot point whether the term “Courtship” can be accurately applied. -They are governed by no conventions, for them there is neither modesty -nor immodesty. Desire with them is not made to walk delicately, -veiled according to custom; nor is it artificially fostered as among -civilized communities by stimulating food and the crowding together of -large numbers of both sexes in artificial surroundings. Rather it is -a natural, rhythmical, highly emotional state, which gathering force -inhibits the ordinary emotions, or, rather, overrides them, begetting -an intensity of passion which brooks no control. It demands, without -parleying, or mincing matters, what is really the object of courtship -among the civilized human communities—the consummation of the nuptial -ceremony. The term “Courtship” is a Euphemism. Nevertheless, bearing -this in mind, it may conveniently be used in these pages. - -We cannot hope to understand the springs of courtship in the human race -from what we observe in present-day society, or even from what we have -gleaned thereon from the records of remote ages. We must get back, so -far as is possible, to the very dawn of the human race: to that period -of man’s evolution when his conduct was controlled by purely savage -instincts. But even then the mark of the beast must have been fading -out. His most valuable asset, his larger brain, even then gave him an -advantage over the Apes, his near relations, and over the beasts of -the field which he had begun to bring into subjection. We may assume -that like his anthropoid relations, he was of a solitary, nomadic -disposition, wandering in small parties from place to place as fancy -or food determined. His advance to this stage started when, by the -activity of his enlarging brain, he began to be oppressed by the gloom -of the forest, and drawn by the fascination of more open country, and -the ever-varying scenes which exploration brought him. But this life -begot new needs and new desires. Hitherto, hunger, self-preservation -and self-perpetuation were the only stimulants which roused his -activities; and they were also the three forces, and powerful forces, -which shaped his love of solitude. The proximity of his fellows -threatened his three most vulnerable points—they competed for his food, -they endangered his life, and threatened the possession of his family. - -This more varied and adventurous existence roused new centres of -activity in his brain; he began to perceive, though dimly, the -possibilities of a larger life, though doubtless one which would -minister to his own comfort rather than to that of his family—the -natural and only road to better things. He began to devise more -expeditious means of securing food, and circumventing his enemies, -among whom the most formidable was his fellow-man, because in him he -met his match. In the course of his wanderings he had learned the use -of stones as weapons—which he could never have done in the forests—and -he had also discovered the value of his family as ministers to his -comfort, if only by setting them to collect such food as did not -require strength and cunning in its capture. An inherent love of the -chase for the sake of the excitement which this afforded probably made -him nothing loth to regard hunting as his own peculiar duty. A little -later the advantages of neighbourliness were borne in on him, largely -for the sake of the greater ease wherewith the animals of the chase -could be captured by their combined efforts; but this begat comradeship -and some of the graces which follow therefrom. - -Thus was laid the foundation of Society and “civilization” with all its -attendant barbarities. Then, as now, whatever discordant notes were -heard, were those struck by the twin Demons Envy and Jealousy. These -disturbers of the peace are parasites on Society, their very existence -depends on it. They have played a larger part in fashioning its rules -and regulations than is generally realized. Their influence is as -powerful to-day as ever in the past. It expresses itself in varying -degrees in different individuals, and is roused by varying causes. But -the most potent of all is jealousy in regard to sexual matters. - -Amorousness, a word with a deep meaning, was, and is, the underlying -factor which shaped, and is sustaining, human society; and is no -less powerful among the beasts that perish. The motive force in this -has not been the desire for offspring, but for the satisfaction of -the elemental animal passion, the gratification of the purely sexual -emotions which at their height are irresistible. There may be some -who will see in this contention a degrading aspect of life. But this -view will obtain only among those who prefer the man-made sophistries -of life to its Divine mysteries. This dominance of what are popularly -called the animal passions is the outcome of a perfectly natural -process, whereby those in which these passions were defective died -without offspring, while those who tended to excess were similarly -eliminated. The desire for offspring for its own sake may exist among -our own species to-day but, normally, offspring follows as an effect -not as a cause. Many of our social problems would straighten themselves -out if these facts were once faced and acknowledged; we are apt to -concern ourselves with what should be—according to our ideals—rather -than what is. Let it be granted that this rendering is true, and much -else that mystifies becomes clear. - -Whether primitive man was monogamous or polygamous, or whether he -practised promiscuity, are themes which have exercised the minds of the -most ingenious since the custom of making books began, and the most -diverse conclusions have been arrived at. In coming to any conclusion -on this subject probability based on what we know of the higher apes -can be the only standard of argument. In these animals monogamy is -the rule, the male and female with their young roaming at large in a -family party. Occasionally, however, a male is seen accompanied by two -females, and this is only what we should expect. The Apes are not very -prolific animals nor are they numerous in individuals, hence, should -any male be killed either in combat with a rival or by any other means, -his mate probably wanders in search of another male, by whom, when -found, she is probably readily adopted even if he should be already -mated. - -In like manner lived our half-human progenitors. But with them family -parties no longer wandered aimlessly searching for food, but with -a purpose. No longer forest dwellers, or vegetarians, food would -require more zeal and discrimination in collecting, and shelter of -some kind had probably to be devised, partly as a protection against -predatory animals, and partly for personal comfort, since it would -now have become apparent that this could be appreciably increased by -the exercise of a little effort and ingenuity. This appreciation of -creature comforts formed a cement holding the family together; a sense -of safety in Society helped still further. Rude tools chipped from -flints were among their earliest and most cherished possessions for -the sake of the advantages they secured. Here was the earliest form -of wealth and the birth of labour and a further step on the road to -progress. Little would now occur to derange the harmonious routine -of the daily life, save only the ever-present jealousy of the head of -the family which was assailable both from within and without. His sons -and daughters were probably now regarded as a portion of his wealth, -for they ministered to his comfort, and aided in the daily work which -had now become a necessity. As his sons attained to maturity, so they -became rivals to be watched with a jealous eye, and finally driven -off, while his daughters at the same time became potential mates. -This danger of close inter-marriage was a real one, though it cannot -be supposed that it was in any way realized. The risk was evaded by -perfectly natural means. The jealousy of the head of the family which -drove him to expel his sons as they attained maturity provided the -means. These young bachelors sought their mates from neighbouring -families, and it is probable that they would not be hard to lure from -their parental control, but in such matters force was able to effect -where persuasion failed. - -These mate hunting excursions are to be regarded as extremely powerful -factors in securing the betterment of the race. They were adventures in -which all must fail who did not possess courage, cunning, and brawn, -for, paradoxical as it may seem, evolution depends, not so much on the -qualities of the individual as on the elimination of the unfit. As -yet might was right. But the strife of combat, fierce and merciless, -had its beneficial results not only in weeding out the physically and -mentally deficient, but in stimulating affection between the victor and -his prize. - -As the advantages of neighbourliness dawned upon these children of -nature, rules and regulations, for the control of the individual on -behalf of the good order of the community, came into being; and among -the earliest laws to be framed, we may be certain, were those for -the regulation of marriage. These, as we may gather from the history -of savage races to-day, did not concern themselves with chastity, at -any rate before marriage, it was enough if they secured the right -of possession, and excluded the dangers of close intermarriage. -Promiscuity in the past was never the practice of any race, its -existence to-day, among both savage and civilized people, is due in part -to imperfections in the social scheme, and in part to the vagaries of -individuals. - -That the sexual instincts form the bed-rock on which depends the -survival of all races of animals, which, for their propagation, require -the co-operation of separate sexes, is beyond dispute. And it is no -less certain that in so far as the evolution of man is concerned, -jealousy has been a powerful integrating factor. - -Among the higher animals apart from Man, both polygamy and polyandry -are met with, and this with no apparent detriment to the race. It -is significant, however, that polyandry is never met with among the -mammals, and but rarely among the birds, when, as will be shown, -this form of sexual relationship has been accompanied by a profound -modification of the behaviour of the sexes in regard, not only to -courtship, but to the offspring. The male has lost his masculinity, -and the female her femininity. In human society both forms of marriage -prevail, and there can be no doubt, from the history of such customs, -that of the two types, polygamy is much to be preferred. It is certain -that no race which practices polyandry can do more than hold its -own, and that in a low grade of development. This cannot be said of -polygamy, which might indeed be commended as a solution of some of our -own social problems, were it not almost certain that the remedy would -prove as bad as the disease. - -The subject of “Courtship” in so far as it applies to the human race -is one concerning which little can be said. Westermark, Letourneau, -Sutherland, and last but by no means least, Darwin, have brought -together a mass of facts bearing on the status of women among -communities, savage and civilized, ancient and modern, and from these -much may be inferred. To this harvest, however, Darwin himself still -remains the most important contributor on all that directly concerns -the “Sexual Selection” theory. Other writers seem to have paid more -attention to the laws governing the possession of women than to the -discussion of the motives which may have controlled the choice of -mates. Instances of amatory dalliance, such as are met with among -the inferior apes, and the birds, seem to be wanting. This negative -evidence seems to show that, even among the most ancient, the most -Ape-like, half-human races of man such dalliance was unknown. And this -because primitive man, in his love-making as in everything else, was -accustomed to take what he wanted, or die in the attempt. It is to -this forcefulness of character that the human race owes its progress -throughout the ages. But did he, when desire possessed him, exercise -any sort of choice, when this was possible? What were his standards? -These are unanswerable questions; at most we can but infer what his -behaviour may have been from observations on existing races of mankind. -These seem to demonstrate that while some races profess admiration for -certain of their physical peculiarities, these cannot be attributed to -the action of sexual selection. - -It has been suggested that the low, beetling brows, protruding mouth, -and flat, broad nose which characterized the earliest human peoples, -were slowly eliminated by the æsthetic taste displayed by the females -in their choice of mates. Now in the first place, it is highly -improbable that they had any choice allowed them, and if they had, -these are just the characters which were most marked in the males and -might, or probably would, in consequence, have been deemed “manly” and -desirable, for it is hardly to be supposed that such people would be -capable of conceiving ideas of a possible refinement of their personal -appearance if they could but add to the height of their foreheads -and reduce the size of their faces. These graces settled down on -them as the brain enlarged and habits changed. But the process of -transformation must have been infinitely slow, and quite imperceptible -from one generation to another. - -The absence of secondary sexual characters in man, such as the brightly -coloured areas which are so conspicuous a feature of many of the lower -apes, is to be explained by his fundamentally different mode of life. -Such vivid hues obtain only in species which live in troops, and -they serve as aphrodisiacs, ensuring mating to every female forming -a part thereof, which would be by no means certain were there no -external signs of her condition. Primitive man, like the higher apes, -was instinctively monogamous, and of necessity solitary, till he had -acquired a tolerable measure of self-control and neighbourliness. When -lust possessed him, he was obliged, in making his maiden venture to -scour the country in the search for the object of his desire. This -found, and won, probably only after desperate conflict with the head of -the family, the nuptial ceremonies would be short. - -The greater physical strength of the male and his higher brain capacity -are probably the result of Natural, rather than of Sexual Selection. -The former would weed out all the weakly and dull-witted in the -ordinary course of the struggle for existence, the latter, during the -early days of man’s development, would award the prizes of life to the -most amorous and cunning, and to the most ambitious of the competitors. - -The secondary sexual characters of the female are chiefly negative -characters, the absence of those which are conspicuous in the male. She -retains more of the primitive characters of the race. This is the rule -in regard to the animal kingdom. Wherever we desire to find the onward -tendency of evolution, the latest developments of the race, we turn -to the male; when we desire to learn something of the past history of -the species we turn to the female and young. This standard, of course, -yields by no means uniform results, for we find every gradation of -progress on the part of the latter, till male and female and young are -externally indistinguishable. But the order is almost invariably the -same—first the male, then the female, then the young. Thus progress is -more or less automatic or “Orthogenic,” as the scientific text books -have it, new characters, as they appear, tending to go on increasing -in amplitude till checked by Natural selection. It is to be noted, -however, that this transference is limited, for the female never -inherits characters which are concerned with aggressiveness to the same -degree as in the males, as witness, for example, the brow-ridges and -huge canines in the case of the gorilla. - -Darwin believed that the beards of men have developed by the selective -choice of the women who preferred bearded men, while the secondary -sexual characters of the women indicate the lines of male choice. -There is, however, no evidence to show that in the past—for these -characters are as old as man himself—woman had any choice whatever in -the choice of her mate, save under exceptional circumstances. He was -led to this conclusion by one or two striking instances apparently -demonstrating this choice, and on these he seems to have based his -version of the influence of sexual selection in man. The first of -them is furnished by the Hottentots wherein, in both sexes, there is -a marked “Steatopygy,” or accumulation of fat on the buttocks. In the -female this is excessively developed, and it is said that such females -are highly prized by the males. Darwin cites an instance of a woman -in which this accumulation was so enormous, that she could only rise -with the greatest difficulty from a sitting position. But there is no -evidence to show that less favoured females remained unmarried. - -In other tribes the breasts attain excessive proportions, so much so -that they can be slung over the shoulder to feed the infant strapped -to her back. These may have been increased by sexual selection, the -preference of the males for such mates as possessed this feature in -the most marked degree; but there is good reason to believe that such -characters, which, it must be remembered, are the outward manifestation -of germinal variations, once having appeared, would of themselves, of -their own inherent vitality, have gone on developing. They won favour -from long familiarity, which has imparted a semblance of increment -from choice. These increments of growth in any given generation would -be imperceptible, but variations in excess of the average would be -conspicuous, and excite admiration from their very strangeness. - -The part which sexual selection has played in determining the physical -characters of the human race has without doubt been overestimated. -Its influence may be said to have ceased with the development of the -emotional side of his nature. This momentous process began with the -male and had its roots in the ebullitions of his inherently amorous -nature which has been the dominating factor in his career, and will be -to the end, however much its influence may be disguised by the complex -conditions of civilization. - -These emotions, varying in kind and intensity, are such as are embraced -in the term “Love” in the highest sense. They control the selection -of mates, but this selection takes no account, save by accident, of -qualities which have any value as factors of race-survival. In the -lower animals these are determined by natural selection, and sexual -selection adopts as it were the material furnished thereby. It -“selects” only in so far as it eliminates the non-sexually inclined, -and those which lack the qualities essential to ensure reproduction, -such as weapons for example. In human communities natural selection -is largely avoided, and “mate-hunger” seems now to be swayed by more -than the mere desire for its satisfaction. With the development of -human faculty new factors have been introduced, complex emotions have -come into being, whose influences are as yet only vaguely understood. -Whither are they tending? What will be their effect on race-progress? -These are matters of grave importance to us all, and to the student of -Eugenics in particular. - -Of man’s higher emotions, which, it is contended, now govern his -conduct, probably the earliest to assert itself was the æsthetic. His -quickening mentality could not fail to be captivated by the bright -hues of birds and butterflies, and flowers, the glorious colour-effects -of dawn and sunset, the seasons in their changes and so forth. And as -this sense of the beautiful slowly gathered force he would seek to -decorate his naked body with such of the more brightly-coloured objects -around him as were suitable or rather with such as could be affixed -thereto. - -As a signal mark of his favour and affection, he would occasionally -transfer some one, or another, of his most lasting ornaments to -his mate, and the additional charm this would give her ensured a -continuance of such gifts, and paved the way for tribal fashions. But -then, as now among savages, the males take the lead in this matter -of ornamentation, but in proportion as affection grows, they are -transferred from him to her, so that among civilized races to-day, -the custom is entirely reversed, the women, not the men, wearing the -finery. So soon as families began to be neighbourly and to combine -for the sake of company and mutual help, the spirit of rivalry, so -essential to progress everywhere, would tend to increase the number -of such gifts, and to set “fashions.” With the foundation of society -“selection”—by the elimination of the unsocial, would ensure, not -only the survival of such fashions, but their multiplication and -diversification, producing results which, to our eyes, have often been -hideous. The immediate effect of this form of selection, however, was -not a change in physical characteristics, but in the evolution of -personal ornaments and development of the æsthetic sense. Progress -in this direction must have been infinitely slow, and the lower races -of to-day furnish us with instructive object-lessons in its course. -In many cases uglification rather than refinement has attended their -efforts. - -It is indeed more than probable that the various types of ornamentation -obtaining among savage races had their origin in outbursts of sexual -exaltation. One of the earliest methods of personal decoration was -probably to daub the body with paint, as is the custom during the -performance of various religious and semi-religious rites among the -Australian aborigines. A desire to find a permanent substitute for -paint led to the practice of cicatrization, and the later and more -refined custom of tattooing. But personal mutilation has taken many and -strange forms, such as knocking out the front teeth, filing them to -saw-like points, inserting gold or jewels, or staining them. No less -extraordinary are the various types of lip and ear ornaments, and the -suspension of ornaments from the nose. The various fashions of dressing -the hair are also traceable to this origin. - -That these modes of personal decoration designed for special occasions -should in course of time become permanent, and should, in many cases, -have lost their original associations is but natural. To-day among -savage and barbaric races many of these modes of transfiguration have -become associated with religious and semi-religious ceremonies, but -many have been retained solely to enhance the personal appearance, even -though in our eyes an exactly opposite effect has been attained. Among -the natives of the Congo, for instance, the face is covered with raised -patterns formed by cicatrization; that is to say, by cuts made with -a knife, which are made to form scars on healing by means of pungent -juices or heated iron. Further, the teeth are filed to form saw-like -cutting edges, producing a revolting effect according to European -ideals, but charming according to the standards of those thus patterns -which adorn the tattooed face of the Maori present a result more nearly -pleasing. Many of the natives of East Africa pierce the lobes of the -ear and hang ornaments therein so heavy, that in due course a hole -large enough to run the arm through results. These are mutilations of a -purely ornamental character. Curiously enough, precisely similar forms -of mutilation occur among people dwelling in different continents, as -in the case of the lip and ear ornaments worn by natives of Africa and -South America. There can have been no means of communication between -these races, and hence we must conclude they were independently derived. - -More striking still is the practice of deforming the head which -prevailed among the Peruvians, the Caribs of the West Indies, and the -natives of Vancouver, and the Chinook Indians, wherein it attained -its maximum. Among some tribes, the head was depressed from above -downwards, giving the skull a cone-shape, the apex pointing backwards; -among others the pressure was applied to the back and front of the -head, giving a more or less globular shape, and causing the sides of -the head to bulge ominously. Now these distortions are to be attributed -solely to the whim of Fashion. But how could this have arisen? No adult -could have started it, for the form of the skull cannot be altered -once its growth is completed. The conception of this diabolical custom -apparently then arose in the brain of some fiendishly ingenious person, -who realized that to effect its realization pressure must be applied -to the head of the infant at its birth and for some considerable -time after, by squeezing the head between boards, or tying it round -with thongs of hide. That disastrous results would follow from this -tampering with the brain would seem an unavoidable conclusion; yet -such was not the case. During the moulding process, travellers who -have witnessed it tell us, children display no sign of suffering, even -though their eyes seemed to be starting from their sockets from the -pressure. But they cried when the thongs were loosened. On attaining -to man’s estate, such victims to parental folly seemed to be in every -way as intelligent as the men of neighbouring tribes which had no such -insane customs. - -How deeply rooted was the prejudice in favour of this extraordinary -fashion is shown by the fact that when, during infancy, from sickness, -or other cause, the bandaging was neglected or omitted, and the child, -in consequence, attained to man’s estate with a head of the shape -designed by nature, he was seriously hampered in the struggle for -existence, for no honours among his tribe were possible. Indeed, as -often as not he was sold as a slave. But thus did Public Opinion bring -disaster on its advocates, for those misguided people have been swept -off the face of the earth by their own folly. Those who survived the -ordeal, it is true, seemed in no way mentally deficient, but the infant -mortality must have been great, and none of the adults could ever have -attained to their full potentiality. - -These people were, however, not the only lunatics at large. For this -extraordinary practice found its devotees in many other widely sundered -parts of the world. Deformed heads of various types have been found -in rock-tombs near Tiflis, in the Crimea, Hungary, Silesia, in South -Germany, Switzerland, and even in France, Belgium and England! How did -it spread from one nation to another? Since means of communication -were extremely limited centuries ago, one can only suppose that in -most cases it arose independently. It is possible that the idea -started with the unintentional deformations of the head which follow -the practice of carrying the child during early infancy. It is well -known that if a child be constantly carried on one arm, so that one -side of the head continuously presses against the shoulder, a more or -less marked asymmetry of the skull results. It would be enough for -the head of one of the chief’s children to show a rather unusually -marked asymmetry of this kind for every mother to endeavour to copy the -defect, for imitation ever was the sincerest form of flattery! - -To place these superficial, non-transmissible, artificially created -features, such as deformed heads, mutilated teeth and ears, and so -on, in the same category as the “secondary sexual characters” of the -lower animals which are physical, inherent and transmissible features, -is to ensure confusion of thought. The one represents a physical, the -other an emotional development. The persistence of certain forms of -mutilation esteemed beautiful in human society is not to be attributed -to Sexual selection, or to “preferential mating,” for these things -are not only non-transmissible features, but outside the sway of the -amorous instincts, as is shown by the case of those individuals who, -living in a community where deformed heads are _de rigueur_, have heads -of normal shape. So soon as such perversions become a part and parcel -of everyday life, they become essential to the general well-being and -comfort of their possessors, enabling them to follow their normal -avocations without exciting the dislike or wounding the prejudices of -their neighbours. The absence of the “tribal sign” alienates the esteem -and comradeship of his neighbours and brings an unenviable notoriety. -In like manner albinos among birds, for example, are hunted down by -their fellows and killed, and birds of exotic species conspicuous by -reason of their unfamiliar appearance are treated in the same manner. -The sexual instincts have no part in this. - -It will have become obvious in the course of this chapter that Sexual -selection as a factor in shaping the evolution of the human race has -not played a very conspicuous part. Nevertheless, the balance of -opinion to-day is probably in favour of the view that the physical -peculiarities by which we distinguish one race from another are, for -the most part, due to the influence of this form of selection. A more -careful survey of the facts will show that this view is untenable. -And there is no more striking demonstration thereof than that it has -been inconsequently applied to account for features in one race, which -in another are attributed to environment or to Natural Selection. -It may safely be asserted that colour, the shape of the nose, the -prominence of the jaws, and the character of the hair, are no more the -result of “Sexual Selection” than stature, for example. These are the -manifestations of inherent growth forces, or “tendencies,” which owe -their survival, and development, to the influence of Natural selection. - -Sexual selection has brought about the dominance of the male, by -the struggle between males for mastery, originally for females. It -“selected” for survival, in primitive races, those males with the -thickest skulls and the strongest physique; it determined the survival -of the keenest witted and most aggressive and most amorous males, and -it eliminated those in which the latter features were too active. It -assured victory, in short, to those only who possessed just those -qualities on which life or death depend in moments of conflict. In the -case of the females, it assured survival only to those who possessed -strongly developed maternal instincts and submissiveness. - -It is by no means realized that the incidence of moulding forces -has changed and is changing with the environment of the race. So -long as physical force, as between man and man, determined survival, -as among savage races to-day, so long does it ensure to such races -strong men and strong children, for in conflict with neighbouring -tribes victory rests with the most powerful of physique and endurance -and the most prolific. This last is an all-important concomitant if -repeated conflicts are to be successfully waged. Among civilized -peoples such contests began to lose their value in this regard when, -by the introduction of arms, physical personality became a steadily -diminishing factor. Victory now rests rather with those peoples who are -most skilful in devising engines of destruction. The brain, not brawn, -tells. But man cannot live by brains alone. With the inevitable decline -in his physical nature man’s hold on existence is seriously imperilled. -Civilization is making for extinction as much as over-specialization in -the case of the lower animals. Hitherto, save in the case of decaying -nations, women have played but a minor part in what we may call the -“tribal” affairs of the race. Among the civilized nations of to-day, -in proportion as the “maleness” of the community becomes more and more -effete, the victims of sophistry, and the slaves of shibboleths, so -the influence of the females asserts itself. And recent events among -us show plainly enough that that influence is the reverse of good. -Having its roots in personal vanity, and the love of notoriety, it is -intolerant alike of reason and self-restraint, and that way madness -lies. - - - - -CHAPTER III - -MAN’S COUSINS THE APES - -The Man-like Apes and their mode of Life—Their “Courtships”—Musical -Chimpanzees—How the Orang-utan improves his voice—His likeness to -Caliban—The truculent visage of the Gorilla—“Ornament” in the lower -Apes—The Concerts of the Howler Monkeys. - -We are none of us given to boasting of our poor relations, and most -of us indignantly repudiate our kinship with the Apes. But facts are -stubborn things: the relationship is there, whether we admit it or -not: and those who love truth for truth’s sake will not shirk the -comparison between themselves and their remote cousins. Unhappily, -from our present point of view, this cannot be carried very far, for -the “Love idylls” of the Apes have yet to be written. Such facts, -however, as have been gleaned are interesting. Of the higher, man-like, -or “Anthropoid” species only the most meagre information is to be -obtained; but this nevertheless is interesting. For the most part we -have to be satisfied with inferences drawn from a study of the external -differences between the sexes—from the “Secondary Sexual Characters,” -in short, and from the records of travellers who have encountered these -creatures in their native wilds. - -The species which throw most light on this theme are the Gorilla, -the Chimpanzee and the Orang-utan. Of these the Chimpanzee has most -in common with the human race. But it may satisfy the qualms of many -to know that between the Ape and the Man there is a great gulf fixed. -The brain of the largest Ape is less than half the size of that even -of the lowest of mankind. Man is a reasoning, and for the most part a -reasonable, creature; he is a tool-making animal. This is more than -can be said of any of the apes, even the most intelligent. Their teeth -and immensely powerful arms must serve their every need. No ape ever -fashioned for himself either a knife, a vessel to carry water, or any -means of transport; and herein we have a measure of his brain capacity. -The huge jaws and great canine teeth are no less conspicuous “marks of -the beast.” - -These, however, man himself has but recently lost, as was proved by -the sensational discovery of the skull of an ape-like man at Piltdown, -in Sussex, during 1912. Herein the jaw was essentially that of an ape, -while the base of the skull was as markedly human. The cheek teeth, or -molars, were of the human type; but the canine was ape-like, though -much inferior in point of size. That the men of this remote age—which -was possibly that of Pliocene times and certainly not later than early -Pleistocene—had begun to use rudely-fashioned tools, is proved by the -roughly-chipped flints found with the remains. With the invention of -tools the decline in the size of his “eye” teeth began. - -In all the large apes these “eye” teeth are of great size. Their -purpose, it would seem, is primarily to serve as weapons in conflicts -between rivals. Such conflicts are apparently unintentionally, and -unavoidably, provoked by the loud cries uttered by the males in their -endeavours to discover the whereabouts of females desiring mates. Of -necessity roaming far in search of food, the unmated have no means of -making their whereabouts known, save by thus giving tongue to desire. -Evidently the normal methods of voice production do not suffice for -their urgent needs, for the carrying power of the voice is immensely -fortified by means of great air sacs, or chambers, formed in part by an -enlargement of the body of the hyoid, or the bone which supports the -tongue, and in part by dilatations of the inner walls of the larynx. -The females, it is to be noted, are by no means so well equipped -in this matter. It is not necessary that they should be. All that -those desiring mates have to do is to follow up the cries of avid -males, a by no means difficult task, especially when under the spell -of the emotions which possess them. But the mechanism which serves -the Chimpanzee and the Gorilla by no means fulfils the needs of the -Orang-utan. In this uncouth creature the system of resonating chambers -is immensely increased by great, thin-walled, membranous pouches -extending round the neck and under the armpits, so that when inflated -these areas have a most extraordinarily swollen appearance. When the -Orang chooses to lift his voice even the deaf must hear. - -Where fighting instead of fondling is the sequel to these impassioned -cries the conflict is probably not of long duration, for it is -certainly severe. This is attested by the fact that captured specimens, -if adult, are commonly found to be minus one or more fingers, which -have been bitten or torn off in these love affairs. - -[Illustration: Plate 2. - -_From a drawing by I. Thornton._ - -THE GORILLA PREPARING FOR HOSTILITIES. - -Note the “beetling” brows, the large size of the canine teeth, and the -great development of the arms in these arboreal creatures, which play -an even more important part in locomotion than the legs. The latter in -this illustration are, however, relatively too small. - -[Face page 42.] - -An added ferocity of expression is given to the male Gorilla by the -development of enormous brow ridges and the huge canines. The former -are regarded by some authorities as adaptations to afford increased -powers of mastication. But if this were so, then such ridges should -be equally developed in both sexes, and this is far from being true. -Hideousness, rather than ferocity, has been given to the Orang-utan by -the out-growth of enormous ridges on each side of the face, and these, -when the great wind-bags encircling the neck are inflated, impart a -repulsiveness of expression attained by no other animal living. - -Of the normal every-day life of the great Apes but little is known. It -would seem, however, that they live in family parties—an adult male -accompanied by a female and one or more young of different ages, of -which one is commonly an infant in arms. It is difficult to procure -positive evidence on the point, but it is commonly believed that the -young remain with their parents till they are several years old, when -they are gradually driven off to fend for themselves. This is a common -procedure with all animals. The dominant impulse in this is something -akin to greediness, an indefinable perception that too large a family -party will entail too great a strain on the food supply, hence the now -no longer helpless young are regarded as a danger to the safety of -the family, and are turned adrift. Incidentally this procedure is of -immense benefit to the race, for it ensures its distribution, enlarges -its chances of survival, and lessens the danger of in-breeding. - -Attention must now be turned to the lower Apes. In these it is to be -remarked the secondary sexual characters differ conspicuously from -those of the man-like species. Manes and beards and brightly-coloured -areas of bare skin are now the dominant feature. But canine teeth, in -proportion rivalling those of the Gorilla, are found in the Baboons, -while in some of the New-world monkeys voice production of quite -remarkable power takes the place of ornament. - -The precise part played by ornament among these animals can only be -inferred from Darwin’s observations on captive animals, and then only -in so far as they refer to colour. Manes, beards and moustaches, such -as are shown in the adjoining illustrations, are borne only by the -males, and sometimes take extravagant forms. - -Darwin suggested that the mane of the Baboons, for example, served as -a shield when fighting with rivals, protecting the great blood-vessels -from injury. Incidentally this end may be attained, but from what we -know of similar developments in other animals, this cannot be regarded -as the primary function of the mane. One is tempted to look upon it -as a protective device because of its position, but it is probably -no more so than is the long flowing hair which adorns the flanks of -the Guereza. This is of a purely ornamental character, although, -according to some, it is to be reckoned as an instance of protective -coloration, the long white hair matching the long pendant masses of -lichen which hang from the boughs of the trees in the damp forests -where these creatures live, and so concealing them from their enemies. -Of beards and moustaches many examples might be cited, but the most -striking must suffice. These are furnished by the Satan Monkey or -Black Saki (_Pithecia satanas_), and the little Tamarin Monkey (_Midas -imperator_)—one of the Marmosets. In the first-named the beard is thick -and full, but in the latter scanty. This, however, is atoned for by the -enormous upwardly curled moustache giving the face a most comically -human appearance. - -[Illustration: Plate 3. - -_From drawings by I. Thornton._ - -THE BAROMETER OF MALENESS—AMONG THE APES. - -All the Man-like Apes possess great canine teeth and powerful voices. -In the Orangutan the Compass of the voice is enormously heightened by -means of a huge wind-bag which encircles the neck. The wind-bag is seen -in fig. 1, which also shows the great folds of skin developed by adult -males on each side of the face. In other species, as in the Tamarin -Marmoset (_Midas imperator_) (fig. 2), and the Satan monkey (_Pithecia -satanus_) (fig. 3), “ornaments” in the shape of beards and moustaches -are developed, while in the Mandrill (fig. 4) the face is vividly -coloured. - -[Face page 44.] - -In the development of brilliantly-coloured areas of bare skin -the monkeys stand alone among the Mammalia. The hues displayed are -remarkable for their brilliancy, and this varies in intensity, waxing -and waning with the varying moods of their possessors, and attaining -their maximum during periods of sexual excitement. Blue, green, red, -and violet are the dominant colours, and these are confined to the -face, buttocks, and genital organs. The same hues are commonly present -in both sexes, though in the female they are less brilliant. Normally -the male appears to be unconscious of the conspicuous patches of -colour, but when under the irrepressible stimulus of sexual excitement -he seems to endeavour to make the utmost possible capital out of such -adornments, more especially presenting his buttocks to his mate in an -apparent endeavour to stimulate her desire. In some species, as with -the Baboons for example, the naked area of this hinder part of the body -is a much more conspicuous feature in the female than in the male, -becoming enormously swollen and carunculated, and from its vivid red -colour presents a positively revolting appearance, according to our -standard of what is beautiful. The most vividly coloured species of -all is the Mandrill, which, in this matter exceeds all other living -Mammals. The face, in the male, is produced forward to give the head a -dog-like shape, while the whole of the upper surface of the muzzle has -been transformed into a swollen, deeply fluted mass by the excessive -inflation of the underlying bone. The bare skin covering this is of a -brilliant cobalt blue, with lines of violet in the furrows, while the -nose is of a bright scarlet. The naked skin of the buttocks, and the -genital organs, are suffused with brilliant tints of scarlet and blue. -In spite of the purity and brilliance of the coloration the effect is -to make the creature really hideous. - -Of the display Cuvier writes: “La partie postérieure du corps n’est ni -moins extraordinaire ni moins révoltante. Sous une courte queue sans -cesse relevée est un anus entouré d’un gros bourrelet d’écarlate; de -larges fesses nues, que l’animal semble montrer sans cesse avec autant -de lascivité que d’impudence, sont colorées d’un rose vif nuancé sur -les côtés de lilas et de bleu. Les parties genitales enfin sont d’un -rouge de feu d’autant plus tranché qu’elles sont absolument nues, et -qu’elles viennent a la suite d’un abdomen revêtu de poils blancs.” - -While we cannot suppose these animals to possess any standard of -beauty or ugliness, it must not be forgotten that they are more or -less conscious, not only of the existence of these brightly-coloured -areas, but of the effect they produce, as Darwin showed long since in -the cases of a captive Mandrill, and some other smaller species of -Monkeys, among them a Rhesus Monkey. These, when shown a looking-glass, -at once presented their hinder ends to what they supposed to be the -new arrival. A similar mark of friendliness was shown towards their -keeper, and visitors introduced by him. Periodically, under the sexual -stimulus, this desire becomes intensified and becomes an invitation to -mating. - -In this connection it is interesting to note that in some of the -Macaque Monkeys we have signs of a reversal of the usual sequence of -coloration. For in the Pigtailed Macaque the young of both sexes are -more brilliantly coloured than the adults, in regard to the bare skin -areas, while in the Hairy-eared Macaque (_M. lasiotis_) and the Rhesus -Monkey (_M. rhesus_) the face of the female is brighter than that of -the male. This surely means that this coloration is in process of -suppression, for according to the rule the male is the first to develop -new characters, then the female, and finally they are transmitted to -the young. The extra brightness in the young, then, is to be regarded -not as an incipient, but as an ancestral character in process of -elimination. - -As a rule, among the Mammals at any rate, brilliant coloration and -weapons of offence are not associated in the same animal. The Baboons, -and the Mandrill in particular, are exceptions, for these animals are -provided with most formidable “tusks,” the canines of both upper and -lower jaws being of great size, and opposed one to another in such a -way that they wear away to form sharp, angular cutting-edges, more -murderous than the fangs of the Tiger. - -Reference has been made already to the existence of large sound -resonators for the purpose of increasing the volume of the voice in -the Orang, Gorilla and Chimpanzee. Some of the Gibbons are also well -provided in this direction. But the most striking instances of the -kind are furnished by the Orang, and the monkeys known as Howlers. In -these last the base of the hyoid, as the skeleton for the support of -the tongue is called, is fashioned into a deep bony cup, which has the -effect of intensifying the volume of the voice to a most surprising -extent. But more than this, apparently for the protection of this bony -voice-bowl the upright branches of the lower jaw have become remarkably -deepened, and widened, a correlation of growth between unrelated parts -which is fraught with deep significance. “Terrific,” “terrible” and -“harrowing” are terms which have been used by travellers like Bates, -Belt and Wallace in describing the cavernous roar of these animals, a -roar which will easily carry two miles. It would seem that these vocal -efforts are not merely confined to what we may call the “Courting” -season, as is the roar of the stag, but that they are heard nightly -at dusk. They may be resumed again at dawn, and re-awakened when -thunder-clouds gather. They have become the normal method of giving -vent to excitement, and probably are intensified when isolated males -are desirous of discovering the whereabouts of females equally anxious -to find a mate. - -Among the Apes we meet, as with the human species, with both monogamy -and polygamy. But it would be dangerous to assume that the reasons -for polygamy are the same in both. Polygamy, indeed, has by no means -always the same significance. In the most primitive, half-human races -of the past, as with the man-like Apes to-day, polygamy is determined -by accident rather than choice. These extinct peoples, like the great -anthropoids, were normally monogamous, but on the death of a male in -conflict with his neighbour, or from other causes, his mate would -probably of her own free will seek out the nearest male and even if he -were already mated would be at once adopted into the family circle. -This certainly happens in the case of the Gorilla and Chimpanzee -to-day. But among living races of mankind, both savage and civilized, -multiplicity of wives is a matter of choice on the part of the male, -and in many cases to achieve this females from other tribes have to -be secured—either by purchase or conquest. With the lower apes, or -“monkeys,” polygamy only obtains among gregarious species; and either -because the birth-rate of the females exceeds that of the males, or -because a considerable number of young males are killed annually -by exciting the jealousy of the older males, who are exceedingly -pugnacious. - - - - -CHAPTER IV - -AT DAGGERS DRAWN - - -The Birth of Weapons—All Flesh is Grass—Utility and Ornament—The -Fever of Love—The “Challenge” of the Deer—What it means—More about -“Hormones”—“Hummel” Stags—The Age of Deer—The “Courtship” of the -Moose—Types of Antlers—Antlered Females—Fighting Topi—The Lance of -the Oryx in the Lion’s Flanks—Happiness and Hartebeestes—Odoriferous -Suitors—The Bloody Sweat of the Hippopotamus—The Elephant in -Love—Concerning Tusks—Polygamy. - -From Apes to Antelopes is a far cry, but contrasts are always helpful. -Antelopes and Deer, Zebras and Elephants, Rhinoceroses and Swine, are -types, taken at random, of that great and important group of animals -known as the “Ungulates,” or “Hoofed” animals. These illustrate in -a very striking manner what is meant by the term “Secondary Sexual -Characters.” They demonstrate no less forcibly what is meant by the -term “Sexual Selection.” They are valuable in this connection, because -of the often formidable weapons, in the shape of horns and tusks, which -so many species have developed during the struggle for mates. - -But “Sexual Selection” will not explain their origin, and it is -difficult, in the present state of our knowledge, to discover any -clues which will reveal this. In seeking these there are certain broad -aspects of the problem which are not to be lost sight of. In the first -place, horns, at any rate, are confined to the hoofed animals. That the -various types of hoofed animals, living and extinct, have had a common -ancestry, no one at the present day will probably call in question. The -relationship, however, of the various living types, one to another, is -by no means always apparent: the missing links are to be sought in the -records of the rocks. - -When the whole of the evidence comes to be surveyed, and not till -then, it becomes apparent that this wonderful diversity is the result -of complex factors. That the conditions of existence have controlled -the results is beyond question; but it is equally certain that these -conditions have been merely controlling and not causative. In other -words, we must regard each of these different groups or types—Deer, -Antelopes, Horses, Elephants, Swine, and so on—as witnesses of what -we call “Heredity.” They are so many “Diathetic types.” That is to -say, the forms, or individuals, belonging to each type have inherited -certain peculiarities in common; they display a “Diathesis” as the -doctors call it: an inherent, inborn tendency, or habit of growth, in -a definite direction: a tendency which, ever and anon, develops new -qualities, takes new directions. And thus it is that we get Oxen—using -this term in its widest sense and not in its special sense—Antelopes, -Goats and Sheep, for example. These have, among other things, a -“diathesis” in the direction of horn production, and each, too, of -a different type. What is meant by this apparently mystifying term -“diathesis” will perhaps be made clear by taking the case of the Ox and -the Sheep. While very different in appearance, these live on precisely -similar food; yet no one has any difficulty in discriminating between -the taste of beef and mutton. In the marvellous chemical laboratory of -the body the grass gathered in the same field is converted into flesh -which even in its uncooked state is easily distinguishable. Though for -the purposes of this illustration domesticated animals have been used, -the same is true of their wild relations. Sportsmen tell us that the -various types of Antelopes and the Zebra, which may be seen feeding -together, have yet flesh of very different qualities. These qualities -are to be attributed neither to “Natural” nor to “Sexual” selection; -they are “accidents.” Similarly, their horns are the witness of a -horn-producing “diathesis”: the various divergencies in curvature, -and in the form of their spirals, or the number of their encircling -rings—as in the horns of Antelopes—are to be interpreted in like -fashion. These twists and turns vary in the same way that the taste of -the flesh varies, and for the same reason; that is to say, they are not -the outcome of “Sexual Selection,” nor have they been brought about by -“Natural Selection” to serve the purpose of “Recognition marks,” as -Wallace would have us believe. - -But horns, as horns, apart from their “accidents” of curvature -and ornament, must certainly be regarded as the product of Sexual -selection, for having once started into being those individuals had -the best chance of leaving descendants which were best armed. The -possession of horns was not necessary to the maintenance of the -species; but such armature was essential among the males in securing -possession of the females. Other things being equal, the male with the -biggest horns wins the prize. Since these are also used as weapons of -offence, or rather of defence, in warding off the attacks of beasts -of prey, it might be contended that they are as much the product of -Natural selection as of Sexual selection. - -It soon becomes apparent that this interpretation must fail. In the -first place, if it were true, the females should be similarly armed. -In the second, in the presence of many of their enemies they are -useless. The Cape hunting-dog, for example, is more than a match for -any antelope. This ferocious animal kills his victim by running it -down, persistently tearing at its flanks, until at last the entrails -protrude and the horrid chase is ended. Furthermore, the horns are a -comparatively late acquirement of the species, as is shown in the case -of the Deer; for the earliest known fossil species were hornless. That -the females among the Oxen and many of the Antelopes possess horns is -an interesting fact, but it can only be regarded as another instance -of a character first acquired by the male and later, in successive -generations, transferred to the female. And it is to be noticed that -this transference is never found save in the cases where the character -in question has attained its maximum in the male. The transference -of weapons to the female is the more remarkable because there is no -evidence that they play any part in the struggle for existence, either -in securing mates or in warding off the attacks of enemies. Moreover, -these weapons in the female may exceed those of the male, in length, -though they are never so massive. They are to be regarded solely in -the light of ornaments. There are few more striking instances indeed -where the purely ornamental and the strictly utilitarian are so closely -associated. - -[Illustration: Plate 4. - -_By the courtesy of Rowland, Ward, Ltd._ - -WEAPONS OF OFFENCE. - -Horns of various types furnish the most conspicuous of the “Secondary -Sexual Characters” of the ruminants. In the Deer only are these -branched. In the “hollow-horned” ruminants they are either lance-like -or more or less spirally curved, or they may form more or less open -loops. - -1. Black-tailed Deer. 2. Hangul or Kashmir Barasingha Deer. - -3. Greater Kudu. 4. Black-buck. 5. Saiga Antelope, remarkable also for -its curiously swollen nose. 6. Marco-Polo’s Sheep. - -[Face page 52.] - -Attention may now profitably be turned to the behaviour of these -interesting tribes when under the alluring influences of love. - -Tradition and the poets have contrived to persuade us that the fever -of Love becomes epidemic in the spring. This, however, is by no means -true, at any rate in so far as what we are pleased to call the “lower -animals” are concerned. For with many, as for example the Deer and the -Bats, this fever is not aroused till the time of autumn plenty. With -regard to the deer, we can find a reason for this. It is determined in -part by the period of gestation, and in part by the peculiar character -of the most conspicuous of the male secondary sexual characters—the -antlers. The deer, at any rate of the northern hemisphere, carry their -young about eight months. Now it is important that they should make -their entry into the world just as the food supply is increasing and -the temperature is rising. With the summer before them the young have -time to gather strength for the encounter with their first winter. We -have a striking witness to the truth of this contention in the fact -that when the Indian Spotted Deer, or Chital, was first introduced into -Europe, nearly all the fawns perished owing to having been born in -winter; later, the females took to calving in spring, and from thence -onwards the species has held its own among us. - -As touching the stags. The antlers, as everybody knows, are shed -annually, and their renewal entails a very considerable strain on the -system. As a consequence, it is necessary that this period of stress -should fall after the trial of winter is overpast, and with the genial -summer before them. From the end of March, when the old weapons are -shed, till July, the masterful males of the community wander at large, -seeking seclusion and avoiding all occasion of quarrel; for they -are not only defenceless, but threatened with disaster should any -accident befall the growing horns, which, during their formation, are -exceedingly sensitive. Even a slight blow would not only spoil their -shapely proportions, but, further, might render them useless in the -warfare that is before them. - -With some species this desire to go into retreat is more marked than in -others. The Red-deer, and the Wapiti, on the one hand, and the Moose on -the other, well illustrate this. The two first-named pass the winter in -herds, in the case of the Wapiti numbering many thousand individuals; -no other species, indeed, is so markedly gregarious. With the advance -of the spring, however, all is changed, for the males withdraw from -their companions to suffer humiliation in seclusion. As chill October -arrives, a striking alteration in their demeanour becomes apparent, -at any rate in the case of the older males. The new antlers are now -hardened, and the blood supply, which has hitherto been building up the -new weapons, is cut off. As a consequence, the “velvet,” which till now -has been directly concerned with the growth of the antlers, dies, and -peels off the underlying bone. To facilitate this work of cleaning, -the animal rubs them, first against the stems of saplings, and, later, -against larger trees, and even rocks, till at last they are ready for -“battle, murder and sudden death.” The “rutting” season, in short, has -commenced. And with the final completion of the antlers other signs -of that approaching frenzy, which is soon to establish itself, become -apparent. The most striking of these are the swelling of the neck, and -a marked increase in the mane thereof; while the voice enlarges its -compass enormously, whereby the females, so long neglected, are now -feverishly sought for. - -[Illustration: Plate 5. - -_Photo by G. W. Wilson Co. Ltd., from “The Living Animals of the World.”_ - -MANCHURIAN WAPITI “CALLING.” - -The “stags” do not begin to call for mate’s until the horns have more -or less completely shed their velvet. - -[Face page 54.] - -The Red-deer, maddened with desire, scours the country, calling as he -travels with a loud musical roar, ever and anon impatiently listening -for the tremulous response of females hardly less anxious to mate -than himself. One after another is speedily added to his harem, but not -without conflict. For sooner or later he catches the call of another -stag in like case. A jealous fury at once takes possession of him, and -the call, intended as a message to mateless hinds, becomes translated -into a challenge to fight for the mates possessed. Each of the now -infuriated challengers makes all haste to come to blows, and speedily -they are rushing headlong on one another to meet in a crash of antlers. -Then follows a test of strength, a sort of tug-of-war reversed, for -each strives to push the other to his knees, and succeeding, to deal -a deadly sideways thrust at the kneeling adversary’s heart with the -spike-shaped brow-tines. This attempt, however, is rarely achieved. Yet -not seldom such encounters become a duel to the death, and one in which -both die, for in the remorseless tilt at one another the antlers of one -may spring apart, and then close in on those of the other. Once this -happens, it seems to be rare indeed that they can be extricated from -this close embrace. With heads thus locked, they sway, and twist, and -tug, not now for the mastery, but for life itself. But as the hours run -they become more and more exhausted by their efforts, weaker and weaker -from loss of food and rest, till finally death releases both. - -A male having once succeeded in forming a harem, will commonly contrive -to repeat his success year after year, withstanding all comers. But -sooner or later his vigour wanes and he is ousted by another and -younger male. Not else would the stamina of the race be preserved. It -is considered a moot point, however, whether physical strength and -sexual potency run at the same pace; for it is believed by some that -a stag will often contrive to hold a harem against all rivals after -his fertility has declined. This, however, is extremely improbable. A -lowering of fertility means a decline in the potency of the hormones, -and in the development of the secondary sexual characters, among which -are the antlers, which are by no means negligible factors. That they -are not all-important, however, seems to be shown by the fact that, -occasionally, stags appear in a herd which are congenitally unable to -produce antlers—a reversion to the ancestral condition—and such are -said, occasionally at any rate, to be able to oust their formidably -armed rivals. This may be so, but the fact that “hummel” stags, as they -are called, are so rare is surely to be regarded as eloquent testimony -of the disadvantages of their unarmoured state. They become speedily -eliminated, in short, by “Sexual Selection.” - -After this outburst of sexual activity has spent itself, the various -harems, with their lords, amalgamate; all living in peace through the -winter. The stags retain their antlers at this season, partly as a -protection against predatory enemies, such as wolves, and other large -carnivores, which would otherwise play havoc in their ranks, and partly -because the cold of winter and scanty fodder would inhibit the growth -of new antlers or reduce their size. With the return of spring the -dangers of attack are lessened, temperature rises, and food becomes -once more plentiful. Then the inevitable disarmament takes place. - -The Red-deer, though mature at six, does not reach his prime till his -eleventh year, and from thence till his fifteenth or sixteenth year is -at his best. The hinds mature earlier, and appear to be fertile for a -much longer period. At any rate, a wild hind in Jura, known by certain -peculiarities of its ears, during twenty-one years produced twenty -calves. She was killed at last with a calf at her side, but was thin -and haggard-looking. She was, therefore, not less than six-and-twenty -at her death. The calves, it may be mentioned, are born in May and June. - -Old stags shed their antlers, it is remarked, earlier than young ones. -And this is an advantage to the species, since it prevents premature -breeding on the part of sexually precocious but immature males, and -limits competition to the adults. - -What obtains in the case of the Red-deer obtains also with minor -variations due to environment, climate, and so on, in the case of all -other deer. The life-history of the Wapiti, as might be supposed, -differs only in detail from that of the Red-deer. But during the winter -they form vast herds, numbering thousands. It may be that in primitive -times the Red-deer was no less numerous. But in this country, at any -rate, conditions favourable to the maximum development, either in -bodily size, or in the massiveness of the antlers, have long since -passed away. Even in the Highlands of Scotland the conditions of -existence have entirely changed owing to disafforestation. Deer are -essentially forest dwellers. But the “deer forests” are such only in -name, and for the most part the wild stags of to-day must get what -shelter they can from rocks and inequalities of the ground. From -this cause, and from the very natural desire of the owners of such -“forests” to secure the finest heads in each year, the whole race -has deteriorated. How great a change has come over it may be seen by -comparing the heads of British Stags with those from German forests, -where the conditions of existence are more favourable. If we turn to -the records of the past we find that the antlers found in the fens, -turbaries, and caverns of our islands are vastly larger, heavier, and -carry a greater number of points on the sur-royals, than do those of -the existing Scotch stags. - -Having regard to the fact that hundreds, and in the distant past -thousands, of antlers were shed annually, the comparative rarity of -these weapons in the haunts of deer excites comment. This is accounted -for by the fact that they are greedily eaten by their late owners, -apparently, though unconsciously, for the sake of their bone-producing -qualities. - -By way of contrast with the Red-deer and Wapiti, we may take the Moose -(_Alces machlis_), which at no time, and nowhere, attains to large -herds. This is explained by the relatively restricted food supply which -obtains in the haunts of these creatures. For they frequent the margins -of streams, feeding largely on willows and birch. From the shortness -of their necks, and the great length of their legs, they cannot crop -grass and other short herbage, for unless they kneel they cannot reach -the ground. Hence it is obvious that though their geographical range -may be wide, their numbers are kept rigidly in check. They would be -fewer still but for the fact that, unlike other deer, they glean no -small amount of food from the water, wading out to feed upon aquatic -vegetation. The roots of water-lilies are especially sought for, and to -obtain these the animal will often disappear entirely under water. - -As a consequence of the limited food supply the Moose lead solitary -lives. On the Eastern side of America, where the winter is severe, a -few individuals, generally a family party, will “yard up,” or make a -fortress for their mutual protection by trampling down the snow over a -restricted area. But in the Yukon district, my friend Mr. F. C. Selous -tells me this is never done. - -The rutting season of the bulls begins as soon as the antlers begin -to “peel.” What follows is practically a repetition of what has -already been related in regard to the Red-deer and Wapiti. And in this -connection it is interesting to note that the natives take advantage of -the period of desire in the bull to entice him to his death. Generally -this is done by imitating the call of the cow in response to the bull’s -anxious bellowing. But in Southern Alaska the opposite side of his -nature is played upon. This is done by scraping or beating the bushes -with the shoulder-blade of a Moose in such a way as to reproduce the -sound of a bull cleaning his horns. The very suspicion of a rival -enrages him, and, rushing in a blind fury in the direction of the -tell-tale sounds, he speedily falls a victim to the trick which has -been played him. - -That the mating period is the most critical, and most searching in -the whole life-history there can be no doubt. Every faculty during -this time is put to the test, and from the time of sexual maturity -until old age is at last attained it is an annual test. Alertness is -all important. Other things being equal, success falls most certainly -to those individuals with the keenest perception, and quickest -interpretation of sight, sound and smell. - -One is puzzled at what seems a concession of Darwin’s to the Lamarckian -theory of the inherited effects of use in this connection. For in -discussing the bellowing of the stag in “The Descent of Man,” he -remarks that it “does not seem to be of any special service to him, -either during courtship or battles, or in any other way. But may we not -believe that the frequent use of the voice, under the strong excitement -of love, jealousy and rage, continued during many generations, may -at last have produced an inherited effect on the vocal organs of the -stag, as well as of other male animals?” All the evidence goes to show -that the production of sound, and the instant interpretation of its -significance, is a matter of the highest importance. In the case of the -Moose, for example, the noise occasioned by the cleaning of antlers -provokes the same frenzy as at another time is aroused by the voice. -Dullness of perception not only in these matters, but at all times, is -fatal. - -As touching the less conspicuous secondary sexual characters of Deer -more must be said presently. For the moment the antlers must retain -our attention. Time was when the Deer lacked these appendages. When -they first appeared, in the now extinct species of the Middle Miocene -period, they were no more than short prongs. Later, one of the prongs -became elongated, and developed short branches or “tines,” which, in -succeeding species, became more numerous, while at the same time, with -the gradual evolution of more and more species, these antlers assumed -new features both in the matter of size and in the character and number -of the “tines,” a development which has reached its maximum to-day. But -apart from these specific variations, which have given us such types as -those of the Roe-deer, Red-deer, Wapiti, Caribou, Moose, Fallow-deer, -Sambar, Schombergk’s deer, the strange Milou-deer, Elds-deer and -Mule-deer, each species displays a quite remarkable range of variation -in regard to its particular type of antler. Nowhere, perhaps, is this -more strikingly marked than in the case of the Caribou and Moose. No -doubt this feature is due largely to the fact that the horns are shed -annually, and that the variations are due, in part at any rate, to -temporary environmental conditions, such as food and weather. But -these apart, individual peculiarities are constant, reappearing with -more or less exactness each year. - -[Illustration: Plate 6. - -_Photo by Lord Delamere, from “The Living Animals of the World.”_ - -GROUP OF BEISA ORYX. - -The lance-like horns of these animals can be used with deadly effect, -even against lions. - -[Face page 60.] - -In contemplating these facts one asks: What are the underlying factors -of this variability? What is the significance of the branching? What -end is attained by the annual shedding? That the antlers constitute -very effective weapons of offence there can be no doubt, and one is -inclined to regard the branching as the outcome of natural selection, -on the assumption that branched antlers would be less deadly than -lance-like weapons. It would perhaps be tempting to accept this -interpretation as all sufficient were it not for the evidence afforded -by the hollow-horned ruminants. The Oryx and the Kudu, for example, are -lance-bearers, and therefore show conclusively that stags similarly -armed might well have continued to survive in spite of the foils which -the “tines” provide. Darwin, long since, guardedly suggested that -while these weapons primarily served for offensive purposes, their -elaborate systems of branching might have been brought about by sexual -selection. That is to say, the extreme beauty of the weapons may excite -the admiration of the females as well as our own. Granting this, he -inferred they might have played an important part in elaborating the -branching by constantly displaying a preference to mate with those -males possessed of the largest and most branched antlers. But there -are many and serious objections to this suggestion, and the most -important of all is the fact that the female is allowed no choice in -the selection of her lord and master. We can, then, only regard the -antlers of deer as another instance of the survival of a “fortuitous” -but inherent variation, which survived because, whatever the defects -thereof, they proved advantageous in the struggle for existence. - -Having regard to the fact that so many of the females among the -hollow-horned ruminants have acquired horns, it is somewhat remarkable -that in the Reindeer alone among the deer are these weapons normally -possessed by the female. The gradual transference to the female of -features which were originally secondary sexual characters in the -male is an occurrence which is met with in every group of animals. In -writing “The Infancy of Animals” I gave a number of instances of this -kind. But the case of the Reindeer affords a more than usually striking -illustration of this curious sequence; and this because rudiments of -antlers are to be met with among the females in several different -species of Deer to-day. They have been found in the females of both -Roe- and Red-deer, though such cases are rarely met with. As a rule -this assumption of the male secondary sexual characters by the female -occurs only in very aged animals, or as one of the sequelæ of diseased -ovaries and consequent sterility. But at least one instance is on -record of a doe Roe-deer which possessed small antlers while pregnant. -Thus, then, we gain a further insight into the process by which the -female slowly assumes the outward attributes of the male; that is -to say, the secondary sexual characters appear first in the male, -and as seasonal characters. Sooner or later they become permanently -established. By the time they have become firmly fixed in the male, and -apparently not till then, they appear in a dilute form during senility, -or in consequence of ovarian disease, in the female. Having once -started, however, they appear earlier and earlier in the life-history -of succeeding generations of females, and at last in the juvenile -stages of both sexes. - -The hollow-horned ruminants, which must now be considered, afford some -very striking facts in regard to these “secondary sexual characters,” -more especially in so far as horns are concerned. In the first place -these weapons are permanent structures, taking the form of a bony core -ensheathed in horn, with which we may compare the temporary covering -of velvet in the deer: in the second, they are unbranched. The only -exception to this rule is furnished by the Prong-horned Antelope, -wherein the sheath is both annually shed, and branched. The branching, -however, is very slight, taking the form of a short forwardly directed -prong about half-way up the sheath, which is borne on a long bony -pedicle recalling that of the Muntjac. The shedding is due to the -formation of new horn material at the base of the old sheath, which -is gradually forced off by the growth of the new tissue. Structurally -the horn of this remarkable Antelope differs somewhat from that of its -relatives. - -As may be seen in Plate 4, in the form of the horns the typical -hollow-horned ruminants present an exceedingly varied range, and one -often of great beauty in the matter of curvature. That they serve as -formidable weapons of offence was demonstrated during 1912, when, -according to the Annual Report of the Government Game Reserves, -published by the Pretoria Government, the game warden, Major Stevenson -Hamilton, reported of the Antelopes that “many carcases of males of -almost all species, killed in single combat with rivals, were found -during the mating season, untouched by anything except vultures.” As -a rule, however, these animals, like the Sheep and Goats, and their -larger relatives the Cattle, seem to avoid a duel to the death. One -or two instances as to the general character of these combats for -the possession of mates must suffice. Thus the late A. H. Neumann, -a hunter of experience, remarks that he once or twice saw conflicts -between the Topi (_Damaliscus jimela_), an ally of the Hartebeestes. -The two rivals would stand a little apart, affecting, apparently, to be -unaware of one another’s presence. Suddenly they would rush headlong at -one another, bringing their heads together with a clash, each, at the -same moment, falling on his knees. - -Major Powell Cotton, again, once witnessed an affray between two Beisa -Oryx. Here the master bull of the herd was infuriated by the advent of -an intruder in his harem. Time after time they dashed at each other, -their foreheads meeting with a thud; then, with horns interlocked, -they wrestled fiercely; then, separating, they charged again. Yet -neither, he remarks, tried to use his lance-points, as they do when -cornered by man or beasts of prey. Nevertheless, encounters of a more -sanguinary character appear to be by no means rare, for it is no -uncommon experience of hunters to kill bulls of this species in which -one eye has been burst by a horn-thrust. Another peculiarity of these -animals is the extreme thickness of the hide of the neck and withers, -which seems to afford a shield against such spear-thrusts during -these battles. How powerful is the thrust of these weapons, and how -efficiently they can be used, is shown by the fact that lions in making -an attack on an old bull are often severely wounded, or even killed. -And there are many instances on record of cases where both the lion and -his intended victim have died together, the Antelope having been unable -to withdraw his horns from his adversary’s body. The beautiful Pala -Antelope fights furiously with rival rams, and the vanquished, as with -so many of the Antelopes, form herds by themselves, till one by one -they gather strength and skill enough to establish their right to mate. - -[Illustration: Plate 7. - -_Photo by courtesy of the Duchess of Bedford, Woburn Abbey._ - -ELAND COWS. - -Among antelopes the females commonly bear horns, which may be even -longer than in the males, though less massive. - -[Face page 64.] - -[Illustration: Plate 8. - -_Photo by courtesy of the Duchess of Bedford, Woburn Abbey._ - -AMERICAN BISON - -The “Secondary Sexual Characters” of the male are here conspicuously -developed, and are seen in the massive fore-quarters and enormous -head.] - -The Elands present some puzzling features, for both sexes bear large -horns, and they are very massive in the bulls. Yet these animals -are generally described as the most inoffensive of all the horned -ruminants. That the horns are used to any extent in conflicts between -rival males seems doubtful, inasmuch as this species is remarkable for -the development of an enormous “dewlap,” a thin pendulous fold of skin -which runs from the throat to the chest. Such a form of “ornament”—for -in this light we must regard it—would be dangerous, indeed, when much -fighting was to be done. Nevertheless, it would be contrary to all -our experience to conclude that weapons so well developed as are the -horns of the bull Eland were entirely useless. This is a matter which -decidedly calls for further investigation. - -That our knowledge of that most important period of life of the larger -mammals, the period of sexual exaltation, is lamentably incomplete will -be realized by anyone who seeks enlightenment on this subject. Most -of the meagre information we possess has been collected by travellers -and sportsmen, neither of whom have the time to devote to the long and -laborious watches that a fuller history demands. Every now and then a -glimpse is afforded of this period of the life-history which brings -home in a very convincing fashion, how little is really known. It -seems certain that the fighting hitherto described is to be regarded -as but a phase of a cycle of events which takes place at this time. -Thus, for example, the old naturalist and traveller Schweinfurth tells -how he once encountered a herd of Hartebeest which were apparently -effervescing with animal spirits, for they kept running around in -couples, like horses in a circus, using a clump of trees as a pivot. -Others, in groups of three or four, stood by, interested spectators. -After a time these, in turn, took their places and ran round, two at a -time, in their own circuit, and in the same fashion. Their evolutions, -he says, were so regular as to suggest the guidance of some invisible -ring-master. These gyrations may be regarded as an erotic dance. The -Sambar, under like excitement, will stalk about with erected tail, -outstretched muzzle and everted face glands, and the Black-buck, among -the antelopes, behaves in like fashion. - -It cannot be supposed that these quaint performances are peculiar to -the species in which they have been observed, but rather it may be -inferred that similar antics, besides others yet to be discovered, -are performed by all. Their purpose seems plain enough, for they must -be regarded surely as aphrodisiacs, excitants to pairing. They recall -the erotic dances of savages, or the ceremonial orgies of ancient -civilizations. Such performances, on an even more elaborate scale, are -to be met with among the birds. - -So far, in describing the horned ruminants, the horns only have been -considered; but these animals display yet other secondary sexual -characters, which, while less conspicuous, are yet no less important -during this critical period of life. Some, as for instance the canine -teeth possessed by some of the deer, are decidedly puzzling. While -absent, or vestigial, in most, in a few they are greatly developed, and -this, too, in species which possess relatively large horns, as in the -Muntjac. It seems difficult to believe that the co-existence of these -very different kinds of weapons can be of vital importance to their -possessors; yet unless this be so, one or other would surely have -degenerated. It is significant that in the hornless Musk-deer these -teeth attain to a very considerable length, at their maximum as much -as three inches. That they are used by rival males, and with effect, -is shown by the fact that the hides of these animals are often found -scored by deep lines cut by these tusks. In those aberrant ruminants, -the Camels, quite formidable tusks are present both in the upper and -lower jaws, and these are used with effect whenever occasion demands, -and often when it does not. - -The armoury necessary for successful love-making contains yet other -weapons, evolved to supplement physical force, and more subtle in their -effect. Such are certain skin glands which, at the rutting season, -secrete a copious flow of a creamy, or semi-fluid matter, and pungent -odour. In the deer the more important of these are found in the deep -pit, or “larmier,” which opens in front of the eye. In the Musk-deer, -however, this secretion has a most powerful odour of musk, and is -formed in a pouch, or “pod,” of about the size of a small orange, under -the skin of the abdomen. The secretion, which is formed by the male -only, is of a chocolate colour, and of about the consistence of moist -gingerbread. It has a most pungent scent, and when diluted forms the -basis of many of our most powerful and most highly-prized perfumes, on -which account, it may be mentioned, this animal has for generations -been submitted to a most unrelenting persecution. But that is another -story. - -In most of the antelopes the principal scent gland is seated in a pit -in front of the eye, as in the deer. In some, as in the Gnu, it forms -instead a swollen, tumid area, oblong in shape, instead of lying in a -pit. In the Reedbuck it is placed around the bases of the horns; and -in the Rocky-Mountain Goat it forms a great bare cushion behind the -horns. All have more or less well-developed glands seated in the skin -between the toes. But, wherever placed, the secretions thereof are more -or less completely suspended save during the breeding season, when they -are poured forth abundantly. The precise rôle they play is by no means -certainly known. It seems reasonable to suppose that, in the first -place, the odour they disperse enables the males to announce their -whereabouts to the females seeking mates, should they fail to hear -their bellowing. But the antelopes, for the most part, unlike deer, do -not, the year round, lose touch with one another; so that it must be -concluded that these odours serve as excitants to the act of pairing, -and we know that the sense of smell plays a very important part at -this time, which, so far as these animals are concerned, is the only -period which comes more or less exactly within the meaning of the term -“courtship.” - -That scent among the antelopes holds a really important place is shown -by the fact that the bull of the common Eland intensifies his natural -odours by micturating upon the mass of long hair which grows upon the -forehead. To do this the head is bent down and turned tailwards, in -order that the tuft should receive its due urinary spray! And goats in -captivity exhibit the same curious habit. In them, indeed, it is often -pushed to such an excess that blindness results, so that the animal has -to be slaughtered. - -While in many cases these odours are imperceptible to human nostrils, -in others this is far from being the case. Among the ruminants the goat -is particularly odorous. So also are the giraffe and the water-buck, -both of which may be detected by their smell at considerable -distances. And these emanations are most noticeable in the males and at -the breeding season. The bull elephant, both in the Indian and African -species, during the breeding season produces a copious flow of aromatic -matter from a gland which opens above the eye in the form of a tubular -aperture large enough to admit a pencil. This aperture in the African -elephant is remarkable for the fact that it is invariably found to be -“plugged” with numerous spines of the acacia, which have from time to -time found their way in as the animal was forcing its way through the -dense undergrowth. This extraordinary fact was first noticed by Mr. F. -C. Selous, and has since been confirmed by Dr. Einar Lonnberg. - -It is probable that the “bloody sweat,” which at times covers the hide -of the Hippopotamus just after leaving the water, is associated with -the period of rut. This mysterious exudation is accompanied by small -crystals; but though red in colour, it contains no blood. So far no -reasonable explanation for this remarkable phenomenon has ever been -given, but probably it will be found to be associated with the sexual -activities and is possibly odoriferous. A precisely similar exudation -occurs in the neck of the male of the Red Kangaroo. - -That these secretions play an important and perhaps variable part -in the selection of mates seems demonstrated in the case of an -incident related to me by my friend Mr. John Cooke, who some time -ago was watching a flock of some three hundred sheep while it was -being driven by the shepherd and his dogs into a field. As soon as -they were securely shut in, and the shepherd had gone, three rams -who were included in the flock at once began a three-cornered fight. -One, presumably the youngest, was soon vanquished. The other two -soon settled their differences, and the clashing of horns was at once -followed by a very different performance. The master ram began to run -in and out among the ewes, sniffing at each, and driving out those -whose odour most pleased him. Having at last satisfied himself with a -harem of about one hundred, the second ram was allowed to make a like -choice, and behaved in a like manner, leaving the remainder to the -ram which was first vanquished. May we take it that the strongest and -oldest rams selected the youngest ewes, and the oldest were left to the -youngest, and first conquered ram? By some such rough and ready method -of selection Nature may contrive that the immature male shall do as -little harm to the race as possible by mating with the oldest, and in -many cases barren females. - -Our survey of the “hoofed” animals has so far been confined to the -ruminants. Space must now be found for a brief review of what obtains -under like circumstances in the case of the great pachyderms—the -Elephant, Rhinoceros and Hippopotamus; the Pig and the Camel. - -[Illustration: Plate 9. - -_Photo by Lord Delamere, from “The Living Animals of the World”_ - -ELEPHANTS. - -The sexes differ but little in general appearance: and chiefly in the -superior size of the male and his more massive tusks. - -[Face page 70.] - -As to actual “courtship” among these animals practically nothing -is known; but the varied and formidable weapons which they possess -are enough to show that the secondary sexual characters play a very -important part in the preliminary capture of mates. That they may -also be used for the more prosaic purpose of securing food is nothing -to the point. In the Elephant, for example, the tusks are sometimes -of enormous size and weight, specimens of eleven feet in length and -weighing as much as two hundred and fifty pounds are on record. They -are used for cutting through the bark of machabel trees, which is then -seized by the trunk and torn off, for elephants are extremely fond of -this bark; and they are also turned to account in breaking up roots -which have been exposed by digging with the fore-feet. But this is -certainly not the main purpose of such weapons. On the contrary, their -use is primarily as weapons of offence between rival bulls. As one -would expect, they never attain to a very large size in the female, but -that they are large enough to serve her at need is shown by the fact -that a portion of a tusk, evidently of a cow-elephant, was once found -embedded in the jaw of a bull. There can be little doubt but that this -was broken off in an endeavour to repel the advances of a too amorous -male, for, as with all animals, pairing is impossible without the -consent of the female, and this is never accorded until she is desirous -that it should take place. As a preliminary to this, an amorous -dalliance is perhaps the invariable rule among animals, and this takes -many and often strange forms. The Elephant affords a case in point. For -the late A. H. Neumann once came upon a pair which were evidently, as -he says, “love-making.” Creeping upon them noiselessly, he found the -male fondling his mate with his trunk, and then, standing side by side, -they crossed their trunks, and put the tips thereof into each other’s -mouths, the elephantine form of kissing. Deer, cattle and horses, cats -and dogs, constantly lick one another under like circumstances. - -Superficial secondary sexual characters are wanting both in the -Hippopotamus and the Camel. Both, however, possess a formidable -armature of teeth which are capable of inflicting very severe wounds. -In the Hippopotamus the canines are of enormous size, and their -punishing power is further strengthened by the fact that they work -in opposition to a pair of similar teeth in the lower jaw; they cut -like a pair of shears, the upper closing upon the lower pair with -the precision of scissors-blades. In addition, the lower jaw develops -two long, blunt-pointed, ivory spikes, which are scarcely less to -be dreaded. With these weapons the bulls fight furiously, and it is -no uncommon thing to find vanquished males frightfully mauled, the -hide being lacerated from head to tail. Protection, in a measure, is -afforded by its enormous thickness, but the great folds and pleats of -skin seen in the Rhinoceros are never developed. The females, however, -are similarly armed, and the teeth are nearly as large as in the males, -which is a rather unusual occurrence. - -The Swine, which are near relations of the Hippopotamus, in like manner -develop huge pointed canines, and these reach their maximum in the -great Wart-hogs of Africa. But in the swine the mechanism differs, -for although the canines are closely opposed, the shaft of the upper -teeth curves upwards, and the lower teeth are much smaller than the -upper. In fighting, these animals do not bite, like the Hippopotamus, -but use the upper canines to rip up their antagonist with a sudden, -swift upward and sideways movement of the head. How dangerous is the -wound thus inflicted those who have hunted the wild-boar know well. A -curious exaggeration of this arrangement of the teeth is seen in the -Babiroussa. Herein the upper canines grow directly upwards, actually -piercing the upper lip as in the case of the downwardly growing tusks -of the elephant. That these teeth, however, are of any service in -fighting is doubtful, for the upper tooth curves upwards and backwards -in a semicircle so that the points are harmless. The tusks of the lower -jaw, however, are extremely long and pointed, though their wounding -power is limited by reason of the upper teeth. This may account for -the fact that the head, the part mostly attacked by enraged -boars, presents no sort of armature designed for defence; while in the -Wart-hog, on the other hand, great solid bucklers of hide stand out -on either side of the head below the eyes, giving the animal a most -repulsive appearance, but affording him a very present help in time -of trouble. In the wild-boar, where the tusks are shorter, no such -protective armature is needed. - -[Illustration: Plate 10. - -HEAD OF MALE WART-HOG. - -In the “Swine” family the canine teeth are always greatly developed, -but they attain to their maximum, relatively, in the Wart-hog.] - -[Illustration: - -_Photos by Scholastic Photo Co., from “The Living Animals of the World.”_ - -MALE AND FEMALE BABIRUSA. - -A characteristic of this pig is the peculiar development of the tusks -in the male, the upper pair of which grow through the lips and curve -upwards. - -[Face page 72.] - -[Illustration: Plate 11. - -_Photo by Lord Delamere, from “The Living Animals of the World.”_ - -SOMALI ZEBRAS. - -The Zebras, unlike their cloven-hoofed relations, have no weapons, save -for inter-tribal conflicts. Yet they have been as successful in holding -their own against lions and other predatory animals as species provided -with horns. - -[Face page 72] - -[Illustration: Plate 12. - -_Photo copyright by A. H. Bishop._ - -GIRAFFE. - -The horns of this animal can prove formidable weapons of offence on -occasion, though they are useless against predatory animals.] - -While the ungulates, or hoofed animals, are peculiar in the development -of horns as weapons of offence, they are by no means singular in the -use of teeth for this purpose. In some cases, as in the Muntjac, both -forms of armature are present. The only other instances where teeth -in this group of animals are used for offensive purposes are those -furnished by the Camel and the Horse. But here they do not exhibit -that excessive size which is met with in the Elephant, and some of -the Swine. In both the Camel and the Horse it is the canine which is -used, and both jaws are similarly armed. Since the camel has no upper -incisors, the part played by the teeth is beyond dispute; but it has -been contended that the horse uses his incisor or “front-teeth” alone -when fighting. But this is not so; the canines can, and do, inflict -ugly wounds, as is shown by the necks of zebras. - -A further method of defence among the larger Ungulates, at any rate, -is resorted to when hard pressed: and this is the use of the hoof in -kicking. Giraffes kick both after the usual fashion and in striking -downwards with the fore-foot. And an interesting demonstration of this -has been furnished by Mr. F. C. Selous in his delightful “African -Nature Notes.” He relates that on one occasion he came across a calf -only a day or two old, with its back broken. From scratches on the -calf, and the footprints on the ground in its vicinity, he was at once -enabled to gather the cause of its terrible plight. In a word, it had -been attacked by two leopards, and the mother, in an endeavour to beat -off the assailants with a blow of her fore-foot had accidentally struck -her offspring. Horses, Cattle, Antelopes, Camels and Elephants can all -kick with precision and effect. So far as the evidence goes, however, -this is a method of defence used against beasts of prey, and is rarely, -if ever, employed in conflicts between rival males. Females persecuted -by the undesired attentions of amorous males, however, do, as we know -from the case of domesticated animals, use this device to defend -themselves. - -It is not difficult to account for the origin of such secondary sexual -characters as manes, beards, tusks, and brightly-coloured areas of -skin, though whether our interpretations are really correct is another -matter. But no attempt to explain the origin of horns has yet achieved -a like degree of persuasiveness. These weapons appear only in the -Ungulates, a group which has, in past times, given birth to some very -extraordinary types of head armature of this kind. These must be -excluded from the present discussion; suffice it to say that, as usual, -they were the adjuncts of the males. According to current theories it -is supposed that these weapons arose as the result of the action of -sexual selection. It is assumed that the hornless ancestors of now -horned ruminants fought for their mates by “butting” with the forehead. -Naturally, other things being equal, the thickest skulled combatants -obtained the mastery. Any tendency to develop frontal “bosses” of bone -would further enhance the chances of success, and would, indeed, soon -become necessary for survival. And from such “bosses” the passage to -horns and antlers forms an easy transition. Just such incipient horns -or “bosses” actually make their appearance in the domesticated horse: -but these animals never butt at one another. If, however, we regard -horn-production as an inherent diathesis of the ungulate somatoplasm, -we have an intelligible basis for the explanation of horn development. - -The formidable horns of the Rhinoceros are of a totally different -character, being solid structures formed by hairlike agglomerations, -firmly fixed upon a roughened area of the nasal region. These weapons -play a very important part in settling disputes between rival males, -but on other occasions demanding offensive tactics the Indian -Rhinoceros at any rate seems to depend rather on his power of wounding -by means of the chisel-shaped lower incisors. These, by means of a -swift lateral movement of the head can be made to inflict most terrible -gashes, as those who hunt with elephants well know. It is quite -possible, however, that the teeth are also thus used during struggles -for supremacy. And this may perhaps account for the enormous bucklers -of skin developed by the Indian Rhinoceros, but only indicated in the -case of the African species. - -All the larger Ungulates, and many of the smaller species, are -polygamous. The Rhinoceros, and all of the swine-group save the -Hippopotamus, among the larger species are exceptions to the rule. The -preponderance of females which this implies is generally supposed to -be due to the losses sustained among the males by fighting during the -struggle for mates. The case of horses, however, seems to militate -against this view, for though they undoubtedly fight furiously, no -evidence is forthcoming to show that such conflicts terminate fatally. - -Were it possible to secure the necessary data it would probably be -found that polygamy, and polyandry, are determined solely by the -numerical proportions of the sexes: the excess of males or females -being due neither to “Natural” nor “Sexual” Selection, but to inherent -peculiarities of the germ-plasm tending to produce an excess of males, -or females, as the case may be. - -Finally, all the evidence goes to show that it is a mistake to -suppose that polygamy is due to the excessive sexual avidness of the -males, which impels them to first essay the overthrow of all possible -rivals, and then to appropriate every female within their sphere of -influence, holding them by force. On the contrary, this plurality of -mates is thrust upon them. And this because the females, impelled by -“mate-hunger,” attach themselves to the nearest male within call: the -size of the harem depending on the number of available males. The -battles which are fought between rival males are no more sanguinary -than in the case of monogamous species. This contention is well -illustrated by the African Wydah-birds (_Vidua_), which are markedly -polygamous, though they have no special weapons of offence. In -districts where males are numerous the harem will not exceed eight, or -ten, females; where males are scarce this number may be increased to -fifty. In like manner the varying number of hinds accompanying a stag -are to be regarded, not as an index of his prowess, but of the scarcity -or abundance of males in the neighbourhood. - - - - -CHAPTER V - -THE LION AND HIS KIN - -A Surprising Relationship—The Lion’s Mane—The Sabre-toothed Tiger—Some -Theories about Origins—Sea-lions in Love—Some Strange Ornaments—Whales -and Weapons. - -That the Lion and the Lamb could possibly have been derived from the -same stock seems incredible: yet such is the case, though the pedigree -is now well-nigh lost in the mists of a hoary antiquity. It is not -surprising, then, that in their present-day garb they should show so -little in common. Nor is it strange that among their many points of -divergence the one should differ so conspicuously from the other in the -matter of secondary sexual characters. For when these are conspicuous -among the Ungulates they usually take the form of horns, of which the -Carnivores have no need, for the teeth and claws whereby they win their -daily portion of meat make equally serviceable weapons of offence when -turned against their own kind. - -Among the larger Carnivora, the Lion alone displays any obvious -distinction between the sexes in the matter of ornament, and this in -the form of the well-known mane. Darwin, and later authorities, have -regarded this as a shield to protect the great blood-vessels from -injury during battles between rivals. But it is not very clear that -this alone is sufficient to explain its presence, inasmuch as the Tiger -in this respect is defenceless. Mr. F. C. Selous long ago pointed out -that the varying abundance of the mane is due to climatic causes. Lions -which live in districts where the nights are very cold, as in high -table-lands, have large manes; those which occupy lower ground, where -the nights are relatively warm, have but a scanty mane. It is clear, -however, that the abundance of the mane is not determined by the need -for warmth, otherwise it would have been as well developed in the -female. Rather we must regard a low temperature as conducive to the -growth of long hair when a natural tendency to produce this is present. - -There are few men who can claim to have so great a first-hand -acquaintance with Lions as Mr. Selous, and he has pointed out to me -one significant fact which seems to show not only that the mane has -not been developed to serve as a shield when fighting, but that fights -between rival males must be rare. And this because of the absence -of any evidence in the shape of scars on the skin. With claws so -formidable as those of the lion, ugly wounds would certainly be made in -any prolonged conflicts, for the skin of this animal is very thin. - -In the now extinct Sabre-toothed Tiger the upper canines were of -enormous length, and it is not improbable that they, on this account, -exceeded the bounds of usefulness; that, while as weapons of offence -they may have proved exceedingly effective, yet they hampered the -animal when feeding. In many ways one is reminded by these weapons of -the huge tusks of the Walrus. These are blunt-pointed, and are said -to be used very largely for digging up the large clams and other -burrowing shell-fish on which this animal mainly feeds. They are also -used as levers to drag the huge body out of the water on to the ice. -As fighting weapons they are formidable, and the wounds they inflict -are sometimes serious. The polygamous habits of this huge creature may -account for the fact that they are so much larger in the males, wherein -they may attain a length of thirty inches, and a weight of eight pounds -a-piece. - -In connection with the monstrous tusks of the Sabre-toothed Tiger there -is a point which so far seems never to have attracted the attention it -deserves. And this concerns two small flanges of bone which project -from the lower border of the end of the lower jaw. In themselves they -are unimportant: they lie, it is to be noticed, parallel with the -points of the great upper teeth which descend on either side of them. -Their full significance is not apparent till we turn to the skull of -another extinct animal of quite another type—the huge Dinoceros, one -of the Ungulates. This animal was also armed with an enormous pair -of tusks, which also, when the mouth was closed, descended on either -side of a flange. In this case, however, the flange was developed to -such an extent that its free edge descended to the level of the point -of the tusk, thus affording it protection against injury. The really -striking feature of this curious down-growth is not apparent till an -attempt is made to explain its presence. What determined its growth? -It seems to furnish us with another of the many instances which are -to be found of the correlation of growth between unrelated parts, for -there is apparently no traceable connection between the growth of this -pair of teeth in the upper jaw and the development of the flanges of -the lower border of the jaw which are embraced by these teeth. In the -Sabre-toothed Tiger the inciting cause to this flange growth, whatever -it may have been, seems to have been much weaker than in the case of -Dinoceros. - -Naturally one asks, can the whole thing be explained by the theory of -Kinetogenesis promulgated years ago by Cope? That is to say, are these -curious down-growths the result of a response to a stimulus set up in -the lower jaw by constant lateral blows dealt by the tusks against -the side of the jaw during the lateral movements of the jaw when -feeding or ruminating? Such movements in an Ungulate would be frequent -and constant: hence perhaps the more striking result. On account of -the scissor-like action of the jaws in the Sabre-tooth such lateral -movements were far less extensive, and less powerful. But though this -explanation sounds plausible, it presents many difficulties. In the -first place it seems to commit one to the admission that the responses -of the Somatoplasm during the life of the individual are transmitted -to the germ-plasm: that, in short, the characters acquired by the -individual during its lifetime are transmitted to its offspring. And -there are insuperable difficulties in the way of such a theory. Yet, -it must be admitted, it is no less difficult to believe that this -correlation of growth is due solely to fortuitous variation, for one -cannot really conceive of a variation of this kind taking place in two -such different structures independently. Such a conception would have -been less difficult if the case of Dinoceros alone were known to us. -We could have supposed that, somehow, the lower jaw started to produce -its flange just as the teeth began to develop an excess of growth which -carried their points beyond the level of the jaw. But the Sabre-tooth -shows that the tusks had assumed a growth relatively exaggerated as -in Dinoceros, and yet the flange never attained to more than feeble -development. We cannot rest content with the theory that the flange is -due to the constant stimulus of blows struck against this region of the -jaw during the lateral movements which take place when feeding. Were -these animals alive to-day it could be tested by extracting the tusks -during infancy, when, the stimulus being removed, the flanges should -not appear. - -There are yet other aspects of the skull of Dinoceros which may well -be considered here. The first concerns the excessive armature of -horns, there being no less than three pairs supported on massive bony -cores; and the second the ridiculously small brain cavity which is -proportionately smaller than that of any other known mammal, recent or -fossil. This poverty of brain-power was probably one, if not the chief, -factor among the causes which brought about the extinction of this -strange beast. Even more formidable horns were borne by the extinct -Arsinoetherium. But this animal did not display the double armature of -horns and tusks. - -Among the Carnivora monogamy is the rule, though the Lion is -occasionally polygamous. But the Eared-seals (_Otaria_), or Sea-lions, -and Sea-bears afford a striking example of polygamous species and -of the ferocity they display when sexually excited. These animals, -moreover, are capable of the most astonishing powers of endurance and -vitality, exceeding indeed that of all other mammals. Since the habits -of the Northern Fur-seal (_Otaria ursina_) have been more carefully -studied than those of any others, it may serve as a sample of the rest. - -Living for the greater part of the year in the open sea, the old -bulls—animals of six or seven years old—are the first to seek the -“rookeries,” or breeding grounds, taking up their territory a full -month before the cows arrive. Later, the younger bulls appear, and the -more daring endeavour to force their way through the ranks of those -who have already taken up positions. This often leads to fighting, but -more usually nothing further than “bluffing” is indulged in, though -it is commonly supposed that very severe engagements take place. This -seems, however, to be only occasionally true. In due course, generally -about the second week in June, the cows begin to arrive, at first in -straggling numbers, but soon the main body puts in an appearance, and -before the end of the month many thousands of both sexes are crowded -along the foreshore. But yet, contrary to the generally accepted -belief, no serious fighting takes place. The bulls quietly seize the -females as they arrive. It would seem that the first arrival serves as -a focus of attraction for all later comers landing in the vicinity. The -bull holding the most advantageous post—that is to say, that nearest -the best landing-place—starts the collection and, unintentionally, the -distribution of the cows. Having seized the first arrival, he places -her by his side. As the later females arrive he gives each a most -cordial welcome, and then proceeds to round up his harem. But soon he -has more wives than he can continue to control. Do what he will, he -cannot be in two places at once; and thus it is that in rushing off to -chastise some covetous neighbour, one or more bulls on the opposite -side of his harem proceed to make captures from his horde. And this -system of abduction goes on over the whole rookery till all the cows -have been appropriated, leaving a crowd of envious bachelors in the -background who have not yet developed either courage or strength to -secure mates for themselves. - - -[Illustration: Plate 13. - -_Photo by New York Zoological Society, from “The Living Animals of the -World.”_ - -CALIFORNIAN SEA-LIONS, OR EARED SEALS. - -The “bulls” of the Eared Seal are much larger than the “cows”; they -have otherwise no very conspicuous “Secondary Sexual Characters.” - -Face page 82.] - -But within forty-eight hours of their landing the cows give birth to -their “pups.” And it is for this purpose, and not for mating, that they -come to land. Within a few days of the birth, however, the females -are “in use” again. This is the critical period in the life in the -rookery. For the bulls now become frenzied with excitement and fight -most viciously one with another, each hoping to possess himself of his -opponent’s harem. Each tries to seize the other by the fore flipper, -and, failing in this, the fangs are buried in the back. They hold -tenaciously, each trying to force the other to relax his hold; but -commonly this vice-like grip is maintained till the skin gives way, -leaving great bleeding rents. Sometimes the contest rages till one or -both is fatally wounded. Often during such duels an idle bull, hitherto -unable to secure a harem, will rush in and capture that of one of the -combatants! - -In the management of the harem the bull is an adept. Whether he has -five cows or fifty, he is, says Dr. Lucas, “master of the situation.” -His will is law. Not that it is always tamely accepted as such, but -the result is the same. If a cow becomes restless, and moves about, a -warning growl usually quiets her. If the movement is persisted in and -an attempt to escape evident, the bull is up at once with a show of -fierceness and in chase. He may simply strike her down with his open -mouth. Often in doing so his sharp canines tear a gash in her skin. He -may even seize her in his mouth and deliberately throw her, or carry -her back into the harem. If the cow thinks she has a chance to get away -she may try to outrun him. If she miscalculates the distance he seizes -her, after a few swift bounds, by the skin of the back, or by the hind -flipper, and tosses her, often torn and bleeding, into the family -circle. As a rule, however, she avoids this seizure by turning and -facing her lord and master, and biting him in the breast and throat. -But all to no purpose. In spite of her violent protests he pushes her -backwards before him into the fold. - -Sometimes in her efforts to improve her position she runs up to, and -is seized by, a rival bull. Her lord speedily asserts his ownership by -getting a grip wherever he can on the would-be truant. Then begins a -tug-of-war between the two bulls, during which the wretched victim of -their rage may be torn in pieces. By the elimination in each generation -of the more querulous and discontented, the peculiarly gentle and -passive nature so characteristic of the females has been developed. - -After the first ten days’ sojourn ashore the female is allowed to go -to sea to feed, returning presently to suckle her young. The bull, on -the other hand, can enjoy no such privilege. For three long months he -must keep watch and ward fasting—at first, in order that he may retain -his territory; later, that he may retain his harem. This fast, having -regard to the loss of energy and blood which this strenuous period -entails, is wonderful; for in the case of all other animals fasts -are always associated with absolute rest and sleep. Not so with the -Sea-lion; he arrives at the breeding-ground fat and well-liking, he -leaves a starved and battered wreck. - -The foregoing summary of the habits of these most interesting and much -persecuted animals is taken from the exhaustive report of Dr. F. A. -Lucas and Mr. Charles Townsend. These two distinguished naturalists -accompanied the United States contingent of the Fur-seal International -Commission despatched in 1896-97 to inquire into the threatened -extermination of these animals. Major Barrett Hamilton accompanied -the British contingent, and also made a report. And it is curious to -note that on some points he is diametrically opposed, not only to -the American naturalists, but to all other writers on this theme. He -contends, for example, that “nothing could better illustrate the fact -that it is the cows, and not the bulls, which have the real control of -the harem-system.” He traced the rapid growth of two harems from four -or five to as many as eighty cows. And he tells these were completely -out of control and free to move about as they wished. “The bulls, in -spite of all their bluster, had the flimsiest of nominal dominion, and -the cows were always able to, and frequently did, leave the harems -daily to dally with the cowless bulls on the outside. Yet ... as long -as they chose to sit massed together on the ground which had been -appropriated by the two stronger bulls, no weaker rivals could approach -to within ten yards of them. The master of the harem had no control -over its occupants, but he was absolute lord of the ground on which -they sat.” This is certainly curious, but more so is the fact that -these females were allowed to return by the “cowless bulls” outside -the charmed circle. Later in the season he tells us he witnessed an -even better illustration of this singular behaviour. At this time “the -division of the cows into harems was a very unequal one, the smaller -bull being only able to keep a very few cows, while the larger one -claimed the greater part of the rookery. But the cows could pass over -to the smaller bull’s ground as often as they liked; and he probably -was father to a great many more of the pups born in 1898 than those of -the half-dozen cows over whom he claimed control.” In regard to two -other bulls in another cart of the island, there came a time when the -inequality of the harems reached such a pitch, that the newly-arriving -cows “had to lie in scattered groups outside the main mass, and thus -permitted the weaker bulls to form new harems out of the reach of the -two strong old bulls.” But perhaps the most singular feature of all -was the indifference which one old bull displayed towards a little -bachelor, permitting him to enjoy the most intimate relations with one -of his cows without displaying the least sign of annoyance, as if he -could scarcely regard one so young as a rival. - -There is much evidence to show that the erotic side of the male-seal -develops early. “I saw,” he says, “the little black pups acting to each -other in a way that made it certain that their sexual feelings had -already made themselves felt.” This one can well understand, for only -animals of strong sexual tendencies could survive the strenuous life -which the period of sexual activity entails. - -The very different interpretation of the behaviour of these animals -at this very important stage of their life-history must be due to the -fact that different colonies were studied which were living, too, under -somewhat different conditions. It seems clear, for example, that the -landing of the females so graphically described by Dr. Lucas was a -landing under exceptional circumstances, the master bulls having taken -up positions at the only spot where access to the desired breeding -quarters was to be found; while Major Barrett Hamilton was probably -fortunate in seeing phases which were wanting in the “rookeries” -examined by Dr. Lucas. And both these observers again differ in the -accounts they give of the life of such “rookeries” with those by Mr. -Elliot, who explored these teeming colonies some years earlier when the -number of animals forgathered there was far larger and the fighting, -apparently in consequence, was far more severe. - -In the matter of secondary sexual characters the most remarkable of the -seal-tribe are those of the Elephant Seal and the Bladder-nosed Seal, -and this because of the extraordinary development of inflatable tissue -above the muzzle which these animals display. Of their life-history -we know little enough, and this despite the fact that for generations -the Elephant Seal was mercilessly hunted and slain for the sake of its -oil. Millions were slaughtered during the last century, yet only scraps -of information on the economy of the creatures has come down to us. -All that is of any value, and especially in regard to the “Courting” -period, we owe to Mr. Charles Townsend, of the New York Aquarium, and -this in regard to the northern species, _Macrorhinus angustirostris_ -of Guadelupe, though it may safely be inferred that the Southern, -Antarctic species, _M. leoninus_, differs in no essential respects. - -According to Mr. Townsend, the adult bull, having taken possession of -his territory and formed a harem, is constantly called upon to wage -duels for both with less fortunate rivals. And the severity of such -combats was attested by the deep wounds and festering sores of the -necks of these old warriors—which, at their maximum, attained in the -days of their prosperity a length of nearly thirty feet and a girth of -sixteen feet; but the last survivors of the race to-day seem rarely to -exceed twenty-two feet. The weapons used in fighting are the canines, -and the only armour they possess is that formed by the thickening of -the skin of the neck, which forms a great massive shield, so that -really dangerous wounds are rare. The great fleshy proboscis, the most -vulnerable part, is carefully guarded by the upturned position of the -head. The use of this trunk-like organ, which may attain a length -of about fifteen inches, is not clear; it seems to serve mainly as -an “ornament,” at times, too, furnishing a very definite indication -as to the temper of its owner. While the animal is slowly moving its -great carcase from place to place, this remarkable organ is relaxed, -and pendent; but when fighting it is closely contracted so as to be -out of harm’s way. Whether it plays any useful part in the capture of -food is not known; but it is probably much displayed during phases of -sexual excitement. In young animals, it is significant to notice, as -well as in the adult female this trunk is entirely wanting, which seems -to suggest that this peculiar feature has only been recently acquired, -the young and the adult female, as is the rule, standing nearer to the -early forebears of this strange type. There is an enormous difference, -it should be remarked, between the sexes in the matter of size, the -female not attaining more than half the bulk of her lord. A further -interesting point concerns the coloration of the young, which are -black, while the adults are brown. Doubtless this is connected with the -requirements of the young, the black coat attracting more heat than the -lighter-coloured coat of the adult. - -As touching that curious creature, the Crested, or Hooded Seal -(_Cystophora cristatus_), a native of the colder regions of the North -Atlantic. This animal is remarkable for the development, in the males -alone, of a great crest or casque on the head, which is formed by a -large inflatable air-sac over the ridge of the nose, and communicating -with the nostrils. When fully inflated, it covers the head as far back -as the eye. Its purpose is a matter of conjecture. It seems to be -inflated either when he animal is greatly excited, as when challenging -rival males, or when threatened with danger from other causes, as -when attacked by man. The males are exceedingly pugnacious, and fight -with one another for the possession of females with great ferocity, -such contests being accompanied by cries which can be heard for miles. -From the difficulty which Esquimaux and sealers find in killing the -animal with clubs it certainly seems as if this strange wind-bag were -more than merely ornamental. - -[Illustration: Plate 14. - -_Photo copyright, W. P. Pycraft._ - -ELEPHANT SEAL. - -This is a young animal. Note the great size of the eyes, and the -general “seal-like” character of the head as compared with that of the -adult.] - -[Illustration: _Photo by courtesy of Charles Haskins Townsend, Director -of New York Aquarium._ - -NORTHERN ELEPHANT SEAL. - -Adult male and female, and yearling. The male shows the enormously -inflated snout. - -[Face page 88.] - -That those extraordinary creatures the Cetacea—the Whales and their -kin—are derived from the same common stock as the typical carnivora -there can nowadays be no doubt, widely as they have departed from -their land-dwelling relatives in almost every possible feature of -their organization. In the matter of their “Courtship” we know -nothing, but we may infer certain incidents in this critical period -of their life-history from the peculiar nature of the secondary -sexual characters which some species display. Thus in the Pilot -Whale (_Globicephalus_) and the Bottle-nose Whale (_Hyperoödon_) the -forehead, in the bulls, is enormously swollen by a mass of fibrous -tissue so dense as to turn the blade of the sharpest knife, as I -know well from attempts to dissect this region. Now the only use, -surely, for such a cushion is that of a battering-ram by rival males -in charging one another, as rams and other horned animals will do. In -the Bottle-nose Whale this cushion is backed up by an enormous mass of -solid bone thrown up by the maxillæ. The origin of this bony growth -is interesting, for it appears first as a slight swelling in the rare -species _Berardius_; it is seen at a further stage of growth in the -female “Bottle-nose” (_Hyperoödon_), and attains its maximum in the -male, where it stands unique. There are two other species which demand -notice here. The first is Layard’s Beaked Whale (_Mesoplodon_); the -second the Narwhal. The former is the only vertebrate which in a wild -state wears a muzzle! In this species the teeth have totally vanished -save for a pair in the lower jaw, which are found towards the end of -the jaw. These in the adult, or perhaps we should say senile, male -grow upwards and inwards, finally meeting one another above the upper -jaw, so as to make it impossible for the animal to open its mouth more -than the fraction of an inch! Surely here we have a secondary sexual -character carried to an excess, and so proving not only disadvantageous -to the animal, but positively disastrous, for it seems clear that so -hampered the creature can feed only on the most minute forms of animal -life, which could only be captured and swallowed with difficulty. It -is true that the Rorquals feed on excessively minute Crustacea, but -they are able to take in enormous quantities at a time, the “whalebone” -serving the office of a sieve to prevent their escape. The Mesoplodon -has no such aids. One is tempted to believe that the skulls displaying -this most curious feature are abnormal, comparable to those, say, of -rabbits wherein the teeth have grown so excessively long as to close -the mouth, on account of the displacement of the cutting surfaces by -accident. But there is nothing to afford support to this view, and one -must therefore fall back on the suggestion of senility. - -The Narwhal has long been celebrated for the enormous size of the -canine teeth, the only teeth present in the jaws. As a rule, only -one leaves its bony socket, the other, commonly the right, remaining -as a mere vestige, seven or eight inches long within the skull. The -protruding tooth, which is spirally fluted, may attain a length of -nine feet. Occasionally both teeth are developed, and in this case -the spiral is the same, differing in a very striking manner from -the spiral horns of ruminants, wherein one presents a right, the -other left-handed spiral. But what purpose do these teeth serve? This -question has never yet been definitely settled. Some hold that it is -used to break open breathing holes in the ice, for the animal lives -in the far north: others that it is used as a spear in hunting prey. -Some aver that it serves as a weapon of offence, being used by rival -males in their struggle for mates. Scoresby, the explorer, indeed, says -he has seen young males in mock-battle, fencing with these remarkable -weapons. But until we have more satisfactory data, we must regard this -armature of the Narwhal as affording another instance of a secondary -sexual character of doubtful value to its possessor. - - - - -CHAPTER VI - -COURTSHIP AMONG BIRDS - - -Generalities—Darwin v. Wallace—The Peacock in his Pride—The “Display” -of the Peacock Pheasant—The Splendour of the Argus Pheasant and the -Marvel of its Eyes—The Frill of the Amherst Pheasant—Birds of Paradise -in the Toils of Love—Inflated Suitors—Ruffs and Reeves—Fearsome Weapons -and their Uses—Birds which dance—Musical Birds—The Bird’s Voice-box—The -“Lek” of the Capercaillie—Instruments of Percussion—The Curious -Performance of the Wood-pecker. - -The fact that so little is known about the mammals during that period -when the all-important work of securing mates is going on, and of the -subsequent events, is largely due to the difficulties which close -observation of this phase of their life-history entails. With the birds -matters are far otherwise; their haunts are more accessible; they -are far more numerous, and much more easily kept under observation. -Consequently, we have a tolerably complete knowledge of the lives -of some species, at any rate, during the reproductive period; that -is to say, as to the sequence of events from the beginning of the -reproductive activities onwards; but the interpretation of what is seen -is another matter. No attempt which has yet been made to fathom the -psychology of sex has yielded more than a slight insight into what is -taking place. Nevertheless, this is an aspect of the subject which -has a far more important bearing on the problems of evolution than -is generally realized. But these pages are concerned rather with the -relations between the sexes, than with the subtle forces which have -fashioned and control conduct in this regard. - -In all that concerns the problems of sex, which is to say of -reproduction, birds, speaking generally, display a briefer and more -condensed sequence of events than the mammals; and, moreover, many -species compel the attention even of the most incurious, to their -behaviour at this time, through the development, either of song, or -of fantastic displays of their amorous feelings: while others force -themselves no less conspicuously under notice by their habit of nesting -in large, and often enormous colonies. - -In the matter of the development of secondary sexual characters birds -stand conspicuous among the Vertebrates, and easily eclipse the -mammals; among which bright, strongly contrasted, colours are the -exception. Among the birds they may almost be said to be the rule. -Also, in this category we have to reckon song, and the production of -more or less musical sounds by the agency of internal resonators or of -specially modified feathers; as well as quaint forms of posturing which -may be included under the head of dances. Further, some species have -developed formidable weapons of offence. These things are interesting -enough in themselves, but they become still more so when we reflect -that they formed the corner-stone of Darwin’s theory of “Sexual -Selection,” and that Wallace’s criticisms thereof were inspired by -evidence from the same source. - -The interests of this chapter will best be served if the evidence -on which this theory was founded be first surveyed: when Darwin’s -deductions and the criticism which they have aroused will be the more -readily appreciated. - -Definitions are always liable to exceptions; and concrete cases are -better than abstract terms. Birds, then, perhaps better than any -other group, illustrate what is meant by the term “secondary sexual -characters,” if only because examples are so constantly at hand. Save -among experts, sex among birds cannot be determined except by the -differences in plumage, or sometimes in size, which the sexes display. -But even here, it is only among species which occupy what we may call a -mid-evolutionary phase in which this discrimination is possible. Among -“generalized” species, wherein the plumage is of sombre hue, there is -no external distinguishing mark between male and female; and the same -is true with species which have attained to the maximum of resplendent -plumage; as for example many Parrots and Kingfishers, where again both -sexes, and at all ages, display the same vivid hues. Thus, in the -case of either of the two extremes, the study of behaviour during the -breeding season is one of great difficulty and no less uncertainty. -Where the sexes are sharply distinguished by differences of coloration, -however, as with the Peacock, the matter is otherwise. This bird, from -time immemorial the symbol of vanity, illustrates in a singularly -effective manner the broad features of what is commonly meant by -“courtship” among birds, while it furnishes a no less striking example -of the development of “secondary sexual characters.” - -One might have supposed that birds, under the spell of that -irresistible desire for sexual intercourse, would behave differently in -regard to their “courtship” according to whether they were monogamous, -polygamous, or polyandrous: but while their behaviour during this -period of the life-history presents an extraordinary variety, it is -only at any rate slightly determined by the plurality or otherwise of -mates; and the same rule holds in regard to the brilliancy or otherwise -of coloration. - -The most common manifestation of sexual desire among birds takes the -form of strange posturings which are, in some species, enormously -exaggerated by the display of vividly coloured frills, tufts, or other -conspicuous modifications of the normal plumage. The Peacock affords -a most excellent example of this combination of the contortionist and -the beau, though the nature of this display is by no means generally -understood. This applies more particularly to artists, who from time -immemorial to the present day, in essaying to paint the Peacock in his -pride, have invariably fallen into the error of treating the great -ocellated train as if it were the tail, placing it where, of course, -the tail ought to be, at the end of the body! As a matter of fact it -is nothing of the kind; these gorgeous plumes are really exaggerated -tail-coverts which, when set on end, appear to arise from an oval -shield of metallic green scales—the central back-feathers. When this -trailing glory is erected, the bird throws the body forwards and -downwards, so that the outermost train-feathers fall downwards on -either side in front of the wings, which are more or less trailed: so -that from the front only the head and neck are visible, the rest of the -body being hidden _behind_ the screen, as may be seen by a reference to -the accompanying photographs. The manner of this display is extremely -interesting, for the bird seems to be conscious of the effect produced: -though it cannot be supposed that this is really the case. - -When displaying, the bird gradually approaches the nearest female -and slowly erects these extraordinary plumes. So soon as this is -accomplished he begins to walk backwards towards the object of his -attentions, presenting nothing but a great round shield of dull brown -feathers, backed up by the tail-feathers, and the dull-coloured wings. -So soon as he judges himself near enough, however, he suddenly swirls -round, confronting her in all his splendour, and heightening the effect -with a loud scream accompanied by a rapid, vibratory, motion of the -train-feathers which produces sounds like the pattering of rain on -leaves. Then he stands before her, with bowed head, as if to give her -an opportunity of drinking in his splendour to the full. Commonly, -however, she appears to be utterly indifferent, and either walks away -or continues a real, or affected hunt for food, as if no such thing as -a love-sick suitor were within a hundred miles of her! But sooner or -later his suggestive attitudes beget an answering response, and pairing -takes place. - -The display of the beautiful Peacock Pheasant differs conspicuously -from that of the Peacock, and recalls that of the Argus Pheasant. In -the Peacock Pheasant, as will be seen from the adjoining photograph, -the wings, and tail, are alike bedecked with ocelli. The display is -made by the bird as it crouches close to the ground, with the wings and -tail raised to form a continuous, patterned surface, the head being -swiftly moved during the performance; hence its blurred outline in the -photograph. - -[Illustration: Plate 15. - -_Photo by D. Seth-Smith._ - -PEACOCK PHEASANT. - -The display of this bird differs conspicuously from that of the Peacock -and recalls that of the Pigeon in some respects. The “ocelli” on the -wings afforded Darwin the interpretation he sought for as to the -meaning of the notch in the “eye” of the Peacock’s tail-feather.] - -[Illustration: Plate 16. - -_Photos by the Author._ - -“THE PEACOCK IN HIS PRIDE.” - -In the upper figure it will be noticed the “train,” when erected, -encircles the base of the neck; the lower figure shows the train -supported by the tail and dropping on each side in front of the wings. - -[Face page 96.] - -The Argus Pheasant is an even more wonderful performer than the two -preceding species. In this bird, it should be remarked, the tail and -the secondary wing-feathers are enormously lengthened, the latter to -an extent met with in no other bird, showing that the struggle for -existence cannot be very severe with this species. For if long -journeys had to be undertaken in search of food, or to avoid extremes -of climate, or enemies had to be swiftly escaped, such cumbersome -wings would lead to speedy extermination. But an even more remarkable -feature of these wings is their wonderful coloration. The primaries -have blue shafts, and a most delicately mottled pattern formed by spots -of reddish chocolate on cream-coloured ground, while the secondaries -have their broad webs ornamented with large ocelli, to be described in -greater detail presently. When under the influence of sexual excitement -Darwin tells us, the wings are so spread as to form a deep concavity, -an effect which is gained by pressing the primaries close to the -ground, and turning the elbows upwards. Within this concavity lie the -ocelli, in radiating vertical rows. But to produce this effect the bird -has to turn its head under its wing, so that it lies behind the screen. -Hence it cannot see the female which is the object of these captivating -antics. As a consequence, to discover whether he has an audience for -she will often walk disdainfully away—he has constantly to thrust his -head through the curtain, and hence many of the feathers in this region -get much worn. - -By nature it would seem the Argus Pheasant is a very solitary bird, -though we must assume it is polygamous. As the breeding season -advances, however, the male proceeds to choose some open space in the -depths of the forest—which it never leaves—and therefrom to clear all -the dead leaves, and twigs, for a space of some six or eight yards -square, so that nothing but the bare earth remains, and thereafter -this area is kept scrupulously clean. Here, in solitary state, for a -short season he remains, calling at frequent intervals to advertise the -fact that an eligible male is in the neighbourhood desiring mates. A -dozen times in succession he will break the stillness of the forest -gloom with a loud, “How-how, how, how, how!” Sooner or later comes a -responsive, “How-owoo, how-owoo!” and in a short time, guided by the -sound, one or more females discover the object of their quest. But the -pairing desire has not yet reached its full intensity, and doubtless to -kindle this the display just described is enacted, and not once, but -a dozen times probably, before the desired state of frenzy has been -aroused. Not seldom another male answers the cry, and this inevitably -leads to a duel whereby the fittest and strongest male is speedily -discovered. - -A word as to these ocelli. This pattern is rare among birds, and Darwin -brought to light some extremely interesting facts regarding it. He was -led to investigate the matter by his curiosity as to the meaning of -the notch in the ocelli of the Peacock’s train-feathers. At last he -noticed that among the different species of Peacock Pheasants there was -one (_Polyplectron chinquis_), in which the ocelli were paired, one -lying on either side of the shaft, in another (_P. malaccense_) these -approached and partly fused with one another. Now, to get the indented -ocellus of the Peacock, we have only to imagine the fusion of two such -ocelli, whose long axes inclined obliquely to one another, to get the -“eye” of the Peacock with its indented lower edge; for such fusion -would give a continuous upper and an indented lower border. - -The “eyes” of the Argus Pheasant are more interesting still, for, as -Darwin pointed out, these have the appearance, if the feathers are -held more or less vertically, of a number of balls lying each within -a socket, or cup: for each of these balls has a light area which -exactly simulates the light glancing across the upper pole of a sphere, -leaving the rest in shadow; and, singularly enough, this effect is -produced in the living bird only when the feathers are erected for -display. The probable steps in the evolution of these ocelli from -simple spots, and through elliptical bars, Darwin traced with his usual -skill and insight, and those who would follow this up should turn to -that wonderful book, “The Descent of Man.” - -[Illustration: Plate 17. - -PATTERNS WHICH PUZZLED DARWIN. - -The notch in the “eyes” of the Peacock’s train-feathers puzzled Darwin -till he met with the ocelli of the Peacock-pheasant. The left-hand -lower figure represents the ocellus of the Argus, the right-hand that -of the Peacock-pheasant. - -[Face page 98.] - -It is probable that the erroneous interpretation of the display of the -Peacock is due to the more lasting and easily remembered impression -of what obtains in the case of the Turkey under like emotions. This -bird in his exultant moods, most people have seen. Herein the tail -plays a very important part, being raised and spread to form a great -half-circle, while at the same time the back-feathers, or at least -those of the lower back, are set on end, and the wings are trailed on -the ground. The effect is heightened by the suffusion of blood to the -bare skin of the head and neck, and the sudden inflation of a long, -pendent, fleshy wattle from the forehead, which hangs down over the -beak. Great display is made with this, and an additional importance is -added by the spasmodic vocal efforts which can best be described by -the “gobble” rapidly repeated, as the bird struts about with mincing -gait, turning the wheel-like tail now to one side now to the other. -But the Turkey possesses yet another “ornament” which commonly escapes -notice. This is the curious tuft of long, black, coarse, bristles which -projects forward in front of the breast. It is difficult to discern -what part this tuft may play, since it is quite inconspicuous. It seems -as though this must be added to the number of structural characters -which appear to survive without any apparent use. - -The game-birds, it is significant to remark—and significant because -they are commonly polygamous—afford a quite remarkable series of -displays, only some of which can be summarized in these pages. In every -case, too, they are accompanied by conspicuous coloration and a more -or less excessive development of brightly-coloured plumes, or areas -of bare skin. In some, as in those wonderful birds the Tragopans, the -development of bare skin, vividly coloured, and produced into pendulous -folds, has attained a degree met with nowhere else among this group. -These flaps, or finger-like wattles, as the case may be, under the -influence of sexual excitement become turgid, and their hues enormously -intensified: though beyond this fact but little else is known of their -performances. In Swinhoe’s Pheasant the face is bare, the skin being -covered, as in the case of the common Pheasant, with tiny villi of a -vivid red colour. But when excited by the presence of a female the -upper part of this face area rises high above the head like a pair of -horns. With these turgid, and erect, the bird makes a series of short, -semicircular rushes around his prospective mate, accompanying each of -these gyrations with an angry hissing sound. The Golden, and Amherst -Pheasants are among the most gorgeously clad of birds. Not their least -conspicuous ornament is a cape-like frill of long, highly coloured -feathers of which the birds seem to be extremely conscious; for when -endeavouring to excite the female nearest him to the necessary pitch of -sexual desire, he places himself sideways before her, drawing the frill -round to the side facing her, and dropping the wing, in order, as it -would seem, that she may miss nothing of his resplendent livery. This -side of his nature he reserves for her. Intruding rivals are treated -after quite another fashion, for like most of the gallinaceous birds -his legs are armed with formidable spurs which can, and do, inflict the -most terrible wounds: as, indeed, has been shown from the evidence of -the Cock-pit in the case of game-cocks. - -[Illustration: Plate 18. - -THE “STRUTTING TURKEY.” - -This should be contrasted with the Peacock. Herein the tail itself -is the principal ornament, the effect of which is heightened by the -erection of the back-feathers, and the vivid play of colour of the -“wattles” of the head.] - -[Illustration: _Photo copyright by W. H. Quentin._ - -THE DISPLAY OF THE GREAT BUSTARD. - -This is effected by the inflation of a great wind-bag in the neck, and -the eversion of the wing and tail feathers as described in the text. - -[Face page 100.] - -By way of contrast with the several displays just described, it would -be hard to find a more striking illustration than that afforded by -the Lesser Bird of Paradise (_Paradisea minor_), inasmuch as here the -display is associated with rivalry between a number of individuals. -For much of our knowledge on this subject we have to depend on the -descriptions of natives; but happily this has now been supplemented by -observations made by Mr. Ogilvie Grant on a captive in the Gardens of -the Zoological Society of London. - -Impelled by the surging wave of sexual desire, as yet only seeking -consummation, these birds gather together at frequent intervals, on -certain of the forest trees of the Aru Islands, selected apparently -because they present an immense head of spreading branches, and large -but scattered leaves. Here ample space is found for the revels, which -take the form of “Sacaleli,” or dancing-parties, comparable to the -erotic dances of many barbaric races. - -By the time the ball opens, the birds, to the number of twenty or more, -have worked themselves up into a state bordering on frenzy, and each -commences his performance with quivering wings and loud, penetrating -cries which may be syllabled as _walk—walk—walk—walk—walk—walk_, -rapidly repeated. Then the wings are suddenly held out on either side, -the tail is bent forward under the branch, and with a quick, barely -perceptible rustle, the gorgeous, golden, diaphanous side-plumes are -thrust upward and forward on each side of the body, forming an arched -cascade above the back. With every muscle tense the performer will -remain in this attitude from ten to twenty seconds, slightly quivering -the wings, and from time to time imparting a tremor to the upraised -plumes. Then follows a second phase. Each bird, seemingly possessed, -commences to dance and hop wildly backwards and forwards along the -bough, and with head bent forward, wings spread horizontally, and the -side plumes raised to their utmost, he gives vent to a series of loud -harsh cries—“ca! ca! ca! ca!” For some seconds he remains in a sort -of ecstasy, rubbing his beak on the bough, and occasionally glancing -backwards below his feet, and with the back fully arched. The climax -passed, he reverts once more to the earlier, more erect stage of the -display, when the paroxysm either gradually subsides or is renewed. - -No less extraordinary is the behaviour of the King-bird of Paradise -(_Cicinnurus regius_), which has been described by Sir William Ingram, -who for a time had a captive in his aviaries. As the illustration -shows, its posturing is quite remarkable. Before this is described, -however, a brief description of its coloration should be given, -which, it must be remarked, cannot possibly convey more than a very -vague idea of its sumptuous character. Picture a bird no bigger than -a thrush, but of a wonderful cinnabar red, with a gloss as of spun -glass: the head clothed in short, velvety, orange-hued feathers; and -with a white breast, having the softness and sheen of satin, and -crossed by a band of deep metallic green, contrasting with the red -of the throat. Add a yellow beak, and legs of cobalt blue, and you -will have the features which catch the eye at the first glance. But a -little closer examination will reveal yet other points for wonderment. -Along each side of the body the upper flank-feathers become elongated -and delicately tinted, and, furthermore, they are erectile: so that -they can be raised up on each side of the body to form an almost -circular shield of delicate ash grey, bordered with buff and emerald -green. These play a most important part during the sexual frenzy, -and the effect thereof is not a little heightened by the middle pair -of tail-feathers, which have been modified to form a pair of slender -stalks, some ten inches long, bearing at the ends a curious disc of -emerald green formed by coiling upon itself—like a watch-spring—the -only piece of the vane of the feather which remains. - -So much for its fine feathers; now for the manner of their use. “He -always commences his display,” writes Sir William Ingram, “by giving -forth several short notes and squeaks, sometimes resembling the call -of a quail, sometimes the whine of a pet dog. Next he spreads out -his wings, occasionally quite hiding his head; at times, stretched -upright, he flaps them, as if he intended to take flight, and then, -with a sudden movement, gives himself a half turn, so that he faces the -spectators, puffing out his silky-white lower feathers; now he bursts -into his beautiful melodious warbling song, so enchanting to hear but -so difficult to describe. Some weeks ago I was crossing a meadow and -heard the song of a skylark high up in the heavens, and I exclaimed at -once: ‘That is the love-chant of my King-bird.’ He sings a low bubbling -note, displaying all the while his beautiful fan-like side-plumes, -which he opens and closes in time with the variations of his song. -These fan-plumes can only be expanded when his wings are closed, and -during this part of the display he closes his wings and spreads out -his short tail, pressing it close over his back, so as to throw the -long tail-wires over his head, while he gently swings his body from -side to side. The spiral tips of the wires look like small balls of -burnished green metal, and the swaying movement gives them the effect -of being slowly tossed from one side to the other, so that I have -named this part of the display the ‘Juggling.’ The swaying of the body -seems to keep time with the song, and at intervals, with a swallowing -movement of his throat, the bird raises and lowers his head. Then comes -the finale, which lasts only a few seconds. He suddenly turns right -round and shows his back, the white fluffy feathers under the tail -bristling in his excitement; he bends down on the perch in the attitude -of a fighting cock, his widely-opened bill showing distinctly the -extraordinary light apple-green colour of the inside of the mouth, and -sings the same gurgling notes without once closing his bill, and with a -slow dying-away movement of his tail and body. A single drawn-out note -is then uttered, the tail and wires are lowered, and the dance and song -are over. - -“The King-bird has another form of display which he very rarely -exhibits, and only on three or four occasions have I seen him go -through this performance. Dropping under the perch, the bird walks -backwards and forwards in an inverted position with his wings expanded. -Suddenly he closes his wings and lets his body fall straight downwards, -looking exactly like a crimson pear, his blue legs being stretched -out to the full length and his feet clinging to the perch. The effect -is very curious and weird, and the performance is so like that of -an acrobat suddenly dropping on to his toes on the cross-bar of a -trapeze that I have named this the ‘Acrobatic’ display. It has been -witnessed on different days to his ‘Juggling’ display. While giving his -‘Acrobatic’ performance he sings the whole time, but never shows his -side-plumes, and when he is in the pendulous position his body sways -gently as if it were influenced by a fitful breeze. The whole of this -performance takes but a very few seconds.” - -[Illustration: Plate 19. - -_From a Drawing by Roland Green, Jun., adapted from G. E. Lodge and -others._ - -SOME OF FORTUNE’S FAVOURITES. - -The Birds-of-Paradise have few rivals in the matter of ornament. In the -centre of this plate are seen the Lesser and the King Bird-of-Paradise -displaying (after G. E. Lodge). The first-named is distinguished by -the enormous development of the side plumes, which can be raised high -above the back. In the second, the ornaments take the form of erectile -frills on each side of the breast, and strangely modified tail-feathers -which end in curious discs. At the top left-hand corner is the -King of Saxony’s Bird-of-Paradise; on the right is the Long-tailed -Bird-of-Paradise; at the bottom of the page, from left to right, are -Hunstein’s, the Six-wired, and Superb Bird-of-Paradise. - -Face page 104.] - -Naturally one needs to witness such a display to appreciate its beauty -and its weirdness; but the wonderful sketches which my friend, Mr. G. -E. Lodge, made during one of these performances, should go far towards -helping the reader to visualize what really takes place. - -While it would be untrue to say that the Birds of Paradise are of a -more amorous, or more excitable disposition than other less resplendent -birds, one cannot but be impressed with the fact that they exhibit -a range of variation in the matter of feather-ornament probably -unequalled, and certainly unsurpassed, by any other group of birds. -From what has been observed of the few species which have been kept -in confinement, they seem to enjoy no less distinction in matters of -display. On this latter subject no more of importance can be said, and -exigencies of space forbid any attempt to describe the exquisite beauty -of coloration which a survey of all the known species reveals. It would -be hardly more profitable to attempt to describe the varied character -of the shields, crests, frills, streamers, which are to be met with -in different species: but a glance at the accompanying illustrations -will show that it would be hard, indeed, to exaggerate the splendour -of the ornamentation which these birds have developed. Even here, -where no indication can be given of the glowing, vivid colours, often -indescribably beautiful, it is obvious that these birds well deserve -their name. St. John’s imaginary Paradise would probably have been -described in far more enticing language had he known of the existence -of these wonderful birds. - -Among all the known species the dullest is Wallace’s Bird of Paradise, -the general coloration being of a dull brown hue; but even here, a pair -of wing-coverts are produced into long, broad streamers, unique among -birds; while the feathers of the throat and flanks are of a marvellous -metallic green, the flank-feathers being produced to form a long, -pointed tuft. - -To what factors must we attribute the growth of these wonderful -colours, these strange outgrowths, frills, and tufts, and streamers, -the like of which is almost unparalleled? In a group numbering some -fifty or more species there is not one that does not display some -strange feature. We cannot attribute it to the environment, for in such -case the results should have produced uniformity; nor can we invoke -the aid of sexual selection save in a very indirect manner, and in a -sense other than generally understood by this term. It seems, then, -not unreasonable to suggest that they are the expression points of the -internal metabolism: the manifestations of that tendency to vary which -is inherent in every fibre of the organism. But no attempt shall be -made to elaborate this theory till more evidence has been taken. The -humming-birds, and the game-birds, are perhaps the only other groups -which exhibit quite such a prodigality of ornament; of the latter, -instances have already been cited. - -So far the displays which have been described have been such as -are confined to the use of more or less resplendent plumage. There -are, however, many species which contrive to secure most startling -results, not so much by the parade of coats of many colours as by -grotesque changes of shape produced by wind-bags of various kinds. The -Pouter-pigeon affords a case in point. This bird possesses the power of -inflating the gullet to an enormous size, so as to produce a strangely -distorted form, at any rate, to our eyes. The “Pouter,” it is hardly -necessary to mention, is an artificial product of the “fancier,” who -has taken advantage of the natural tendency, seen in the Wild Pigeon, -to inflate the neck during moments of excitement. By the selection from -each generation of the finest performers in his stock, the Pouter of -to-day has been developed. But there are many birds which, while not -even remotely related, have developed the same strange device. The most -striking illustration of this kind is furnished by the Great Bustard, a -bird once common on the fen-lands of Great Britain, but now, unhappily, -exterminated within these islands. - -The means of inflation in this case is afforded by a large thin-walled -sac of a very remarkable character. Opening by a small slit just under -the tongue, it is continued down the front of the neck immediately -under the skin, which in this region is thickened by an accumulation -of fat and blood-vessels. Between the arms of the furcula, or -merry-thought, its cavity is constricted, to expand again immediately -to form a pear-shaped termination. How it is filled is something of -a mystery. But once inflated, the bird draws its neck downwards and -backwards, so that the head is brought to rest between the shoulders -and is there almost buried, partly by pressure on this curious -air-cushion and partly by the erection of a number of bristle-like -feathers, which in calmer moments project backwards on each side of -the head. At the same time the tail is drawn forwards to lie upon the -back, thus exposing a billowy mass of white feathers forming the under -tail-coverts. The tips of the wing-feathers are used to hold the tail -in position. Meanwhile the scapulars are set on end, and the long inner -secondary quills are similarly erected. The feathered contortionist, -having completed his preparations, now approaches his partner with a -mincing gait, then halts before her and solemnly utters a series of -low grunts like “oak, oak, oak.” Having thereby declared his passion, -and commonly without gaining any answering response, he returns to his -normal shape again! - -It is curious that a near relative of this bird, the Great Australian -Bustard (_Eupodotis australis_), also captures the wind to declare -his love; but it is disposed of after a quite different fashion, -being drawn into the gullet, though the precise manner in which it -is manipulated demands further investigation. The display Dr. Murie -described many years ago. It begins, he tells us, with a swelling -of the throat, while the head is thrown upwards. Immediately after, -the neck swells, and the feathers of the lower part of the neck, set -all on end, are carried downwards, apparently surrounding a huge bag -which reaches nearly to the ground. During all this time the head and -neck are held rigid and point skywards, the head surmounting a great -feathery column. Meanwhile the tail, as with the Great Bustard, is -drawn forwards over the back. In this peculiar attitude the bird struts -about in a stiff, waddling manner, the elongated neck-bag swaying to -and fro and the feathers of the throat standing out in the shape of a -great rounded swelling. The acme of inspiratory effort completed, the -bird begins to snap the jaws together, producing loud noises, which are -accompanied by a soft dove-like cooing. - -The Pectoral Sandpiper in like manner inflates its gullet. But, unlike -the Pigeon and the bird just described, the neck is not markedly -straightened, nor is the body raised. As the air is drawn in, the -gullet expands, till it forms a great spherical drum. Then the excited -performer runs along the ground uttering a resonant “too-u tooo-u” -repeated seven or eight times in rapid succession, all the while he -approaches nearer to the apparently very much-otherwise-engaged female. -This effort failing, he will then often rise on quivering wings twenty -or thirty feet into the air, and dive gracefully down again immediately -afterwards, deflating this curious balloon to await a more favourable -opportunity. - -In some species where wind-bags are employed as aphrodisiacs the outer -skin is brilliantly coloured and exposed during the display. The -Prairie-hen affords a case in point. In this species the air-chamber -is furnished, not by the gullet, but by the air-sacs of the neck. When -these are inflated they appear as two large orange-coloured bodies -standing boldly out among the feathers. Their effect is heightened by -a tuft of long stiff feathers which are thrust forward like a pair of -horns, on each side of the head, while at the same time the feathers of -the back are set on end, the tail is spread like a great fan, and the -wings are half opened and trailed like those of the Turkey. - -The displays take place in the early hours of the morning, when parties -of from a dozen to fifty, of both sexes, meet on some slight knolls -where the grass is short. Having duly assembled, the more ardent cocks -immediately begin to prepare for the morning revels, the first part of -the performance apparently being of a comparatively passive nature—the -parade of the air-sacs and the erection of the feathers. - -Then some “proud cock, in order to complete his triumph, will rush -forward at his best speed ... through the midst of the love-sick -damsels, pouring out as he goes a booming noise ... which may be -heard for at least two miles in the morning air. This sound is by no -means harsh or unpleasant. When standing in the open prairie at early -dawn listening to hundreds of different voices pitched in different -keys, coming from every direction and from various distances, the -listener is rather soothed than excited. - -“Every few minutes this display is repeated. I have seen not only one, -but more than twenty cocks going through this funny operation at once; -but then they seem careful not to run against each other, for they have -not yet got to the fighting point. After a little while the lady birds -begin to show an interest in the proceedings by moving about quickly, a -few yards at a time, and then standing still a short time. - -“The party breaks up when the sun is half an hour high, to be repeated -the next morning, and every morning for a week or two before all make -satisfactory matches. It is towards the latter part of the love-season -that the fighting takes place among the cocks, probably by two who have -fallen in love with the same sweetheart....” - -There is much that is extremely interesting in this account and a -little that seems to have been misinterpreted. The fact that these -antics are repeated during many days until at last the females are -moved to display some interest is just what we should expect if this -demonstrative behaviour on the part of the males acts, as we believe, -as an aphrodisiac. And that actual fighting occurs is highly probable, -but there can be no doubt that in such case the whole aspect of the -bird must be changed, for anything in the nature of fighting with the -delicate air-sacs inflated would greatly endanger the most important -aid to success in achieving this object which these birds possess. - -[Illustration: Plate 20. - -_Photo from The Museum of Natural History, New York._ - -THE LOVE-MAKING OF THE PRAIRIE HEN. - -During the “display” large, yellow, air-sacs in the neck are inflated. -The bird in the foreground shows one of these, and the ornamental -feather frill, very clearly. - -[Face page 110.] - -No less remarkable is the performance of the Frigate-bird (_Fregata_), -a tropical species allied to the Pelicans and Boatswain-bird, and to -our own more familiar Cormorants and Gannets. It might well be called -a marine Swift, having excessively short legs and small feet, and a -wonderful expanse of wing. As with the Swifts, of course most of its -time is spent on the wing; the feet are only useful for supporting the -body when ashore, they are never used for walking, at any rate, for -more than a few steps. The wings afford the only means of locomotion. -Our knowledge of these birds when under the stress of sexual excitement -we owe to Dr. C. W. Andrews, who had the good fortune to study the -species known as the Great Frigate-bird (_Fregata aquila_) during his -task of surveying Christmas Island (Indian Ocean). - -“About the beginning of January,” he remarks, “the adult males begin -to acquire a remarkable pouch of scarlet skin beneath the throat; this -they can inflate till it is nearly as large as the rest of the body, -and a dozen or more of these birds sitting on a tree with outspread, -drooping wings and this great scarlet bladder under their heads are a -most remarkable sight. When a hen bird approaches the tree the males -utter a peculiar cry, a sort of ‘wow-wow-wow-wow,’ and clatter their -beaks like castanets, at the same time shaking their wings. When -they take to flight the air is allowed to escape from the pouch, but -occasionally they might be seen flying with it partly inflated.” - -Here again there can be no doubt about the purpose, or perhaps one -should say the stimulus, of this strange performance. This pouch, -I have been enabled to ascertain from dissection, is not formed by -inflating the gullet, but, as in the case of the Prairie-hen, by the -enlargement of the air-sacs of the neck. - -These air-sacs, which are present in all birds, are only enlarged to -further the ends of sexual display in a few species, and, curiously -enough, these are in no way related one to another. The Adjutant -storks, it may be remarked in this connection, have used the air-sacs -which are fed by the nasal system instead of those fed by the lungs, -as in all the species so far described. When deflated this pouch forms -a quite inconspicuous conical swelling in front of the neck; under the -stimulus of excitement, it awakens as it were into activity, and is -suddenly transformed into a great red or red-and-black bag, encircling -the neck and projecting far downwards in front of it, only to be -deflated an instant later with a speed which leaves one gasping. - -The specialization of the air-sacs, that is to say their transformation -to perform new functions subservient to the ends of sexual activities, -is not exclusively confined to display. In at least one instance an -air-sac has been specially developed to act as a voice resonator. -This is furnished by the Emu, wherein the wind-pipe, near the middle -of its length and on its anterior aspect, has a number of incomplete -rings forming a long slit. The lining of the windpipe escapes from -this slit in a hernia-like pouch, and takes up a position beneath the -skin. Even when inflated this pouch gives no very obvious sign of its -existence, but it serves to produce a curious hollow, drumming sound, -like the boom of a big drum softly beaten. But why it should have been -developed, when the Ostrich and the Cassowary produce similar but -louder “music” without any special apparatus whatever, is a mystery. At -least one species of Cassowary can emit a roar which would do credit -to a lion. - -In the males of all healthy animals the periodic stimulus to -reproduction finds expression in more or less striking eccentricity -of conduct. Sometimes, as the foregoing instances have shown, this -has been exaggerated by the development of long, resplendent plumes: -sometimes by brilliant coloration, displayed either by the plumage or -by bare areas of skin, or by both, while in not a few cases attitudes, -to our eyes grotesque and made still more so by the aid of inflatable -pouches, are the outward and visible sign of the raging fires within. -For the completion of this chapter yet other instances of this kind -must be cited, instances which reveal a further elaboration of some -of the more striking of these tricks of posturing; or which concern -the growth of the aggressive instincts, which are proclaimed by the -development of armature often of a very formidable character. As the -sequel shows, however, there are no hard and fast dividing lines -between these several modes of expression. - -That remarkable bird, the Ruff (_Machetes pugnax_), now, alas! no -longer to be met with in our fens, exhibits a curiously composite -character in the phases of its love display. - -Preparations for this are begun in the early spring by the assumption -of what is called a “nuptial dress,” which is worn only by the male, -and which contrasts in a very conspicuous manner with the plumage worn -during the rest of the year. The most striking features of this dress -are the great, erectile, Elizabethan ruff which encircles the neck -immediately behind the head, and the long, tongue-shaped “ears” which -surmount the head itself. These exhibit a most remarkable diversity -in their coloration, and it is no exaggeration to say that no two are -ever alike. Red, cream-colour, buff, black, white; spotted, streaked, -freckled and barred are the only descriptions that can be applied to -them, for the combinations of their hues and patterns seem infinite. -Having grasped this fact, the eye next turns to the colouring of the -rest of the body, and it will be found that here too is the same -diversity, though less conspicuously so; and finally it will be noticed -that at this time the feathers around the base of the beak have been -replaced by yellow or orange-coloured papillæ. The females also now -wear a dress differing from that of the so-called “winter plumage,” -but it does not present any very striking features nor any form of -ornamentation comparable to that of the males. - -The Ruff is a polygamous bird, which, in its display, presents some -curious and puzzling features, one of which consists of a sort of -tournament between rival males. At the break of day the performers, -selecting such eminences as the fen-lands afford, assemble apparently -to display their finery, for a couple of males will often stand facing -one another with frills erected and beaks touching the ground, silent -and immovable, for perhaps half a minute. Sooner or later, however, -they will commence to spar, and this presently leads to blows, during -which one of the combatants will attempt to seize the other by the -wings. However, no damage seems to be inflicted during such encounters, -which are by no means aimless or profitless, for during such bouts the -weaker, less vigorous birds are driven from the field, and the victor -in consequence wins for himself a larger harem. - -When the actual pairing time arrives the parade of the frills begins -again. The amorous instincts, it is important to notice, are awakened -earlier in the males, so that by the time the females have attained to -a like condition the least mettlesome males have been driven off. What -follows is not the selection by the females of the finest performers -so much as a process of sorting out, whereby the females discover and -cleave to those males which are readiest for mating. This display -succeeds in revealing both the most mettlesome males and the most -amorous females, who, however, would seem to require great persistence -and much demonstration on the part of the males before they can be -finally aroused to the pitch necessary for pairing. Again and again a -male may be seen to approach an apparently very unconcerned female, -and then to crouch down before her with his beak pressed to the ground -and his frill and “ears” set off to their fullest. For some seconds -he will remain lost in apparent contemplation, then with a dazed, -far-off, expression he will look up, to find, as often as not, that -she is still apparently feeding, quite unmoved by his protestations; -or that she has even flown off and left him. Pursuit speedily follows, -and the performance is repeated until at last she too catches the flame -of passion and permits, or rather invites, the final act of sexual -congress. - -Though these birds on occasion will fight, and savagely, they cannot -inflict serious damage on one another by reason of the relative -feebleness of their beaks and legs, which are but ill-adapted for -violent measures. Inasmuch as the Ruff is a polygamous species, -these bloodless battles have a peculiar interest. They show that -the preponderance of females, which polygamy implies, is not, as is -commonly supposed to be the case, due to a high death-rate among the -males by fighting. The same is true of the Wydah-birds, and their kin, -the only polygamous species among the Passeres. - -In this connection it is to be remarked that fighting, of a more or -less sanguinary character, is apparently universal among birds, the -conflicts being waged not so much in the way of squabbles for the -possession of females as for the acquisition and retention of territory -and all that this entails during the breeding season and, to a much -less extent, in the defence of the eggs and young. But to this point -we must return. For the moment it will be more profitable to focus -attention on the character of this fighting. In the first place, it -is by no means necessary that the combatants should be armed. The -“dove of peace” at this time of the year appears in a new and not -always pleasing light, for not only will he fight his neighbours, -but he does not always show that gentleness towards his wife with -which tradition has credited him. The little Humming-bird would seem -to be as little capable of fighting as a bird could be, yet few are -more pugnacious. The naturalist Gosse tells of a pair which had torn -one another’s tongues out in their blind fury; and everybody knows -that Robins and Tits fight savagely to preserve their chosen haunts -from invasion by their neighbours. In some birds this pugnacity has -become an overmastering passion. Some of the Quails, and a species -of Rail (_Gallicrex cristatus_), a near relation of the Moorhen, are -commonly kept by the natives of the East, as our forefathers kept -Fighting-cocks, for the sake of seeing them fight one another. Yet, -save in the case of the Fighting-cock, neither of these birds possesses -any aggressive weapons. - -Among the game-birds, however, powerful armature, in the shape of -long, pointed, spurs on the legs are met with. In the Jungle-fowls -and Pheasants only a single pair are found on each leg, but in other -species, as in the Francolins, there are several pairs, and these -birds, it is instructive to notice, are notorious for the ferocity -of their encounters. It is said that in the Indian Swamp-Francolin -(_Francolinus gularis_) nearly every individual is marked by scars and -wounds received in duels with rivals. - -Certain members of the Plover-tribe, and certain Anserine birds, have -developed spurs of a very formidable character on the wings. Among the -Plover-tribe the best example of such armoured species is the Egyptian -Spur-winged Plover (_Hoplopterus_). This bird, after the fashion -of its unarmoured relatives, such as the Common Lapwing, fights by -turning suddenly in the air and striking with the wings. In the case -of the formidably armed Egyptian bird the result is often fatal; but -with our Lapwing a fatal result is rare, since but slightly swollen -knobs take the place of spurs. In Hoplopterus and in the Jacana this -spur arises from the base of the thumb, but in the Spur-winged Goose -(_Plectropterus_) it is borne by one of the wrist bones (the radial) -while in the aberrant Geese-like birds (_Palamedea_ and _Chauna_) there -are two spurs on each wring, one at each end of the metacarpus. That -these weapons have come into being in response to need seems a very -natural conclusion, but it is one which presents many difficulties -when more closely examined. The wing spurs, differing widely in their -nature as they do, in one case borne on a carpal bone, in others on the -metacarpus, seem rather to owe their origin to fortuitous variations -which have become, so to speak, adopted by selection, than to a -response to the oft-repeated stimuli incidental to fighting. The latter -explanation is Lamarckian and to-day finds favour with but few. The -stimulus theory seems to be effectually discounted by the existence -of the spurs on the legs of gallinaceous birds. That these owe their -origin to impacts, or blows, seems more than doubtful: and one can -hardly see how they could have served any useful purpose until they had -attained a sufficient length to serve as weapons. Even if we suppose -that the spurs of, say, the Jungle-fowl or the Francolin have been -derived from tuberosities such as are found on the legs of the French -Partridge (_Caccabis rufa_), we should still lack evidence that the use -of the legs in fighting caused the origin of the tuberosities. - -There is yet another puzzling feature in regard to the armature of -the wings, and one which may yet help to a better understanding of -the puzzles presented by spurs. A Jacana, one of the Plovers, has the -radius broadened or flattened out from its middle onwards to form a -flat plate or blade, but the use thereof is unknown. It may possibly -serve as a weapon of offence, enabling the bird to beat its rivals with -its wings, but from the nature of the structure, and of the effect -such a use of the forearm would have upon the hand, it seems doubtful -whether it serves any aggressive function. If used at all in fighting -it is probably during fights in mid-air, when, after the fashion -pursued by the Spur-winged Plover, and even in the case of our own -Lapwing, a blow is struck by the uppermost bird at its rival, and often -with fatal effect. It is significant to remark, by the way, that in -the Lapwing a tubercle answers to the spur of Hoplopterus just as the -tubercles of the French Partridge (_Caccabis_) answer to the spur of -the Jungle-fowl or Pheasant: but the flattened radius of the wing of -the _Metopidius jacana_ has no parallel. - -With birds, as with men, there must always remain the ability to -appeal to force when some important end cannot otherwise be gained. -The species which adopts the crazy tactics of the Quaker is doomed to -extinction, sooner or later. The foregoing instances display force, as -we may say, aggressively. But even the peacefully disposed birds can -fight when aroused. - -Reference has already been made to dancing in this chapter; but so far -no very striking instances thereof as a form of sexual display have -been cited. The subject has been deferred because this peculiar type of -activity is not always directly associated with the _furor amantium_. - -With some species, which, it should be remarked, also lack distinctive -colouring, the erotic state is manifested apparently not so much by the -display of expanded wings and tail as by frenzied dances. The Jacanas, -aberrant members of the Plover tribe resident in South America, are -expert performers, displaying moreover a curious spontaneity during -such outbursts. A flock will be apparently sedulously feeding when -suddenly and with quick, excited gestures all will cluster together in -a group and go through a singular and pretty performance, holding their -wings outstretched and agitated, some with a fluttering and others with -more leisurely movement, like that of a butterfly sunning itself. The -performance over, all scatter and feed again.. The Honourable Walter -Rothschild, in his “Avi-fauna of Laysan” tells us of the stately -Albatross, which breeds, or rather bred there—for the Japanese display -a singular callousness in regard to animal life where commercial -interests are concerned in thousands: “First they stand face to face, -then they begin nodding and bowing vigorously, then rub their bills -together with a whistling cry. After this they begin shaking their -heads and snapping their bills with marvellous rapidity, occasionally -lifting one wing, straightening themselves out and blowing out their -breasts; then they put their bills under the wing or toss them in the -air with a groaning scream, and walk round each other often for fifteen -minutes at a time.” - -Cranes are much given to dancing. Mr. Nelson, an American -ornithologist, has described with much vigour the dancing of the -Sandhill Crane in Alaska. As he lay in a “hunting-blind” he was -suddenly aroused by the arrival of a crane, followed speedily by -a second, uttering his loud note as he came, until he espied the -first-comer on the ground, when he made a circuit and dropped close -by. Both birds then joined in a series of loud rolling cries in quick -succession. Suddenly, the last-comer, which seemed to be a male, -wheeled his back towards the female and made a low bow, his head nearly -touching the ground, and ending by a quick leap into the air. Another -pirouette brought him facing his charmer, whom he greeted with a still -deeper bow, his wings trailing loosely by his sides. She replied by an -answering bow and hop, and then tried to outdo the other in a series -of spasmodic hops and starts, mixed with a set of comically grave and -ceremonious bows. The pair stood for some moments bowing right and -left, when the legs appeared to become envious of the large share -taken in the performance by the neck, and then would ensue a series -of skilled hops and skips, like the steps of a minuet. Such antics -are characteristic of the Cranes of all species, and sometimes a -whole flock will join in such dances. But, it is to be noted, they are -not necessarily signs of the _furor amantium_: they certainly always -accompany this, but frequently they are indulged in, apparently, solely -as an outlet for exuberance of feeling. - -Before the theme of dancing can be dismissed the performance of a small -species of perching bird, one of the South American Manakins, must be -described. The natives call it the “Bailador,” or dancer. In an account -of his travels in Nicaragua Mr. Nutting tells us: “I once witnessed one -of the most remarkable performances it was ever my lot to see. Upon a -bare twig ... at about four feet from the ground, two male ‘bailadors’ -were engaged in a song and dance act that simply astonished me. The two -birds were about a foot and half apart and were alternately jumping -about two feet in the air and alighting exactly on the spot whence they -jumped. The time was as regular as clockwork, one bird jumping up the -instant the other alighted, each bird accompanying himself to the tune -of to-le-do—to-le-do—to-le-do, sounding the syllable to as he crouched -to spring, le while in the air, and do as he alighted. This performance -was kept up without intermission for more than a minute, when the birds -suddenly discovered they had an audience and made off.” Here again we -have no evidence of the _furor amantium_; nor that any females were -spectators of the scene. - -It is important to notice that Mr. Howard, in the course of his study -of the Warblers, witnessed a performance having some likeness to this -on the part of three young Sedge Warblers but newly escaped from -the nursery. And this not in some solitary instance, but on several -occasions. Just after leaving the nest, he remarks, they are very -playful, “their games sometimes taking the form of a tilting match. -Three take part; two sit on convenient twigs facing one another, and -the third, from the central position, might almost be called an umpire. -Numbers One and Two lower their heads, each in anticipation of the -other moving; one of them, call him Number One, then springs into -the air and darts at Number Two: Number Two dodges and occupies the -position vacated by Number One; each of them then faces round ready to -continue the fray, the change of positions becoming quite rapid.” But -no recurrence of these antics has been noted during the course of the -adult sexual display, which is confined to posturing and displaying the -outspread wings and tail. Nevertheless there can be no doubt but that -such games in later life are incorporated, in the case of many species, -with the love display. - -That the reproductive glands have played, and still play, a by no means -unimportant rôle in Evolution is shown by the history of the secondary -sexual characters. Among the birds, at any rate, the early stages of -physical changes belonging to this “figuration” are to be seen in -various forms of posturing, which in their more elaborate developments -we call “dances.” In many cases, as for example among the Warblers, the -periods of sex-emotion are marked by posturing alone. But in a number -of species, as has already been shown, the products of the sexual gland -seem to have undergone some further elaboration which has resulted in -the additional phenomena of gaudy coloration, in hypertrophied plumes, -and in weapons of offence. - -But not yet is the list of such sexual products exhausted, for no -mention has so far been made of the development of the many wonderful -devices for the production of peculiar and arresting sounds, musical -and otherwise. These are of two kinds: one wherein certain feathers -have been modified to produce rhythmical notes either by percussion or -by vibration; the other wherein the internal organs have been modified -to produce musical notes or loud, resonant cries. - -Instances of the latter kind are innumerable, and as a consequence no -more than one or two can be cited in these pages. The facts associated -with the production of vocal, as distinct from instrumental, music are -both curious and puzzling. To begin with, this music is produced by -the lower end of the trachea or windpipe, which has become modified in -various ways, though not so strictly in relation to the sounds produced -as is commonly supposed. The anatomical details of these modifications -cannot, or rather need not, be described now, save in the most general -terms. - -Briefly the syrinx, or organ of voice, of birds, is formed in part by -the lowermost rings which form the tubular windpipe, and in part by -the smaller pair of tubes which, running therefrom to the lungs, form -the bronchi. These last are formed of semi-rings only, the inner wall -of the tube being formed by very delicate translucent membranes. As -air is forced from the lungs along the bronchi and up the windpipe, -the modulation of the voice is effected by muscles which regulate the -amount of air driven through the syrinx, and the height of the column -in the tube; the latter being effected by muscles which alternately -lengthen and shorten it. - -So far so good. Next it is to be noted that this syrinx presents -a great variety of modifications, or types, differing not only in -plan, but also in the number and distribution of the muscles for its -manipulation. The most accomplished performers are to be found among -that great group of birds known as the Passeres, or perching birds, -wherein the number of these muscles is never less than five pairs, -and generally rises to seven. This association of musculature with -performance is exactly what we should expect. In Nature, however, -it is always the unexpected that happens. In the first place, the -females are, so far as the dissecting-knife and the microscope can -show, as well provided as the males, yet they do not sing. In the -second, the Nightingale and the Crow are equally endowed, so far as -we can discover, yet it is unnecessary to state that the talents -which the Crow possesses are never used! More disconcerting still -is the reflection that the Parrot, which is far less generously -endowed by Nature in so far as singing muscles are concerned, is -a much more skilful performer, inasmuch as it will reproduce with -equal fidelity the human voice and the song of the Canary! The latter -feat, at any rate, has been accomplished with amazing accuracy both -by the little Budgerigar (_Melopsittacus undulatus_) and the Quaker -Parrot (_Myopsittacus monachus_). In their wild state the Parrot -family are notorious for their discordant cries. It is therefore the -more remarkable that such feats should be capable of attainment. But -wherefore the elaborate syrinx of the Nightingale, if the simple type -seen in the Parrot is capable of the same result, and why the elaborate -syrinx in the case of the Crow, which never attains to a greater -perfection of vocal effort than the wild Parrot? - -One speaks of the syrinx of the Parrot as of a simpler type because of -its feebler musculature and the lesser complexity of its framework, but -it is nevertheless a more efficient instrument, since it is capable -of reproducing both the human voice and songs such as that of the -Canary. This fact becomes still more remarkable when we reflect that -the natural voice of the Parrot, as we have just remarked, attains to -no more than a harsh screech. How is it that, capable of so much, it -has achieved so little? The same question may be asked in the case of -the Raven. This bird has a syrinx indistinguishable from that of the -Nightingale, save in point of size; yet the Raven’s voice is never -musical, nor can it be trained to such an achievement. Like the Parrot, -however, it can be taught to speak, though its vocabulary is never -so extensive. One would have imagined that when the syrinx of, say, -the Raven, or any of the Crow tribe, was compared with that of the -Nightingale or the Skylark, some structural differences, commensurate -with the difference in performance, would be discovered; but such is -not the case. - -What interpretation are we to place on these paradoxical facts? One -cannot help asking why seven pairs of muscles should have been produced -by one group of birds to perform what can as easily be achieved in -another by two? It is true that the more generously endowed species -are musicians by birth, the others only by training. But one cannot -make a silk purse out of a sow’s ear. In like manner one asks why male -and female, possessing precisely similar voice-organs, should not sing -equally well, but they do not. Evidently mere mechanism does not alone -answer these questions. - -Some, perhaps, may see in them instances of what is known as -“Hypertely,” wherein the bounds of mere utility seem to be transcended. -Hypertely, however, implies something more than this: it implies a -shooting beyond the mark, the overdoing of a feature, where the -momentum gained, from some obscure cause, keeps on being increased by -cumulative inheritance: and not being checked by Natural Selection, -causes the species in respect of such characters to pass beyond its -congeners. Professor Lloyd Morgan’s theory of “over-production” would -seem better to apply here, though in a somewhat different sense from -that used by him. For in the instances just quoted there is a latent -potentiality for response to new demands which the struggle for -existence may make, but a potentiality varying in degree, and here -selection finds its _métier_. - -Yet further illustrations of secondary sexual characters, such as are -concerned with vocal music, must now be considered. The discussion of -these has been designedly deferred. They embrace instances of voice -production more singular than any yet referred to, and if possible more -difficult to interpret. - -The facts first to be reviewed concern the syrinx of certain of the -Anatidæ. It is noteworthy that each of the three divisions of this -group—the Swans, Geese and Ducks—contains species in which either -the syrinx or the windpipe has acquired some singular feature. In -the surface-feeding Ducks, modifications of the syrinx are most -frequently found. Commonly, as in the Mallard, this takes the form -of a spherical bony case; in the diving Ducks this bony chamber -has enormously increased in size. Furthermore it has conspicuously -changed both in form and character: for it is now roughly trihedral in -form, and its walls present large _fenestræ_ closed only by delicate -membrane, suggesting that the increased size of the chamber has not -been accompanied by a corresponding increase of bony tissue for its -construction. Hence all that is available is used for the construction -of girders to form supports for the now membranous chamber walls. -Some species seem to show that this fenestration has been pushed to -excess, leaving only vestiges of this singular chamber, as is shown in -PI. 21. In some species the bronchi are much swollen, and the syringeal -chamber has entirely disappeared: in others, as in the Merganser and -Goosander, a large syringeal chamber is supplemented by dilatations of -the windpipe. - -[Illustration: Plate 21. - -GRADES OF EVOLUTION IN THE SYRINX OR ORGAN OF VOICE IN THE MALES OF -SURFACE FEEDING AND DIVING-DUCKS. - -1. Wigeon. 2. Common Sheldrake. 3 and 4. Red-crested Pochard. 5. -Red-crested Merganser. 6 and 7. Long-tailed Duck. 8. Steller’s Eider. -9. Common Scoter. - -Face Page 126.] - -Save in the case of the Goosander, these peculiar structures are found -only in the male, but in the species first named the male, in addition -to the syringeal chamber, has two fusiform swellings in the windpipe, -one above the other: in the female one of these swellings is present, -but there is no syringeal box. - -This box is generally, and probably correctly, regarded as a sort of -musical instrument. Nevertheless the males are far less vociferous than -the females which have no such voice resonator. One has only to listen -to, and compare the notes of the Mallard drake and duck to discover -this fact. Here, then, we seem indeed to have a case of “Hypertely.” -Before, however, we build too much on this we must discover whether the -sibilant sounds uttered by the males do, or do not, play an important -part in arousing the sexual passions of the females. - -Certain of the Swans and Cranes afford illustrations of musical -instruments of an even more remarkable kind. Herein the windpipe at the -base of the neck enters a large chamber formed by the absorption of the -diploe sandwiched between the outer walls of the keel of the breastbone -and the enlargement of the space so created until it can accommodate -the tubular windpipe. This, entering the cavity in the form of a -loop, runs the whole length of the keel, the upper limb of the loop -finally running to the lungs. That we have here an indubitable musical -instrument there can be no question, for its possessor is enabled -thereby to utter loud, trumpet-like, if harsh, sounds. Here again only -the males are so provided. - -The profound interest of this really extraordinary association of -unrelated structures has never attracted the attention it deserves. -Originally, no doubt, one would have met with nothing more than a -loop of the windpipe impinging against the anterior border of a -normal, blade-like keel: later there would have been formed a broad -shallow surface on the keel at the point of contact with the loop, and -gradually the depression must have deepened till the bony chamber came -into being. By what nexus of sympathy were these reciprocal responses -made? - -Another very singular type of looped windpipe is that wherein the -trachea forms a series of coils between the body and the skin. It -is surely somewhat surprising to find that precisely similar coils -are met with in widely different groups of birds. Among the Passeres -they occur in the Manucode: among the Plovers in the Painted Snipe -(_Rhynchea rostratula_): among the game-birds in some of the Curassows, -and among the Anatidæ in the aberrant Australian Black-and-White Goose -(_Anseranas_). - -Very little is really known of the part played by these musical -instruments of the Anatidæ, nor, for the matter of that, of most of -the “musicians” among birds. Of some of the game-birds more has been -gleaned, and among these surely the most interesting is the love-song -or “lek” of the Capercaillie. With the advent of April the cock, just -before dawn, repairs to some favourite tree—used year after year—and -there performs a most astonishing if unmusical serenade; with -outstretched neck, drooping wings and spreading tail he gives forth a -weird, uncouth kind of song, more or less divisible into three parts. -He begins with a series of notes which remind one of nothing so much as -the sound made by two sticks knocked together at intervals of ten to -fifteen seconds, getting quicker and quicker, and changing in key till -at last they become bell-like. Then follows a series of sounds like -the drawing of a cork out of a bottle, and these end with bird-like -twitterings. By this time, however, the singer has worked himself up to -an ecstasy of fervour and passion so intense as to deaden him to all -that may be passing in the outer world. During these moments no sound -disturbs him, partly, apparently, because the excitement of the “song” -causes a turgid condition of the blood-vessels which for the time -effectually deafens him. “Sportsmen,” in Swedish and other European -forests, knowing this, select such performances as affording the most -favourable time for Capercaillie shooting, only cocks being selected. - -A survey must now be made of some of the more remarkable cases whereby -more or less musical, or rhythmical, sounds are made by instruments -of percussion; or by rapid vibrations. These are in almost every -instance formed by varying grades of modification in the feathers of -the wings or tail. Their presence, and their use, seem natural enough -until we recall the fact that many other birds without any apparatus -whatever, make sounds in no way less remarkable or less penetrating. -Pigeons, Nightjars and Owls, for example, can produce at will curious -snapping sounds by bringing the wings smartly together over the back. -The White, and Shoebilled Storks make castanets of the beak, throwing -the head backwards till the point of the beak touches the back, when -the jaws are set rapidly clashing one against another, producing a -sound comparable to the “bones” of negro minstrels, but without the -varying rhythm. As this performance is proceeding, the head and neck -are slowly moved through half a circle, till the tip of the beak -touches the ground, when the music ceases. As with the wing-snapping -just referred to, both sexes are equally skilful performers; but while -they seem to indulge in such exercises much more frequently, and with -more vim during the breeding season, they will break out after this -demonstrative fashion at all times of the year. But why, then, the need -for the yet more elaborate contrivances which are to be met with among -the Snipe, the Game-birds, and certain of the Passeres? - -However, be this as it may, in a large number of species a special -mechanism has been evolved to produce sounds which, as has been -remarked, in other species are no less effectually made without that -mechanism. - -One of the simplest of the cases is that furnished by the remarkable -“bleating” or “drumming” performances of many species of Snipe, -generally, if not only, when sexually excited, and especially of -the Common Snipe (_Gallinago cœlestis_) during its love-flights. -Mounting to a great height, this bird, at such times, suddenly turns, -and descends with prodigious speed, meanwhile holding the tail fully -expanded. The outermost pair of feathers are, however, specially -modified so that, in the first place, during this descent they stand -at right angles to the long axis of the body and well apart from the -rest of the tail-feathers. This alone, however, would not produce these -weird sounds, which owe their origin to the fact that these particular -feathers have their shafts conspicuously thickened and peculiarly -curved, while the vane or web of the inner side of the feather is -of great width and structurally differs from the vanes of the other -feathers, whereby the vane becomes more resistant to the rush of air -caused by the wings during the descent. - -But in the case of these Snipe it is to be noted this curious form -of musical instrument is found in both sexes, and there is little -difference in the quality of the sounds produced, but the bleating of -the male is said to be the more resonant. - -The Common Snipe is the best performer among several different species, -and it is to be noted presents, to a casual examination, no remarkable -or peculiar feature whatever—the structural differences just described -are only to be discovered by very patient scrutiny. But in the -Pin-tailed Snipe (_Gallinago stenura_) the number of the feathers has -been greatly increased, while at the same time their webs have been -so reduced that the outspread tail seems to consist of little more -than spines. With such a transformation one expects to find a quite -exceptional performance, far surpassing that of the Common Snipe. Yet -so far as observation and experiment go they effect absolutely nothing! -Here again we have a case where modification of structure has passed -the bounds of need and passed so far as to make the whole tail useless -as a sound-producing organ! - -A contrast and a parallel are afforded by some of the gallinaceous -birds of South America. The Black Penelope (_Penelopina nigra_) of -Guatemala, while on the wing, will, during its “love-flights,” pitch -suddenly earthwards with outstretched wings, and at such times a -crashing, rushing sound is produced, which has been likened to the -sound of a falling tree. Yet there is nothing in the shape of the wing -which will account for this. On the other hand, a near relation of -this bird, the Black-wattled Guan, _Aburria_ (_Penelope_) _aburri_ has -the four outermost primaries deeply incised along their inner vanes, -reducing the outermost portion of the feathers to mere spines. Yet, so -far as is known, this wing makes no especial noise. However, the males -of certain little South American Perching-birds known as Manakins have -the shafts of the secondary quills thickened to an extraordinary degree -so as to form solid, horny lumps, and these, when the wings are brought -together smartly over the back, produce a noise not unlike the crack of -a whip, so that here again structure and function are found together. -In the contradictory cases just cited where specialized parts are found -which are apparently functionless, we must suppose that the habit of -using them has been supplanted by some new stimulant. - -The part played by musical instruments of percussion would seem to -be a variable one. In some cases, and possibly in all, it may serve -as an excitant, or stimulant, to the rousing of a “sex-storm”; in -many, at any rate, such sounds serve as calls to the sexes when -separated. This much seems to be demonstrated in the case of certain -of the Woodpeckers, which in this matter differ conspicuously from -any other species yet referred to, in that they have developed no -special sound-producing mechanism, but make use of hollow trees which -serve them as drums, the beak being used as the drumstick. This is a -very noteworthy fact, for one would have supposed that here at any -rate, where the production of loud and far-reaching sounds is of -vital importance, the means would have been provided by some such -modification of the wing-feathers as we have already seen to obtain in -the case, for example, of the Manakins. More closely examined, however, -this apparent failure of the organism to produce its own mechanism -becomes less remarkable, for Woodpeckers are forest-dwellers and but -indifferent fliers; loud sounds produced by the rapid vibration of -the wings or tail, as in the case of the Snipe, in mid-air, are thus -impracticable, if not impossible, and sounds produced after the fashion -of the Manakins would not have sufficient carrying power. - -One of the most skilled performers among the Woodpeckers is the Great -Spotted Woodpecker (_Dendrocofus major_), whose weird drumming once -heard will never be forgotten. These sounds are produced by blows -of the beak on a branch, delivered so rapidly that the bird’s head -presents but a blurred appearance. The sounds thus made vary with the -resonance of the wood and can be heard at a distance of half a mile. -These strange vibrating notes are most frequently heard during the -courting season, and they will commonly beget a speedy response from -some more or less distant part of the wood, so that their purpose is -clear. They attain the same end as the bellowing of the stag or the -“lek” of the Capercaillie. They are, however, to be heard at other -times, as when the birds are greatly alarmed or when the nest is being -robbed. - - - - -CHAPTER VII - -THE SEXUAL SELECTION THEORY AS APPLIED TO BIRDS - - -Where the Rôle of the Sexes is reversed—Polygamy and how it is -brought about—Coloration and Courtship—Instinctive Actions—The -Importance of Landed Possessions—The Meaning of “Display”—The Springs -of “Behaviour”—A New Light on the Wild-duck—The “Display” of the -Great-crested Grebe—Some Neglected Factors. - -The significance of the varied behaviour of birds—more especially of -the males—during the period of reproductive activity must now be more -minutely analysed. But before this analysis can be profitably begun, it -will be necessary to recall the fact that there are several cases known -wherein the rôle of the sexes is largely reversed. Herein the females -do the “courting,” and fight one another as rivals for the males; while -the males perform the duties of incubation and brooding, and feeding -the young. This is really very remarkable, and demands more attention -than it has yet received. - -What factors have brought about this curious reversal? In any search -for an explanation it must be borne in mind that in all such cases -polyandry is the rule, and in all such cases the female is larger and -more vividly coloured than the male. Here, then, we have exactly the -opposite to what obtains in cases of polygamy. What is the reason -for this preponderance of males? Why is it that when the males are -in excess of the females the latter should be the more brilliantly -coloured and the more amorous? These questions at present are -unanswerable. When polygamy obtains it seems always to be assumed that -it is explained by the excessive pugnacity of the males, which, after -fierce contests for the mastery, take forcible possession of as many -females as may be captured and held in durance; the same argument seems -never to have been applied when polyandry obtains. There can be no -doubt but that it applies in neither case. - -When polygamy obtains, as we have already pointed out, the females -are not seized and captured by the males, they are not victims of a -lecherous lord. On the contrary, they seek the males, and the intensity -of the desire to satisfy their natural cravings extinguishes any -feeling of jealousy. - -The same interpretation must obtain where the numerical values of the -sexes is reversed. Failure to appreciate this accounts for one of the -many futile suggestions made for the suppression of the rabbit plague -in Australia, which was that large hauls of these pests should be -made by netting, and that the females should be slain and the males -released. This, it was held, would lead to the speedy reduction of the -latter, which would kill one another in their fights for the remaining -females. The plan was impracticable, but the suggestion demonstrated -the prevalent belief as to the attitude of the male in this respect. -Had it been well founded, surely polyandrous species, whether of birds -or beasts, would never have existed; for, by the reduction of the -males, monogamy would speedily have been restored. How, then, are we to -explain polyandry? How are we to explain the fact, as it seems to be -the fact, that the excess of males has brought about such a complete -reversal in behaviour—the males, instead of the females, requiring -the aphrodisiac? The solution of this problem probably lies with the -physiologist. We now know that the problem of sex does not rest merely -in the complete development of the primary sexual organs; we know that -fertile unions do not depend merely on the act of pairing, but on the -functional activity of those ancillary glands already referred to. And -it may well be that some change in the character of the secretions -has not only altered the numerical values of the sexes, but reversed -the normal rôle of coloration and behaviour. That is to say, neither -polygamy nor polyandry among the lower animals, at any rate, has been -brought about or is maintained by the excessive death-rate due to -combats for possession of mates, but must be explained as demonstrating -inherent changes in the germ-plasm, disturbing the relative proportions -of the sexes and correlated with a profound transformation, not only in -the behaviour of the sexes during the period of reproductive activity, -but also in their physical characteristics. - -The action of the primary sexual glands and of the ancillary glands -has, then, to be allowed for in all attempts to interpret behaviour -in sexual matters. No less so must this be the case in regard to the -development of coloration and other forms of ornament, and the genesis -of weapons of offence. But at present we are, in this direction, -dealing with an unknown quantity. The recognition of this, however, -should not deter us from attempting to solve the riddle of sex from the -phenomena which have so far been surveyed. - -To-day the interpretation which holds the field is Darwin’s theory -of “Sexual Selection.” But this was framed rather to account for the -existence of conspicuous secondary sexual characters—the antlers of -Deer, the train of the Peacock, and so on; it did not take cognizance -of the unarmed, and the soberly-clad individuals. But whatever -shortcomings we may discover, real or imaginary, in this theory, we -must never forget that he had not only to analyse and present his -facts, but he had first to collect them. This, in his case, was a more -laborious task than most people seem to suppose. Our criticisms to-day -are based, not so much on the revelations of new facts, as on the -harvests of his gleaning. Yet when all is said and done, the theory of -“Sexual Selection” remains, though perhaps in a new setting. - -To attempt to epitomize this theory is to essay a very difficult task. -But, in a condensed form, it may be said to be a theory which accounts -for the development of secondary sexual characters, on the one hand -through the agency of conquest by battle, whereby rival males strive -for the possession of one or more females, who have no choice in the -matter, or who may deliberately elect to follow the victor: and on -the other by display of conspicuous ornamentation, or of more or less -grotesque antics, or of some form of music, using this term in a very -wide sense. Wherever display is the agent, however, its purpose seems -to be to win the affections of the female to whom such attentions are -addressed. She is supposed to elect to mate with the finest performers -of a number of suitors. In this way, it is assumed, the intensity of -the display, whatever its nature, has been gradually increased. - -Wallace strongly opposed this, contending that it assumed too much, -that it assumed a common and uniform standard of perfection shared by -all the females concerned in the selection, which is indeed assuming -too much. But his own theory was no more satisfactory. Indeed it was -very much less so, for he contended that these various exaggerations -of colour and form are to be regarded simply as evidences of a -superabundant vitality, though there is no evidence that “superabundant -vitality,” if it exists, is a transmissible character. - -The revised version of the Sexual Selection theory advanced in these -pages is largely inspired by the work of Mr. H. Eliot Howard who, -in his Monograph on the British Warblers, has not only added very -materially to our knowledge of the life-histories of these birds, -during the reproductive period, but has also done much—both in the -direction of destructive, and constructive criticism, of generally -accepted conceptions on this head—to set us on the right track for -further research. - -A study of his work leaves one with the conviction that, while these -birds exhibit what we may call a nascent intelligence, their actions, -on the whole, may be described as instinctive, or congenitally -definite. That is to say, they follow one another in definite -sequence. Hence we must regard each new phase in the chain of events -appertaining to the reproductive cycle, as following one another in -a definite sequence, so that any break therein throws the orderly -performance of the necessary acts out of gear. There is no realization -of what reproduction means, no deliberate striving to achieve that -end. Each new phase brings its own set of associations and sets a new -train of actions in motion, which are performed mechanically. For -instance, these Warblers, like hosts of other species under similar -circumstances, are scrupulously careful to remove the fæces of their -young from the nest; thereby preserving it in a sanitary condition. It -is certain that any neglect to do this would speedily end in the death -of the young. This act is “instinctive”; it is not performed because -the parents have evolved any views on sanitation, and any strain in -whom this instinct was defective would speedily become eliminated. -Mr. Howard has demonstrated the mechanical character of this sanitary -measure by placing leaves in nests of young. The parents, having fed -their offspring, at once seized upon the leaf and commenced to dispose -of it after their usual fashion, first by trying to swallow it and then -by carrying it away. They did not, evidently, realize the difference -between the texture of the leaf and the milk-white, jelly-like envelope -which always encloses the fæcal matter of the nestling. We shall -probably never know how this most vitally important instinct came into -being; nor can we hope to discover what chain of happenings begot the -instinct, which each parent displays, to gently stimulate the cloacal -lips of their offspring in order to induce the discharge of the fæces -when this does not immediately follow the stimulus of swallowing food. - -We cannot credit these birds with notions on the importance of the -regular discharge of the evacuations. Equally mysterious is the -development of the envelope enclosing the fæcal matter. This is -jelly-like in substance, and of considerable thickness, and is enclosed -within a very delicate skin or pellicle, enabling one to lift the whole -in the fingers without soiling them. How and where it is formed should -not long evade discovery. But how it has come to be is another matter. -We can, at any rate, vaguely account for responses of the organism to -internal stimuli reacting directly on the individual, but here is an -elaborate mechanism evolved in response to extra-personal needs: and -which cannot be regarded as of exactly the same configuration as the -instinct to feed the young. - -A return must be made to the nature of the early phases in the -procession of the reproductive instincts. Mr. Howard’s study of -the Warblers seems to show conclusively that these first manifest -themselves in an overmastering desire to seize upon territory large -enough to ensure an abundance of food for the offspring that are yet -to be. To this end the males arrive from their far-distant winter -quarters at least a week in advance of the females. Since each returns -approximately to the scene of last year’s nursery, the arrivals are -fairly distributed at the first; but nevertheless this distribution -inevitably brings a conflict of interests between one or more males, -perchance young birds about to start in life, and having therefore no -definite objective. But whatever the reason, the competition is there. -The strongest male remains in possession, and immediately commences -to express the ecstasy of feeling which possesses him in continuous -outbursts of song. Such, doubtless, answer to the bellowing of the -male stag. They advertise the presence of a male to the female, who, -as she arrives, would seem to be already stirred by the rising storm -of sexual desire, for having once discovered a male in possession of -the all-necessary site for the nest, and the equally necessary domain, -each settles down to conjugal bliss: within twenty-four hours the task -of building has begun. There is evidently here no sexual selection -in Darwin’s sense: no choice from among a number of males of the -individual which most excites desire within her; but the mating of -the most mettlesome, most virile males has been determined before her -arrival and by a double sieve. In the first place, the duller-witted -birds fail to secure suitable territory, and in the second, the -territory, having been taken, must be held by force, so that only the -strongest males remain to mate when the females eventually arrive. -So far as one can see, selection is less exacting in the case of the -females, which apparently need do little more than respond to the -advances of the males. - -[Illustration: Plate 22. - -From a drawing by H. Grönvold. - -FIGHTING FOR TERRITORY. - -Two Black-caps are here seen fighting for their annual breeding -territory. A Chiff-chaff has been unable to resist the excitement of -conflict. - -Face page 140.] - -With the advent of the females the amorous instincts of the male -speedily gather force; but for their satisfaction it is imperative that -the female should be possessed by a like desire. To provoke this, for -it is essential to the well-being of the race that offspring should be -produced as early as possible, some form of aphrodisiac seems to be -necessary. This fact has never been properly realized, though it is -implied in Darwin’s theory of “Sexual Selection.” Here, however, it was -used to account for the evolution of resplendent coloration, eccentric -postures, and dances which, it was assumed, enabled or induced the -female to choose the most mettlesome males. What obtained among -sombre-clad species, appears to have excited no curiosity among the -students of the evolution theory. Hence it comes somewhat as a surprise -to find that the soberly-clad Warblers behave exactly as though they -too wore coats of many colours. After what has been said in the last -chapter on this head it will be unnecessary to describe these displays -among the Warblers in detail, more especially as my friend Mr. Howard -has kindly allowed me to use some of the illustrations from his book. -These show convincingly enough that the wings and tail are made to -play the same part as though they bore all the hues of the rainbow. To -bring this fact home compare the figures of some of these small birds -clad in sober russet and black with that of the Sun Bittern (_Eurypyga -helias_) in like mood, whose wings and tail when spread, and only then, -display bands of vivid chestnut-red, contrasting with bands of black, -on a background of grey and buff, variegated with delicate mottlings -and vermiculations of black and brown, and streaks of white. In the -case of the Warblers, it is to be remarked, the male, in these ecstatic -moods, will commonly hold a leaf, or a piece of stick, in his beak, as -if suggesting the work of nest-building and its delightful sequence. -This, or its equivalent, is a common phase, for the Great Crested -Grebe, for example, in these paroxysms will dive and bring up weed, -the nest material of the species, as an offering to his mate, or as a -stimulant to her yet slumbering passion. - -It seems clear, then, that the evolution of colour is not the -stimulant to display, for this is present where conspicuous colours -are wanting. Yet it can readily be understood how the association of -ideas in regard to colour and display arose, for there are cases where -this interpretation seems inevitable. Such are afforded by certain -sea-birds like the Kittiwake, Guillemot, Fulmar and Cormorant, wherein -the inside of the mouth is of a lurid orange-red in the case of the -first-mentioned, and of flaming gamboge yellow in that of the others. -During moments of sexual ecstasy the mouth is widely opened, as if -to charm the beholder with its gaudy hue. Both sexes have the same -colouring, and both behave alike. But it is doubtful whether either -is conscious that its own mouth is like that exposed to its gaze: the -action is sympathetic. No doubt it may play its part in stimulating -desire, but we cannot contend from this that it has been evolved -by sexual selection, that is to say, that the hues have undergone -a process of gradual intensification owing to the deliberate -rejection of the less gaily-coloured suitors. The tendency to develop -colour in the mouth would appear to be latent in all birds. - -[Illustration: Plate 23. - -_From a drawing by H. Grönvold._ - -THE DISPLAY OF THE GRASSHOPPER WARBLER. - -The behaviour of this bird under the stimulus of sexual excitement is -precisely similar to that of the Sun-bittern and the Kagu, yet it has -no brilliant colours to exhibit by such actions. - -Face page 142.] - -[Illustration: Plate 24. - -THE DISPLAY OF THE SUN-BITTERN. - -Quite inconspicuous in repose, this bird, in its moments of exaltation, -becomes banded and blotched with vivid colours, revealed by spreading -the wings and tail.] - -[Illustration: _Photos copyright, D. Seth-Smith._ - -THE KAGU IN DISPLAY. - -What is true of the Sun-Bittern is true also of the Kagu.] - -It is significant that whenever bright colours appear, they do so first -in the males, the females and young retaining the dress common, up to -this time, to the species at all ages. In the majority of instances, -at any rate, it would seem that this accession of colour appears with -the seasonal re-awakening of the reproductive activities: it forms a -“nuptial” dress, and is discarded after the breeding season is over for -a livery indistinguishable from that of the female, this forming the -so-called “winter plumage.” But if all the available facts are taken -into consideration there seems good reason to believe that the nuptial -plumage tends to be assumed earlier and to be retained later, as this -disposition to develop ornament gathers force, till finally only the -head and neck go into “eclipse,” as in the case of the Black-cock, -Jungle-fowl and Partridge. - -In the Pheasant we have an instance—one of hundreds—where the -resplendent dress is worn throughout the year. The next phase in the -direction of the growth of colour occurs when the female, towards old -age, develops a more or less well marked tendency to assume the hues -of her lord, and this accession of colour makes its appearance earlier -and earlier in succeeding generations, till finally the adults of both -sexes are coloured alike, save that, as a rule, the female lacks the -intensity of coloration which her mate displays. The original sombre -dress is now only worn by the young. In due course the resplendent -dress is assumed also by the young, as witness the numerous instances -among the Kingfishers and among the Parrots, where adults and young -are all habited in the same vivid hues. There are infinite variations -of these changes which cannot be discussed here, for obvious reasons. -All that matters now is the fact of such sequences, which inevitably -raise the questions: Why, in so many cases, do the females show no -disposition to assume resplendent colours? And to what factors can such -coloration, when it occurs, be attributed? The second only of these -questions is germane to the present discussion, and to this no very -satisfactory answer can be returned. - -To say that the development of brilliance in species hitherto sombrely -clad is due to “changes in the metabolism” is only an affectation of -wisdom. What we want to know is what induces the changes? Time was -when no more than a guess could be hazarded as to this: a suggestion -that ornament, of whatever kind, was one of the many modes of the -expression of that instability of the organism which is characteristic -of living things: that it was one of the outward and visible signs of -that inward, intangible tendency to vary which is so familiar. Later -research seemed to show, fairly conclusively, that ornament was one -of those “secondary sexual characters” which was dependent on the -stimulating juices, or “hormones,” emanating from the primary sexual -glands. To-day it is manifest that this is only partly true, for it is -certain that these glands are not alone concerned and they may only -participate indirectly. It seems to have been clearly demonstrated that -the thyroid and pituitary glands, or the “hormones” therefrom, play a -large part in this matter of the “secondary sexual characters.” - -Castration, it is true, profoundly affects these characters. In the -case of Deer it inhibits the growth of antlers, in Cattle the horns -are increased in length but reduced in thickness—they are longer than -those of the female, but resemble them in appearance, and further, -the whole stature is greatly increased, but it is at the same time -conspicuously less massive, particularly at the neck and fore-quarters. -In eunuchs it results in immense stature and the loss of the more -characteristic male features, such as the beard and the bass voice. The -removal of the testes in birds is always a difficult operation and is -rarely successfully performed. Hence the accounts of changes in plumage -consequent on this operation are inconclusive. It has generally been -supposed that whenever, either by removal or by disease, the testes -are rendered inoperative the plumage, when normally of a resplendent -type, assumes the coloration of the female. This is probably an -erroneous supposition, but what happens is a failure to secrete the -more intense pigments and the more specialized forms of feathers, so -that the resultant dress answers to the juvenile male dress. It is -not a case of “reversion” to this livery, but a failure to assume the -latest acquirements of the species. These, as has already been shown, -are only very gradually developed. The intensity of pigmentation, or -concentration of pigmentation, which results in sharply defined areas -of colour, is a cumulative process. As it loses in intensity at any -given moult, so the individual tends to reproduce the phases of the -earlier and vanishing livery. Sooner or later, however, this earlier -livery disappears more or less completely: is eliminated from the -system, so to speak: and what is commonly called lack of “vigour” -results, not in a return to the earlier, sombre dress, but in the -later-acquired, resplendent plumage lacking intensity. The seasonal, -temporary secondary sexual character has become, as some say, a -“somatic” character. Highly probable as this view appears, it ought, -it may be argued, to receive support from nestling plumages. Young -gulls, for example, should occasionally revert from the mottled to the -earlier striped livery. But we have no evidence of this; and it does -not follow that this sequence of events should occur. The conditions of -control are different. - -What exactly are the factors which govern the evolution of resplendent -plumage is not known. But they would seem to be more complex than -was supposed. That the primary sexual glands play an important part, -through the juices or “hormones” which they liberate, there can -be no doubt but these are only partial factors. The “hormones” of -the pituitary and thyroid glands are also necessary contributors, -controlling as they do both fertility and the more superficial -characters, such as colour and ornament. Evidence, indeed, is slowly -accumulating to show that the problem of the behaviour of animals -during the period of sexual activity, as well as the peculiarities -of structure and coloration which they develop at this time, are all -largely governed by the action of these secretions. - -These, in their turn, are undoubtedly inhibited, or increased, by -the control of the nervous system, though this control is of course -involuntary. This much seems clear from the fact that birds will -display when under the excitement of fear, though the character of that -display is never the same as that in moments of sexual exaltation. If -the nervous system, through the eye, by “suggestion,” played no part, -there could be no use for display, but it is equally certain that for -the realization of the sexual activities a number of other factors have -to contribute. - -The existence of this nexus of conditions is commonly overlooked, but -it is extremely important. Normally, not only among birds, but other -animals higher and lower in the scale of life, “suggestion” does not -suggest until the “hormones” concerned with the sexual activities have, -as it were, saturated the system and rendered it, so to speak, highly -inflammable. Even then it commonly happens that, with the male at any -rate, this inflammable state bursts into flame of its own accord. But -for this, indeed, how could the consummation—of the period of sexual -activity ever be realized? In many cases the sexes are sundered far -apart. What, but the merest accident, could bring them together if -it were not for this consuming fire of desire which impels each sex -to seek out the other? This stage is manifested in the case of the -Deer, where, we have seen, the stag wanders far and wide bellowing to -advertise his errand and listening for a response to his call. He is -possessed by a “male-hunger” which eventually attains to a state of -frenzy. Here no “suggestion” is needed, but the necessity for this -stimulus, for some form of aphrodisiac, occurs with him after the -first relief of his pent-up state has been attained. This stimulus is -applied, both through the eye and the sense of smell, by the females of -his herd. The same conditions apply in the case of the birds. But it is -to be noted that with the females, as in the case of mammals, sexual -desire is commonly less intense than in the males, and hence, in their -case the need for “suggestion” by display of some sort. But apart from -this, a “display” of some kind is necessary. How else can desire be -indicated? And here is “sexual selection.” For males, mate-hungry as -they might be, which resorted to no means of expressing their condition -would go mateless: and the same is true, though perhaps in less -extent, with the females; hence, then, it is clear display is a product -of sexual selection. - -That sexual desire is less intense in the case of the females is to -be regarded as another result of this form of selection. If they -displayed the same intensity of passion the males would speedily become -exhausted, for it is well known that the gratification of the sexual -emotions is far more enervating in the case of the male. It may well -be that polyandry has arisen from this transference to the females, or -development by the females, of increased sexual hunger. - -The fact that birds will repeat, albeit imperfectly, the phases of -the sexual display under the stimulus of fear, or anger, and when no -females are present, must be regarded as an indication, for we can -scarcely call it a proof, that exaggerated movements have become the -normal concomitants of great excitement, at any rate during the season -of reproductive activity. They are purely nervous responses to external -conditions. It must not be forgotten that, at this time, fear begets -other movements, equally striking, such as feigning lameness, and -death, which have no part in the sexual display. - -Interpreted in this light one can understand that to the female not as -yet sexually “ripe” or sexually “hungry,” these movements, when not -interpreted as signs of fear or anger, fail to produce any response. -So soon, however, as this period of “ripeness” arrives, the stimulus -through the nervous system produces the desired response, begetting a -complementary stimulus through the secretions of the sexual glands, -by what we may call the flow of the hormones; just as the sight of -food stimulates the flow of saliva, or “makes the mouth water” before -we are conscious of feeling hungry. In due time hunger will assert -itself without the stimulus of the nervous system through the senses. -But there must in any case be some form of display, some form of -communicating and stimulating desire between the sexes, to secure the -consummation of the reproductive acts. How else could intimation of sex -hunger be indicated and satisfied? - -That the desire for sexual congress is inherently more avid, more -intense, in the male than in the female is often called in question; -and more especially so by those who imagine that they have a mission -to carry on “social reforms” and to regulate the relations between -the sexes of the human race. Such aims and ambitions are commonly -those of the arrogantly ignorant. There are few people who possess a -sufficiently wide knowledge of this theme, or of the factors which -underlie it, to qualify them to become the mentors of their fellow-men -in these matters. However much we may choose to seek refuge in -sophistry, the fact remains that man is still an animal, and if the -human race is to continue he must always remain so. - -A lurid light has just been shed on the fierceness of the sexual -passion in the male by Mr. Julian Huxley, who relates some facts -pregnant with meaning to all who have understanding, in regard to what -obtains among birds. These facts are primarily concerned with the -Mallard (_Anas boscas_). This bird is ostensibly monogamous, and, on -the whole, seems to be a fairly considerate mate. The normal period of -pairing having passed, and the duties of incubation having begun, the -female ceases to harbour any further desire for sexual intimacy. Her -whole energies are devoted to nursing her embryonic young into life. -Not so the male. He is yet far from satiated; in him the sexual fever -still burns fiercely, but somehow he seems never to make any attempt to -provoke in his mate a like condition, as in the days before brooding -began. On the other hand, he does not scruple to savagely pursue -every other female who ventures abroad in his neighbourhood. So soon -as a duck takes wing for a brief relaxation from the arduous work of -brooding she is pursued by ten or a dozen already mated males, till -at last she is obliged to descend on the water, and with her descend -her pursuers, now to mob her without mercy. Commonly at least half of -these infuriated males will eventually succeed in treading her; leaving -their victim only after she has become completely exhausted or killed -outright. This is no unusual occurrence. On the reservoirs at Tring, -where every spring from one thousand to one thousand two hundred pairs -congregate to breed, from seven per cent, to ten per cent, of females -are annually killed in this way. - -It is just possible, however, that an error may have crept into these -observations. One cannot help asking, may it not be possible that these -pursuing males were actually unmated birds? The chief argument against -this is the fact that there is no sort of attempt to “display” apparent -with these birds, simply an overmastering, ravenous desire to satisfy -the craving which possesses them. - -Evidence is not wanting that the evolution of pigment intensification -and the consequent development of vividly coloured liveries, or -the equivalent development of ornament, has been accompanied by an -intensification of the reproductive instincts. For there can be no -doubt but that the display of species which are conspicuous for -their ornamentation is more animated than those of duller hues. As an -argument in favour of this view the case of the display of the Great -Crested Grebe may be cited, wherein each sex has developed both colour -and ornament to a high degree, and are distinguishable only to the -expert. - -The latest and the best exponent of the behaviour of this species -under the spell of sexual exaltation is Mr. Julian Huxley, whose -observations, in a condensed form, are now to be surveyed. The most -conspicuous features in this bird are the great Elizabethan ruff of -bright chestnut and dark Vandyke brown, and the long dark-brown tufts -of feathers, or “ears,” which surmount the head. But the satin-like -sheen of the white breast and the fore part of the neck and face add -not a little to the general effect. These ornaments are worn only -during the breeding season. So soon as the fires within begin to burn, -the parade of this finery commences, and it would seem that a somewhat -protracted dalliance takes place before any actual pairing. During -the early phases of these performances much play is made with ruffs -and “ears.” The courting pair will frequently face one another on the -water, and go through a strange ceremony of head-shaking. To this is -soon added a sort of ghost dance, wherein the male suddenly dives, -leaving his mate swinging excitedly from side to side. In a moment or -two, however, he appears, not suddenly, as usual, but arising gradually -out of the water. He seems to “grow” out of the water. First his head -appears, with ears and ruff extended, and beak pointed downwards; then -his neck, and finally the body arises into view, till only the extreme -tail end remains submerged, so that he looks more like a penguin than -a grebe! All the while he is turning on his long axis, as it were, -till he gradually displays before his mate the dazzling white sheen -of his breast and neck, set off by the rich red chestnut and brown of -his face and frills. A moment more and both subside into their normal -attitude, shake their heads at one another, and then proceed to feed as -if nothing had happened. - -But these quaint antics are only the preliminaries to still stranger. A -pair of birds, engaged, apparently, solely in fishing and feeding, will -suddenly approach one another and begin head-shaking, each striving -to outdo the other. Then the ears, till now erect, are thrust out -laterally, and the ruff is still further erected till it forms, with -the ears, a common disc. Then the hen dives: immediately after down -goes the cock. After some fifteen seconds or so she appears at the -surface again, speedily followed by the cock, who breaks out about -five-and-twenty yards off. Each crouches low over the water, and -each will be seen bearing a tuft of weed in the beak. As each sights -the other a tremendous rush is made, as if they intend to charge. -But when about a yard apart each springs up and assumes the penguin -position, save that the beak, instead of pointing downwards, is now -held horizontally and bears its burden of weed. Still approaching, -they eventually touch one another, treading the water and swaying in a -sort of ecstasy, all the while shaking their heads from side to side. -Then they gradually settle down into the normal swimming pose, though -still keeping up the head-shaking; then this, too, subsides, the weed -is dropped, and the performers drift apart and begin feeding. But no -actual pairing accompanies these strange performances. This final rite -is associated with a quite different ceremonial, and was witnessed -more than once by Mr. Huxley. On the particular occasion which he -describes he was watching a male swimming along near the reeds, -apparently on the look-out for something, and turning his eyes in the -direction of the course, he saw, at some distance off, what he supposed -was a dead grebe lying hunched up in the water, with outstretched neck, -and ruff and ears depressed. Presently the male swam alongside the -body and bent down his head as if to examine it. Then he swam to the -tail end, and suddenly scrambled out of the water on to the body; and -there, with bowed head and depressed ears and crest, he seemed to stand -a moment. Then he waddled forward over its head and into the water. -Instantly the supposed corpse raised its head and neck, gave a sort of -jump, and was swimming by the side of its mate. They had been pairing -on a half-made nest, whose surface lay level with the water. - -[Illustration: Plate 25. - -_From a drawing by H. Grönvold._ - -A MALE-SAVI’S WARBLER - -—in one of his “courtship” attitudes. Note the leaf held in the beak. - -[Face page 152.] - -Mr. Edmund Selous seems to have witnessed some almost incredible -behaviour on the part of the owners of a nest he had under observation, -inasmuch as, on more than one occasion, he declares the male lay prone -upon the nest and the female assumed the position of the male. After -this pantomime both would leave the nest, but commonly the female would -speedily return and pairing would be duly performed. - -This brief summary of Mr. Huxley’s observations, which he was generous -enough to give me the privilege of seeing in manuscript, taken in -conjunction with many other facts of a like kind given in these pages, -seems to lend support to the view that an excessive amorousness is -commonly associated with conspicuous ornamentation, as if these stood -in the relation of cause and effect. - -Finally, it is contended, the facts garnered during recent years -show that the theory of Sexual Selection, as Darwin propounded it, -especially in so far as birds are concerned, is no longer tenable: -but it is not an exploded theory, it has only undergone modification. -So far as the evidence goes, it would seem that the first of the -series of events in the sexual cycle is performed by the already avid -male, when he proceeds to secure a “territory” large enough for his -needs. In insectivorous and carnivorous species this area is fairly -extensive. No other male will be allowed within its confines. The -perfection of this instinct is vitally important, if sufficient food -for the offspring that are to be is to be assured. Where the food is -inexhaustible, as with the Auk-tribe, only a ledge large enough to -hold the egg is required. Only avid males will develop and respond to -this stimulus. The second stage occurs with the arrival of a female in -the area. She does not at once proceed to “select” her mate, passing -on if he fails to provoke her admiration. Her sexual condition is -apparently as yet but half awakened: to rouse this, the male supplies -an aphrodisiac in some form of display to which, in the normal course -of things, she responds, often also with some form of display, or -indication of the desire which has been aroused. The intensity of -the performance seems to vary with the intensity of the sexual -passion, which appears to be greater in some species than others, and -especially so with such as have conspicuously ornamental plumage. There -is, indeed, a variation in the sexual appetite as there is in the -ornamentation. The two are reciprocal, and are determined in degree by -the stimulatory qualities of the hormones of the sexual glands. Where -these have been developed in like intensity by the females, they also -display. Diminution in the quality and quantity of the stimulating -secretions of the ancillary sexual glands, the hormones of the -pituitary and thyroid, or the primary glands—testis and ovary—decreases -fertility, or induces sterility. Where these stimulants are lacking -there will be no desire, no display, and no pairing, and consequently -an end to this defective strain. Here then is Sexual Selection. - -[Illustration: Plate 26. - -Photo copyright by D. Seth-Smith. - -ANOTHER ASPECT OF THE KAGU’S “DISPLAY.” - -Herein two birds are seen facing one another with the great head-crest -fully erected. While in this mood these birds will strut up and down -with mincing gait and drooping wings. This is a posture commonly -assumed during momentary excitement, whereas the posture shown in plate -24 is apparently only assumed during moments of sexual excitement. - -Face page 154.] - -Instances of such impotency on the part of either sex are wanting, -and we can only speculate as to how such cases would be met. Would -a female who had chanced to settle in the territory of a male whose -sexual impulses carried him no further than seizing territory remain -with him throughout the mating season, held by an imperfectly roused, -ill-defined, sexual instinct? Or, eventually becoming mate-hungry, and -failing to stimulate him to perform his part, would she desert him and -seek another mate? On the other hand, would a male, failing to arouse -response in the female he had secured, drive her away and supplant her? - -In other words, are we then justified in postulating differential -effects in regard to display: a minimum of intensity to ensure mating? -A display of some sort is essential. It may be feeble as compared with -that of another species—that of the Sparrow, for instance, compared -with that of the Peacock—but it must be sufficiently good of its kind -to effect its purpose, which is to “hustle” up the production of -offspring. A phlegmatic but virile male, or a too feeble performer, -is almost as certainly doomed to extinction as an impotent male; for -his offspring will probably be eliminated by the adverse conditions of -existence to which their late appearance exposed them. Where a female -settles down with a male which does not attain to the standard of -display characteristic of his race, it is conceivable she may sooner -or later seek a mate elsewhere, deserting the phlegmatic bird as if -under the impression that she had made the mistake of settling down -with one of her own sex. There is no need that the female should have -to “select” the best performer of a number of males displaying at the -same time and place as a number of rivals. - -Finally, the ornamental crests and frills, and the vivid hues which -so many birds display have not arisen, as is generally supposed, as a -direct result of the selection, by the females, of the most vividly -coloured, or ornamented, from among a number of suitors presenting -varying degrees of intensity in ornamentation. Such “frills and -furbelows” are to be regarded as “expression points” of internal -variations in the germ-plasm, which have been free to develop along -their own lines because they have not proved in disharmony with the -conditions of the birds’ environment. Their development is to be -traced to the stimulating action of the “hormones” which control -both pigmentation and structure, as is shown by the fact that both -are modified by any interference with the glands in question. Such -ornamental features then are the concomitants not the results of Sexual -selection. - -The development of ornament, whether of colour or structure, may be -taken then as an index of specialization, and as one of the many -manifestations of that variation which is going on in every part of -every living organism. - -So long as the continued increments in the development of these -characters do not hamper their possessors in the struggle for -existence, they are free to go on developing. Sexual selection, other -things being equal, operates by according the greatest number of -descendants to the most amorous, and not necessarily to those of the -brightest hues. - -[Illustration: Plate 27. - -_Photos copyright, G. Herring._ - -SOME STRANGE ACCOMPANIMENTS OF COURTSHIP. - -THE WHITE-HEADED BELL-BIRD. - -This species is remarkable for the enormous, erectile wattles which -arise from the base of the beak of the male at the courting season. - -THE UMBRELLA-BIRD - -The crest which adorns the crown of the head has many counterparts, but -the long-feather clad wattle which depends from the fore-part of the -breast is unique. - -[Face page 156.] - -Plate 28. - -SKULL OF THE AMERICAN WHITE BEAKED PELICAN. - -The beak of this bird develops at each breeding season an irregular -horny plate which falls off at the end of this period, It is difficult -to regard this as a sexual “ornament,” yet it comes under this category. - -_Photos copyright, H.G. Herring._ - -HEAD OF A PUFFIN, SHOWING THE MOULTING OF THE BEAK SHEATH. - -At the breeding season, in both sexes, a triangular horny plate is -developed over the eye, an oblong plate below it, while the sides of -the beak become deepened by means of larger triangular horny plates. -All these embellishments are highly coloured, and they are shed at the -end of the breeding season. A further ornament is developed at the -gape, in the shape of a fleshy rosette of a bright orange colour. - -Face page 156.] - -But Sexual selection does not begin, and end, with the evolution of -frills and furbelows. “Behaviour” counts for more than is generally -supposed. This is as specific as “structure,” that is to say, it is -as constant for each species as is its coloration, and it is also as -variable. That Evolution may be determined by variation in behaviour, -no less than through structural variations, is a possibility which has -received but little consideration at the hands of students of Evolution. - -The singular history of the Australian Bower-birds lends additional -support to this view, and at the same time provides an additional -argument against the generally accepted opinion that bright colours -have been evolved by reason of the preference shown by the females -for the most vividly coloured of their suitors. For while the males -affect all the tricks and turns which are the common accompaniment of -courtship, they, in addition, introduce very extraordinary features in -the shape of “bowers” cunningly constructed and often gaily decorated, -as will be seen presently. Eight of the total number of species of this -group exhibit this behaviour, and while they differ very conspicuously -in coloration among themselves, they agree very closely in the type -of the bower they build. If the coloration is determined by the -female, then in this they display very different standards, and if -they do select, each according to the standard of the species, then we -must suppose that they also must exercise a choice in regard to the -character of the bower, the favoured male being the best builder. But -why, in this case, is there not as much diversity in the form of the -bowers as in the coloration of the feathers? A survey of the facts will -perhaps make this point clear. - -One of the best known of these bowers is that of the Satin Bower-bird -(_Ptilonorhynchus violaceus_). On either side of a platform of small -twigs a fence of similar twigs is reared, sloping inwards to form a -more or less complete tunnel. At the entrance to this is placed a -platform of sticks, which is strewn with a miscellaneous assortment of -brightly coloured feathers, bleached bones, and occasionally flowers. -The work of construction is almost entirely performed by the male: it -is indeed a little curious, having regard to the circumstances, that -the female should bear any share in its construction at all. - -Really this is a more wonderful piece of architecture than would appear -from the mere description of its main features: for it represents -psychical activities which are difficult to fathom. It does not take -the place of display, but is an extension of this. During his amorous -moments the cock becomes greatly excited, chasing his mate in and out -of the bower, carrying the while, in his beak, a brightly coloured -feather or a leaf. - -At the same time he sets all his feathers on end and every now and then -drops first one wing, then the other, accompanying these actions with -curious whistling notes and pretences of picking up food. - -The Regent-bird (_Sericulus melinus_) differs conspicuously from the -Satin Bower-bird, for while this is of a uniform, deep, metallic -steel-blue, the Regent-bird is jet black, with a golden yellow crown -and hind-neck and a great blaze of golden yellow on the wing. Yet -the bowers of the two species—which belong to different genera—are -practically identical, save that brightly coloured berries are used -more frequently by the Regent-bird. - -[Illustration: Plate 29. - -Photo by W. P. Dando. - -THE SATIN BOWER-BIRD AND ITS BOWER. - -The “Bowers” of the “Bower-builders” are the most remarkable variants -on “Secondary Sexual Characters” yet brought to light.] - -[Illustration: - -Photo by L. Medland. - -THE “BOWER” OF THE BOWER-BIRD. - -The “Bower” must not be confused with the nest, which is placed in a -tree and bears no sort of likeness to the bower. - -Face page 156.] - -The Spotted Bower-birds (_Chlamydodera maculata_ and _C. nuchalis_) -are quite dull-coloured species save for a vivid semicircular crest of -pink and mauve feathers which arise from the nape of the neck. Their -bowers differ from those just described in having a longer run and for -the immense quantities of shells which are deposited at each end of the -run. Some of them are brought from long distances, as is shown by the -large number of sea shells which are to be found in the collections -made by birds living far from the sea. - -By far the most remarkable of all are the bowers of Newton’s Bower-bird -(_Prionodura newtoni_) and the Gardener Bower-bird (_Amblyornis -inornata_). The first of these, a native of the Mountains of -Queensland, is somewhat strikingly coloured, at any rate so far as the -male is concerned, for he is of an oil green above and has a small -yellow crest, while his breast is of a bright yellow; the female, on -the other hand, is brown above and grey below. - -The Gardener Bower-bird, on the other hand, is of a sombre olive-brown, -but the male boasts an enormous crest of a flaming orange yellow. Yet, -widely dissimilar as are these two species, in the matter of their -bowers they display much in common. - -That of the Gardener Bower-bird takes the form of a hut-like structure -of twigs, arranged around a central support, commonly a very young -sapling. As a rule the thin stems of an orchid (_Dendrobium_) are used -in the construction of this curious hut, whose diameter is about three -feet. Before the entrance is a carpet of moss, which is kept clear of -leaves or debris of any sort, and on this the most vividly coloured -fruit, seed-pods, fungi, and flowers are laid, being constantly -replaced as they wither. Newton’s Bower-bird, in like manner, forms -a hut around a central column: a hut which may attain to a height of -as much as six or even eight feet, and the walls of the pyramid thus -raised are generally gaily decorated with flowers and fruit. Around -the central a number of subsidiary huts are not infrequently found, -and in and out of these the birds pursue one another in ecstasies of -excitement. - -We have in these facts some extremely puzzling features, which at -present, at any rate, permit of no more than a very rough analysis. -Probably the whole of these bower-building instincts have their origin -in the habit, which the males of so many birds exhibit, of carrying -a leaf in the beak when under the excitement of love-making. This is -suggestive of nest-building, and in many species this is actually -begun before the arrival of a female in the breeding territory, while -others build what are known as “cock-nests” which are never used. -Among the Bower-birds these “cock-nests” have taken a new and more -elaborate form, and are placed on the ground instead of in the trees, -the normal site for the nest in all these birds. Furthermore, stages -in the evolution of such strange fabrications can be found. These are -furnished by the Tooth-billed Bower-bird (_Scenopaeetes dentirostris_), -the Cat-bird (_Aeluredus viridus_) and the gorgeous Lawe’s Bird of -Paradise (_Parotid lawesi_)—which is not perhaps a Bird of Paradise. -These build no bowers, but are content with clearing a patch of ground, -of about ten feet in diameter, on which to disport themselves. But -while the “displays” of these birds closely resemble one another, in -the matter of coloration and ornament they present the most striking -contrasts. - - - - -CHAPTER VIII - -SOME “COLD-BLOODED” LOVERS - -The Courtship of the Crocodile—Amorous Lizards—Horned Chamæleons—A -flagellating Terrapin—The Frog that would a-wooing go—Semo musical -Frogs—Some marvellous instincts in Newts. - -The measure of the vitality of animals may be estimated by their -response to stimuli; and their behaviour increases in variety and -complexity as the nervous system develops. Our interpretation of -that behaviour commonly leaves out of account the character of this -responsiveness: we are apt to see proof of intelligence in acts which -should be read as instinctive. And instinct is to be regarded as a -co-ordinated response to stimulus, independent of prior experience. - -The complexity of this response stands in very close relation to the -structural complexity of the organism in which it occurs, and this -because an ever-increasing number of mechanisms and actions must be -set in motion to carry out the fulfilment of any given stimulus, as -this is traced from the lower to the higher groups of animals: till at -last we have to distinguish between movements that are merely reflexes, -and those which are “instinctive.” The latter must be fulfilled by -the former—the reflex actions are the agents of the instinctive. -Indifferent performance in either, endangers the existence of the -individual, and in some directions of the race itself. - -The sexual instincts, with which alone these pages are concerned, are -primarily stimulated and sustained by internal forces, generated, as we -have already seen, by the juices of certain glands whose relation to -the reproductive system has only recently been discovered. Though not -commonly realized, and though denied by some, the sexual instincts are -the dominant factors in the animal world. Even Man himself, the lord -of Creation, knowing good and evil, cannot escape their overmastering -rule. Commonly he is by no means inclined to rebel against this control -But there be some who, in their arrogance, imagine that its overthrow -is an end to be desired. Having scaled some slight intellectual -eminence they fondly imagine this feat was accomplished by virtue -of some spiritual grace of their own cultivation, and call to their -fellow-men to emulate their example. But such preceptors are labouring -under a strange delusion: they are suffering from a disease they wot -not of, a “Disharmony,” as Metschnikoff calls it, a disease which -blinds their perception of the motive power which has given them all -that they believe themselves to have created. For these same despised -instincts are the sacred fires of our being, and when they are quenched -all that makes us human, love, ambition, and life itself will be -extinguished. If the continuance of the race be a thing to be desired -it is well that the choice should not be left to us. - -Truisms are sometimes trite, and while it is a truism to say that -no race can continue which does not reproduce its kind, it is more -exact to say that, other things being equal, the race depends for -its existence, primarily, on the efficient working of the sexual -instincts. In the higher animals, the phenomena which these present are -so complex that they often assume something more than a semblance of -intelligent, purposeful behaviour. It is therefore necessary, for their -right understanding, that they should be analysed in animals lower and -lower in the scale of life till at last we come to the very simplest -types of organisms wherein instinct can be said to play a part. - -The lower we descend in the scale of animal life in our survey of -behaviour during the reproductive period, the more the evidence seems -to grow in favour of the interpretation of the Sexual selection theory -adopted in these pages—the view that neither the formal displays nor -the exaggerations of colour and ornament which so commonly accompany -them, are due to female choice; a choice not necessarily conscious, -but rather to be interpreted as the final abandonment to the finest -performer of a number of suitors. On the contrary, this ornamentation, -of whatever kind, is the expression of an intensification of the gland -secretions which is manifested by the process of pigment concentration -and a consequent intensification of coloration. Hand in hand with -these developments it would appear goes an exaggeration of the normal -movements which characterize the species when under the influence -of great excitement, whether of fear or pleasure. At any rate, the -displays of gaudily coloured and highly ornamental species are commonly -more striking than those of sober hue. - -On this rendering, the behaviour of Reptiles, Amphibia and Fishes, is -much more readily interpreted, and this is even truer of the more lowly -groups of animals such as Spiders, Butterflies and Beetles. - -Among the Reptiles, as among the birds and beasts, the desire to -obtain territory seems to be strong. But the information to be -gathered as to their behaviour in the search for mates, and after, is -exceedingly small. - -Sluggish by nature, all become animated under the stimulus of -mate-hunger, and this is especially true of the males. As one -would have expected, from what has just been said, desire is most -demonstrative in brightly coloured and highly ornamented species. But -even the dullest hued and most phlegmatic display quite surprising -agility and animation under the fever of Love. Thus among the -Crocodiles fierce battles are fought by rival males for the possession -of some coveted female: and later the victor strives to dispel the -apathy of his mate by caperings most undignified in a Crocodile. He -will twist and turn, or rather twirl, round on the surface of his -chosen pool, with head and tail raised high in air, and his capacious -barrel of a body swollen out to bursting point. These antics are -performed to the accompaniment of loud bellowings and roars heard at no -other season of the year. But more than this, an appeal is made to the -nose as well as to the eyes of his apathetic mate, for during all this -parade of love he exudes from glands in the lower jaw, and tail, an -almost overpowering smell of musk. At last, however, these antics have -their reward, for sooner or later apathy awakens into interest, and -interest ends in desire. - -The Crocodile is colourless, or at least is monochromatic; not so many -of the Lizards, which rival the birds in the vividness of their hues. -With the birds the colours undergo no changes save such as are due -to the incidence of light; with the Lizards, however, the bare skin -is exposed and this can, as it were, be made to blush with all the -colours of the rainbow. Having regard to what has been said already -as to the sources of this coloration it is not surprising to note -that here also the males are the more vividly coloured whenever the -sexes differ in this particular. And further, it is among the most -vividly coloured males that most animated displays take place when the -endeavour is being made to excite the amorous instincts of the females. - -The males of the genus Sitana are very brightly garnished. They possess -a large throat pouch, coloured blue, black, and red when expanded, and -this occurs only during moments of excitement, whether this is due to -fear or pleasure. And at the same time the vividness of the coloration -is greatly increased. No such secondary sexual characters are present -in the female. - -A variant on the throat pouch, of a much more striking character, is -displayed by the Frilled Lizard (_Chlamydosaurus kingi_), wherein -the tongue bones have become enormously elongated so as to project -backwards on each side of the body almost as far as the base of the -tail. With them they have carried a thin fold of skin, so that whenever -the mouth is opened these bones stand out at right angles to the head -and display a circular fold of skin stretched as it were on rods; or -they may be compared to the ribs of an umbrella. The great Elizabethan -frill thus formed, is displayed only during moments of great -excitement, and the open mouth, at such times, is flushed with a vivid -red, which, contrasting with the teeth, gives a very terrifying aspect -to prospective enemies, and doubtless also proves a valuable asset as a -“secondary sexual character.” - -The display of a vividly coloured mouth during moments of sexual -excitement, it may be remembered, occurs in some birds. Among reptiles -it is a common feature. A good illustration of this is furnished -by the Moustached Lizard (_Phrynocephalus mystaceus_), a native of -Southern Russia. When violently excited it raises itself on its hind -legs, curls and uncurls its tail, and opens its mouth to its widest -extent, presenting, to our eyes, a quite fearsome aspect. This effect -is immensely increased by the fact that the corners of the mouth -are provided with flanges of skin, which at this time swell up into -crescentic plates, the inner borders of which pass gradually into the -rosy lining of the mouth, thereby causing it to appear much wider -than it really is. So far this display has been witnessed only when -the animal is under the influence of fear. But since we find that -birds will make similar displays, both when under the stimulus of fear -and that of sex, we may assume, with no little degree of certitude, -that the same applies in the case of the reptiles, for the origin of -the ornaments is almost certainly to be attributed to the same gland -secretions which produce the secondary sexual characters of birds and -beasts. - -This, however, is no mere assumption, for we have some positive -evidence as to the association of bright coloration with “courtship,” -which has been furnished by Mr. Annandale, a naturalist of long -experience and having a first-hand acquaintance with tropical life. -He has given us a lively description of the courtship of the Malayan -Lizard (_Calotes emma_). “The males,” he says, “are very pugnacious, -and change colour as they fight. At the time of courtship a curious -performance is gone through by the male, the female remaining concealed -in the foliage hard by. He chooses some convenient station, such as a -banana-leaf, or the top of a fence, and advances slowly towards the -female. His colour is then pale yellowish flesh colour, with a -conspicuous dark spot on each of the gular pouches, which are extended -to their utmost. He stands upright, raising the fore-part of the body -as high as possible, and nodding his head up and down. As he does so -the mouth is rapidly opened and shut, but no sound is emitted. When he -is driven away, caught, or killed, the dark spot disappears entirely -from the neck.” - -[Illustration: Plate 30. - -Photo by W. Saville-Kent. - -THE BEARDED LIZARD. - -Paring moments of excitement the Bearded Lizard opens the mouth widely -displaying a vividly coloured interior. - -Face page 166] - -Normally sluggish, the Lizards display, it will have been remarked, a -quite surprising degree of animation when maddened by mate-hunger. Some -exhibit a considerable degree of pugnacity. In _Anolis carolinensis_, -for example, when two males meet they face one another, bob the head up -and down two or three times, expand the throat pouch, lash their tails -from side to side, and then, worked up to the requisite degree of fury, -rush at one another, rolling over and over and holding firmly with the -teeth. The conflict generally ends in one of the combatants losing his -tail, which is eaten by the victor. - -The Chamæleons include among their number species which have developed -quite formidable horns, recalling those of the Rhinoceros or, better, -of the extinct Arsinoetherium, since they are placed side by side -instead of one behind the other. In Owen’s Chamæleon there are three -such horns, two on the forehead and a median horn on the snout, and -these are borne only by the males. - -The marvellous play of colour which many Lizards display is commonly -attributed indifferently to “protective coloration” and to “sexual -selection.” It is unlikely that both have played equally important -parts in their development. If the case of certain of the Geckoes alone -is taken, then there would seem to be no doubt but that “Natural -Selection” was the agent which had determined their elaborations -for protective purposes, and in such and similar cases this may be -largely true. But the material which “Natural Selection” has worked -upon has been furnished by the secretions of the sexual glands to -which reference has so frequently been made already. These seem to -possess a very marked tendency to contain an excitant which promotes -the formation of intense pigmentation, or an excess of tissue which -may assume the form of weapons of offence, or of excrescences in the -form of spines, or other ornamental features. Animals in whom this -tendency to pigmentation and ornament has developed must, so to speak, -obtain a licence from “Natural Selection” if they are to retain it. -That is to say, if such ornament whenever it appears makes the wearer -conspicuous to its enemies, or hampers it in escaping therefrom, or in -fulfilling the ordinary avocations of life, then its further progress -will be inhibited, or the wearer will be exterminated. But the tendency -to produce colour, a by-product of the sexual gland secretions, may -incidentally serve to afford it a protective garb, and in this event -its further elaboration in the required direction is assured. - -In certain abnormal, sexually poisoned individuals among the human race -it is well known pleasure is derived from flagellation. There is but -one instance known to me where this obtains as a normal accompaniment -of desire among the lower animals, and this occurs in one of the -Painted Terrapins (_Chrysemys picta_), whose finger-nails are produced -into long, whip-like ends. I had the good fortune to witness their use -one day when in the Reptile House at the Zoological Gardens in London. - -The unusual activity of a male of this species was the first thing to -attract attention to his movements. Watched more closely, he was found -to be dodging a female and making frantic efforts to swim round so as -to oppose her path. This done, he closed up and immediately commenced -to apply the bastinado to her head. The movements were so rapid that -nothing more than a blurred image of these strange whips was visible. -As soon as she escaped his attentions, he set about circumventing her -again, and again succeeded: and this most extraordinary performance was -repeated many times during my watch. - -Turning to the Amphibia, the descendants of that stock which must be -regarded as the ancestors of the Reptiles, the version of the sexual -rôle which is adopted in these pages, that “Sexual Selection” in the -older, Darwinian sense, does not exist, finds further support. - -Among the tailless Batrachians—the Frogs and their kind—there is no -“display” immediately preceding the act of pairing. The males seize -upon the females and hold them in a close embrace which lasts for a -very prolonged period, covering many days or even weeks, until the -extrusion of the eggs, which he impregnates by successive emissions -of the fertilizing element. What controls the orgasm no one has yet -succeeded in discovering, but this is an important point, for it is -essential that the seminal fluid should not be emitted until the moment -the eggs are set free. The pairing act is here purely instinctive, as -is shown by the fact that if a Frog in embrace be removed and replaced -on some inanimate body, this will be treated as though it were a female. - -With the tailed Batrachians—the Newts and Salamanders—the male -commonly executes a very animated display which is followed by -behaviour of a quite remarkable character. The display, which is always -associated with vivid coloration, or the development of fin-like -frills along the back, takes the form of amorous writhings and other -gesticulations. At times he will hit his mate with his snout, and -at others he will simply rub sides with her, as if to entice her -to respond to his advances. These evolutions may be followed by an -amplexus, an embrace. In some species, however, these performances are -followed by behaviour which leaves one gasping with astonishment. - -To begin with, there is no act of pairing, no coitus, but the male -discharges a number of conical or bell-shaped “spermatophores,” each of -which is crowned by a bunch of spermatozoa, the male germs necessary -to ensure fertilization of the ova. These spermatophores adhere to -the bottom of the stream, and are gathered up by the female, either -directly, by placing herself in such a position that they can be seized -by the lips of the genital opening, or by seizing the spermatophore, -with its fertilizing germs, between her hind legs and pressing it -home! The more one contemplates this extraordinary proceeding the more -one marvels at the evolution of a departure from the normal sexual -relations so inconceivably strange. Here one sees the purpose of the -aphrodisiac in its true light. But for these facts it would have seemed -certain that its primary object was to enable the male to relieve -desire and at the same time to accomplish its end—the fertilization of -ova—without undue waste. And this, in all the cases so far discussed, -is possible only when the female has become inflamed with a like -desire for coitus. But here the male finds relief, without more ado, -by depositing the precious germs upon the ground. The display then is -indeed to serve as an aphrodisiac. For the continuation of the race now -rests entirely on the female. Any defect in the orderly working of her -sexual appetite means the waste of the spermatozoa and the failure to -effect fertilization. We cannot suppose that there is any realization -of these facts, or any deliberate action on her part, but rather that -she derives pleasurable feelings from the necessary passage of the -spermatophore, which, probably, she recognizes by its smell. - -The statement that the Frogs and their kind dispense with a display -requires some qualification. For in the first place they, like their -tailed relatives the Newts, develop secondary sexual characters, but -these are of a quite peculiar kind. Among the Newt tribe, as has been -mentioned, these characters take the form of frills and crests and -vivid colours. They are intended to stimulate through the sense of -sight, and arouse emotion, as a city is beflagged to welcome those it -may delight to honour. The Frog tribe appeal to the musical sense, even -though that music be of a barbaric kind. But, it would seem, when once -the errant females have been drawn to the spot chosen by the males, no -further aphrodisiac is used, the male simply seizing upon the female -nearest at hand and, having once embraced her, she is not released -again until the eggs have been extruded and fertilized. To maintain -his hold, the forearm is often excessively muscular, while one or more -of the fingers may be armed with pads. In some cases, as with the -Himalayan _Rana liebigi_, the inner side of the arm and each side of -the breast are studded with small conical spines. But the absence of -ornament in these cases, as with such of the Newts where there is no -amorous display before embrace, is significant. - -And now, as touching the musical performances of these troubadours. -These commence in the early spring. With many species, as with our -Common Frog (_Rana temporaria_) nothing more than loud croakings are -attained. But with others this “music” is enormously increased in -volume by resonators in the form of air-sacs or wind-bags. We may -surely, with some show of certainty, liken this “music” to the song -of birds, and assign its primary purpose to the same cause—a device -to advertise their presence to wandering females seeking mates. That -birds sing after mates have been found, and later, is no doubt due -to a general feeling of “fitness,” which finds expression in what -has become the usual mode for such emotional states. Most people -must have heard the spring concerts of our Common Frog; but these -are incomparably surpassed in volume by the Edible Frog and the Bull -Frog, which are provided with large, globular, inflatable, air-sacs in -the throat, serving as voice-resonators. Such performances, however, -are mere bawlings compared with some other species, which mew like -cats, or bark like dogs. The most famous of all is the Brazilian -“Ferreiro” or “Smith” (_Hylodes faber_), whose voice is one of the -most characteristic sounds to be heard in Tropical South America. -“Fancy,” says Dr. Gadow, “the noise of a mallet, slowly and regularly -beaten upon a copper plate, and you will have a pretty good idea of -the concert given generally by several individuals at the same time -and with slight variations of tone and intensity.” When seized, the -performer utters a “loud and shrill, most startling cry, somewhat -similar to that of a wounded cat.” Another, a Paraguayan species, -_Phryniaxus nigricans_, at the breeding season, utters a call note -which consists of two clear, musical “rings,” followed by a long -descending “trill” like that of our British Greenfinch. But, it is to -be noted, both sexes in this case perform. - -The period of sexual activity with perhaps the majority of animals -is intermittent and extends over but a short period annually; with -others potency is continuous, at least with the males, though desire -becomes clamant only when aroused by external stimuli. But whenever -this condition be aroused it invariably finds expression in exaggerated -movements or vocal demonstrativeness. It uses the normal channels of -expression, in short, but intensifies them. Now this period of sexual -activity represents the maximum of “fitness” in animals, and it is not -surprising, therefore, to find that when the barometer of vitality -stands high some approach to the maximum of activity is indicated. In -many birds this is revealed in song, though the earlier stimulating -cause is absent. Among the cold-blooded frogs the same obtains. In the -Edible Frog (_Rana esculenta_), for example, the males, which “are -great musicians,” remarks Dr. Gadow, “go on singing for sheer enjoyment -not only during the pairing time, but throughout the months of June and -July. Warm, moonlight nights are the favourite times for the concert, -which takes place in the water, beginning at sunset and continuing till -early dawn. A few individuals utter a single note, ‘gwarr-oo-arr’ or -‘coarx’ but these are only preliminaries. The precentor ... begins with -a sharp-sounding ‘brekeke’ and this is the signal for all the others -to chime in with the same note, varied with all sorts of other sounds, -bass, tenor and alto, each performer filling its resounding vocal sacs -to bursting size, and these bags then look as if they acted as floats. -When there are several hundred of these sociable creatures the din is -continuous, and may be heard more than a mile off.” - -From what has been said of the Amphibia, and especially of the -Newts, it would seem that, among the land vertebrates at any rate, -the sexual instincts in this lowest or simplest form are satisfied -with the discharge of the germinal products. Many, however, have -advanced a stage further and reveal the rudiments of that instinctive -care for offspring which develops to higher and higher grades as we -ascend in the animal kingdom, till at last, in the human race, where -the offspring is desired for its own sake, we ascend to the highest -plane of all. The varied means of expression which these rudimentary -instincts take in the Amphibia have already been discussed in “The -Infancy of Animals,” which preceded this present volume, and hence no -more need be said on this head in these pages. - - - - -CHAPTER IX - -LOVE-MAKING AMONG FISHES - - -Germinal Variations—Fishes and mate—hunting—Some remarkable Sexual -differences displayed by the Teeth of Rays—The Double-eyed Fish—The -Coloration of the Dragonet—Some curious facts about Salmon—The strange -use of the kidneys in the Stickle-back—The Stickle-back and parental -duties—Siamese Fighting-fish. - -Mammals, Birds, Reptiles and Amphibia, as has already been shown, -all exhibit practically the same line of conduct in regard to their -mate-hunting instincts; all use like modes of expression. And this is a -very significant fact. It becomes more so when we turn to the fishes, -for here again we meet with the same behaviour, and here again we meet -with the same rules in “secondary sexual characters.” - -An instance or two of the latter distinction between the sexes should -suffice. As a rule, among fishes, the males are smaller than the -females: commonly there is no other external distinguishing feature -between them. In many cases, however, the males are more or less -strikingly different, thereby showing a departure in the nature of a -higher degree of complexity, or “specialization,” just as obtains among -the birds. And the same sequence to this also obtains. That is to -say, as has already been remarked in the case of the Mammals and the -Birds, the new features first appear in the males, leaving the females -and young of both sexes unmodified. A singular illustration of this is -afforded by some of the Rays, or Skates, as they are often called. In -the Thorn-backed Ray (_Raia clavata_) the teeth of the adult male are -sharp-pointed and directed backwards, while those of the female are -broad and flat, forming a sort of mosaic or pavement. The young male -agrees with the female in this respect. In the Common Blue Skate (_Raia -batis_) the teeth are pointed in both sexes, though more so in the -adult male. In the Spotted Skate (_Raia maculata_) the teeth are fully -pointed in both sexes. Here, then, the normal course in the evolution -of new characters is followed, but it is remarkable that the teeth, -which are so intimately related to the capture of food, should be thus -affected. Whether the change of teeth is associated with a change of -food, or whether neither pointed nor pavement teeth affect the feeding, -is unknown. - -Still more remarkable is the case of the Double-eyed Fish (_Anableps_). -In this fish there is an intromittent organ in the shape of a tube -which is formed by a continuation of the urinogenital ducts down the -front of the anal fin. In the hinder half of this organ a bend is made -either to the right or left. Out of seventeen males, this bend was to -the right in eleven, to the left in six. Further, there is a small -fleshy tubercle at the side of the anal fin-ray, at the middle of its -length. When this prominence is on the left side, the organ bends to -the right; when it is on the right, the bend is to the left. In the -females the genital opening is covered by a special scale, which is -free on one side, left or right, and not on the other. Thus copulation -is possible only from the side, and a left-sided male can only -conjugate with a right-sided female, and vice versa. Here is one of the -most extraordinary cases of specialized secondary sexual characters -known. How do the sexes distinguish their complemental mates? It is -important that they should, for unions are otherwise impossible. - -In the Dragonet (_Callionymus lyra_) the male differs conspicuously -from the female in being much the larger—an exception to the rule—and -in having the fin-rays enormously elongated. Further he wears a -conspicuously resplendent livery, but this is strictly a “nuptial” -livery, the colours waning as soon as the period of sexual activity is -past. That these colours play the same part as with the birds is clear -from the observations of the late Saville Kent. “The male,” he says, -“resplendent in his bridal livery, swims leisurely round the female, -who is reclining quietly on the sand, his opercula distended, his -glittering dorsal fins erect and his every effort being concentrated -upon the endeavour to attract the attention of his mate.... The -female, at first indifferent, becomes at length evidently dazzled by -his resplendent attire and the persistency of his wooing. She rises -to meet him, the pair so—far as is practicable with fishes—rush into -each other’s arms, and with their ventral areas closely applied, ascend -perpendicularly towards the surface of the water.” In the course of -this ascent the ova and sperms are shed, and fertilization takes place. - -The difficulties in the way of the study of the behaviour of fishes -during the critical period of mate-hunting are many and obvious. -Something may be inferred from the nature of the secondary sexual -characters which they exhibit, and more definite information can be -obtained from such species as can be kept in aquariums. From these -two sources enough has been gleaned to show that these cold-blooded -creatures, in many cases, exhibit the same emotions and the same means -for their fulfilment as the higher vertebrates. And it is significant -that wherever anything approaching what may be called “Courtship” -obtains, the males commonly exhibit secondary sexual characters, -whether in the form of ornament or of armature; while among species -which consort in shoals during the breeding season no such distinctions -are present. The ova and milt are shed and fertilization takes place as -they escape. - -Comment is frequently made in works on Natural History on the fact that -among fishes the males are commonly smaller, often conspicuously so, -than the females. Among mammals the males are the larger; but among -birds this is by no means always the case. It is somewhat surprising to -find this discrepancy among the birds of prey, where, as in the case -of the Sparrow-hawk, the male is little more than half the size of his -mate; commonly, however, there is little or no difference. Among the -fishes the differences are often much more marked, as for example in -the Conger-eel, wherein the male never exceeds a length of two feet -six inches or a weight of one pound; females, on the other hand, may -exceed eight feet in length and attain a weight of one hundred and -twenty-eight pounds, though such giantesses are rare, but specimens -of fifty pounds and upwards are frequently met with. The explanation -of this may lie in the fact that among fishes it is no uncommon thing -to and males becoming sexually mature long before they have attained -their full stature. With the Salmon, for instance, ripe spermatozoa -have been found in individuals of not more than a few inches in length, -and in this species also the male is the smaller. Ova take longer -to attain maturity, for in addition to the germ-plasm they must be -provided with a more or less extensive amount of food-material in the -shape of yolk. The formation of this is inhibited until the demands -on the system for the building up of the body have begun at least to -lessen. - -Mate-hunger among the fishes seems generally to find peaceable modes -of expression, either in “display” or in consorting in vast shoals, -though, so far, the factors which govern their conduct in this matter -are as yet unknown. But here, as with the higher vertebrates, there -are some species which adopt more violent methods. A good illustration -of such conduct is furnished by the Salmon, which, during the period -of sexual activity, develops a curious modification of the lower jaw, -which is produced forwards and upwards to form a hook-shaped projection -of fibrous tissue. When the mouth is closed this hook is received -into a cavity formed within the fore-part of the roof of the mouth. -It has been described as a weapon of offence. But this it can hardly -be. On the other hand it has been suggested that it serves to protect -the jaws when charging a rival, for the shock on such occasions is -considerable. It answers, in short, like to the fibrous mass of tissue -which protects the fore-part of the head in Whales like the Black Whale -(_Globicephalus_) and the Bottle-nose Whale (_Hyperoodon_), serving as -a battering-ram. In the Pacific Salmon (_Onchorhynchus_) both jaws are -hooked, so that when the mouth is closed the hooks cross one another -as in the beak of the Crossbill. In this Salmon, too, the front -teeth attain a considerable length, while the body becomes laterally -compressed and a hump forms at the shoulders. Little, however, seems to -be known as to the nature of their battles. - -The combats of the Salmon of our own islands, however, are evidently -severe, and this has long been known, for Darwin speaks of as many -as three hundred, all with one exception males, being found dead in -the Tyne during the month of June, killed by fighting. Such battles -are fought, it is to be noticed, not so much for the possession of -females—for it is a polygamous fish—as for the privilege of fertilizing -the eggs as they are shed. The absence of a “display” here is a -noticeable feature, and it is on this account, probably, that the -reproductive period is not associated with the appearance of any form -of resplendent livery. On the contrary, the marvellous silvery sheen -which adorned both sexes on their arrival at the spawning ground from -the sea has entirely vanished by the time that the consummation of the -journey has been attained, and in its place is naught but a slimy, -dingy copper-coloured hue. But no sooner has the reproductive period -passed than the silver lustre makes its appearance once more. - -These facts are the more interesting when contrasted with what obtains -among other fighting species which must woo the females. Take the -case of the common freshwater Stickle-back. In this species the body -is invested with an armature of bony plates and spines in place of -scales, while the males are arrayed in vivid hues of red and blue. Any -survey, however, of the reproductive activities of this little fish -must take into account certain quite remarkable prenuptial actions -and instincts. Briefly, before the male commences his search for a -mate he constructs a nest of fine fragments of aquatic weeds, which -are held together, not by interweaving as with birds’ nests, but by -a sticky and copious secretion from the kidneys. According to some -authorities, this secretion is to be regarded as a pathological product -caused by the undue pressure of the ripening testes. It is difficult -to accept this interpretation, for it might with as much reason be -argued that the copious secretions of the salivary glands of the edible -Swift—which builds a nest constructed entirely of hardened saliva—are -also pathological in character. But be this as it may, the nest -completed, the male seeks a mate, or mates, for polygamy is the rule of -his tribe. In his search for these he has constantly to do battle with -other males, whom he endeavours to disembowl by swift rushes contrived -to t rip open his rival as he passes, by means of one or other of -the erectile spines which project from his back and belly. With the -females whom he desires he uses the arts of peaceful persuasion, -swimming backwards and forwards before her in his endeavour to excite -her amorous instincts. At last he persuades her to enter his bower and -deposit a few eggs, fertilizing them immediately they are laid. The -first to enter leaves by forcing a passage through the opposite wall -of the nest, a happy contrivance, for thereby a current of water can -be constantly driven through, leaving fresh oxygen to the developing -eggs. One female after another is inveigled into the bower, until the -complement of eggs is complete. These, singularly enough, are now taken -charge of by the male. He it is who creates life-sustaining currents -which bathe the eggs, by the rapid vibrations of his breast-fins, and -he it is who protects them from their most persistent enemies—the -females who laid them. As soon as the fry appear the duties of the male -are still further increased. He must guard them from their mothers, -and other foes, and he must prevent their too extensive wanderings. -Such as stray too far afield are sucked into the mouth and brought -back again to the nursery, where they are set at liberty by a reversal -of the sucking action. That the male of a polygamous species, and -with all the attributes of a polygamous species—pugnacity and vivid -coloration—should take upon himself the duties which under like -circumstances among the higher vertebrates are undertaken by the female -is a very remarkable and puzzling feature. In this species, in short -the male plays successively a polygamous and a polyandrous rôle. - -Strange as these facts are, they are not apparently without parallel -among fishes, for certain of the labyrinth-gilled fish present many -features in common, though as yet proof seems to be wanting. Thus -the small Siamese “Fighting Fish” (_Betta pugnax_) is endowed with -so ferocious a nature that it is kept, as the Malays keep fighting -cocks, for the amusement of native sportsmen, two fish being pitted -against one another and large bets being made on the result. In a -state of quiescence it presents no very remarkable coloration, but if -two be brought together, or if one sees its image in a looking-glass, -it becomes thrown into a paroxysm of rage, the fins are raised and -the whole body becomes irradiated with metallic colours of dazzling -beauty. There can be no doubt but that a like play of colour occurs -during moments of sexual excitement; it is highly probable that it is -polygamous. Of its breeding habits, however, little or nothing seems to -be known. Not so, however, in the case of a closely-related species, -less pugnacious in disposition, but almost as vividly coloured, in so -far as the male is concerned. Now in this species a nest of froth is -made and the eggs, after deposition therein, are jealously guarded by -the male. Hence, on these facts, we may assume with a fair amount of -certainty that the closely-related “Fighting Fish” displays like habits. - -That the Reptiles, Amphibia and Fishes have much in common with one -another, and with the higher vertebrates, in the manner of their -love-making is indisputable. We find no evidence anywhere that the -first faint throbbings of the sexual pulse in the female are quickened -to fever beats after the efforts of several successive wooers, each -more demonstrative than the last, to arouse this state—the conditions -required by the Sexual Selection theory. But successful mating depends, -in each year, on the sexual fitness of the male himself, and the mate, -or mates, which for that year he has taken “for better or worse.” It -is possible, of course, that a male, ambitious but impotent, will -be forsaken by his mate; it is possible that a female of low sexual -vitality may fail to respond to the most impassioned displays; in -either case no offspring result, and thus the failures are eliminated. -It is possible that here, as with the higher vertebrates, coition -may by no means always be immediately preceded by display. But the -“display” has done its work. It has stimulated the sexual appetite, as -the sight of tempting food stimulated the bodily appetite. - -But both the Amphibia and the Fishes reveal a lower plane of the sexual -instincts, when the sexes, dominated by some imperious instinct, -gather in hordes, commingling to shed their precious germs into the -surrounding water, there to effect the work of fertilization and the -achievement of new birth. The all-important union of these germs is -no mere work of chance, as it might seem, but the sperms seek the ova -with unerring surety, guided, in this case, by that very efficient -substitute for instinct, chemotaxis, or the attraction which certain -chemical substances have for lowly organized living bodies. In -this case the allurement is furnished by the ova. It is surely no -unreasonable surmise that here we have the beginnings of the complex -phenomena which the earlier chapters have revealed. On this lower plane -we are probably confronted by instinct alone, but from this level -upwards intelligence plays an increasingly important part. - - - - -CHAPTER X - -SOME OF THE “LOWER ORDERS” - - -Butterflies and Moths, and the Coloration of their Wings—Female -Choice and “Fine Feathers”—When Male Butterflies are Dominant—Sexual -Selection among Butterflies—Abortive Experiments—Wallace and the Sexual -Selection Theory—The Sense of Smell in Butterflies and Moths—Fragrant -Butterflies—Wingless Moths and their Lures to Lovers—Methods of Pairing -among Butterflies and Moths—More Experiments. - -Not the least impressive feature met with in the study of animal -behaviour under the spell of the Sexual Instincts is its uniformity. -This fact becomes the more apparent as one turns to the lower grades -of life. Whether one starts with the vertebrates and works downwards, -or vice versa, the same problems arise and the same interpretation is -demanded. That is to say, the theory of “Sexual Selection” leads one to -the same conclusions whether it be tested by the evidence afforded by -the Butterflies and Moths, or that furnished by Birds or Mammals. - -The accessory phenomena, the vehicles which give expression to these -internal fires, are in like manner curiously similar. These “vehicles” -are the “secondary sexual characters”—colour, and armature, and scent. -These very tangible signs are the phenomena in the Mystery Play of Sex -which first catch the attention of the investigator. To account for -these the theory of “Sexual Selection” was first devised. - -After the birds, probably the group most conspicuous for its splendour -is that which contains the Scale-winged Insects or Lepidoptera, and it -has always been allowed that any explanation of the one must apply also -to the other. It seems impossible to avoid this conclusion. But before -going further it would be well to take note of one or two interesting -features in regard to coloration that have so far not been touched upon -in these pages. - -The Coloration of Animals is generally regarded as a by no means -fortuitous feature, but one, on the contrary, controlled and -determined by various factors. Hence are recognized various kinds -of coloration: Obliterative or Protective-resemblance Coloration; -Warning Coloration; Mimetic Coloration; and Epigamic Coloration, or -the colours associated with courtship. These various types have been -subdivided and accorded technical labels by Professor E. B. Poulton, -in his “Colours of Animals,” but these need not be enlarged upon -here. Suffice it to say that it is generally held that all forms of -coloration can be explained, and all can be labelled, as to their -origin, with more or less certainty. There are those who doubt the -warranty for this classification. Commonly, it must be admitted, the -arguments of these sceptics are not impressive; they are sometimes even -stupid. That such coloration, however it be labelled, is subjected to -some control seems to be shown in the case of the Lepidoptera, for, -generally, in the Butterflies, the upper surface of the wings is much -more vividly coloured than the under surface, and this, apparently, -because when the creature is at rest the wings are brought up over -the back like the leaves of a book, so that the brightly-tinted, and -therefore conspicuous, area is concealed, as, for example, in the “Red -Admiral.” With the Moths the wings, while the creature is at rest, are -held horizontally, and it is the upper instead of the under surface -which is exposed, but the hind-wing is covered by the fore-wing. The -coloration is here very different; for while the exposed surfaces -of the fore-wings are commonly soberly tinted, the hind-wings may -be quite glaringly coloured. These bright colours are exposed only -during flight, or during moments of unusual excitement, as in the -case of the Eyed Hawk-moth. According to Weismann, this insect when -alarmed raises the fore-wings so as to expose the “eye-spots “on the -hind-wings, which, with the increased area of the wings, impart a -terrifying appearance to the body to would-be assailants. This is -as it may be, but for the moment the feature to be insisted upon is -that the bright colours are almost invariably hidden when the insect -is at rest, and by quite different means, determined, apparently, by -the different carriage of the wings. Now, according to some, bright -colours are begotten by strong light, but in the Moth and Butterfly the -surface area of the wing which is most exposed is the surface turned -to the light during rest, and this is the least coloured. The curious -relation between this coloration and the resting position is strikingly -illustrated by the case of one of the “small Blues” (_Lyccenæ_), cited -by Weismann. Herein the male, which has the upper surface of the wings -of a bright blue, rests in the position common to Butterflies—with the -wings raised and concealing the bright colour—while the female, which -has the upper surface of a dull brown, rests with the wings expanded. -As, however, the concealed under surface is not brightly coloured, it -is difficult to believe that these different postures and conspicuously -different colours can have been brought into existence solely by the -action of Natural Selection, which, it is generally contended, has -brought about the extinction of those individuals which neglected, -when resting, and therefore liable to be “caught napping,” to conceal -their arresting colours. There is, indeed, no apparent reason why the -female, which has nothing to conceal, should depart from the custom -common to Butterflies, of resting with the wings closed and raised, -this position effectively protecting the male. The facts seem to show -that the coloration of the exposed surfaces of the wings is determined -primarily by some physiological factor rather than by the incidence -of Natural Selection directly through external agencies. Thus, for -example, the action of light on the surface of the wings when in the -resting posture may well inhibit the production of vivid pigment owing -to some inherent physiological idiosyncrasy. But any individuals which -lack this inhibiting factor—as some species which, though resting, are -brightly coloured, appear to do—will be eliminated, if they live in an -environment harbouring eliminating factors, which the exceptions to the -rule we must suppose do not. But on this interpretation the fundamental -factor in the determination of the coloration is the action of light. -Selection imposes a bar only to certain types of coloration. - -Some Butterflies and Moths, it has just been hinted, when resting -exhibit bright colours. Our “Swallow-tail” the under surface of the -wings is as brightly tinted as the upper. Among the Moths may be cited -many of the gorgeous Atlas Moths, the Hawk Moths, the beautiful Indian -_Dysphania militaris_—wherein the whole of the exposed surface is of a -beautiful and vivid violet and yellow—and the tropical members of the -Burnet Moths, belonging to the family Syntomidæ. In all these cases it -is not the under but the upper surface of the fore-wings which has thus -departed from the usual rule of the tribe. Not the least remarkable -feature of these insects is the fact that while the Atlas -and Hawk Moths are crepuscular in habits, the Dysphanias and Syntomids -and Burnet Moths are diurnal, and revel in the sunlight. - -To revert for a moment to the factors to which these and other bright -and often conspicuous hues are due. That all highly-coloured animals -are descendants of dull-coloured ancestors there can be no room for -doubt. The vivid tints they now display are to be regarded as due to -some change in the metabolism, some clarifying process of the organism -whereby the various pigments became segregated, concentrated and -intensified. But many of the most vivid hues are not due to pigment at -all, but to changes in the surface structure of the coloured areas. -Such are the wonderful metallic colours which all kinds of animals -display. The iridescence is due to the breaking up of the light by -reflection from finely-grooved surfaces. - -Whatever their nature, one still asks what is their origin, what -brought them into being. They cannot be regarded simply as adaptations -which have arisen to meet the demands of the environment, as are the -structural peculiarities of the skeleton for example; for in this -case both sexes, and all stages of growth, should display the same -hues, and this is rarely the case. Furthermore, we should not in this -case be left with a vast assemblage of forms which certainly cannot -be “pigeon-holed” as to the nature of their coloration. Such, for -example, as the marine types of birds. - -The metallic and iridescent tints to which reference has just been -made, occur among animals to which they can be of but doubtful value, -as in the Golden Mole, for example, or the inside of the Oyster shell. -Their existence in such places well illustrates what we may call the -fortuitous, or apparently fortuitous, beginning of colour of whatever -kind, regarded from an analytical point of view. That is to say, we -are not concerned with the fact that animals are coloured—that is -inseparable from their existence; but with why this coloration should, -in some cases, assume so conspicuous a brilliancy and vividness—a -coloration varying in its character with every species, but apparently -unchanging among the individuals of that species. - -No answer to this, likely to find general acceptance, seems to be -forthcoming at present. But it is significant to remark that all -coloration of the kind now under consideration has its origin, as -have most other structural characters, in the male. It is as true of -coloration as of, say, skeletal characters. One turns to the male -for what is new in the history of a species, to the female and young -for indications of past history. It is equally true that in their -coloration one finds the same sequence of development—the male first, -then the female, then the young, till both sexes, and all stages, are -once more alike in hue. And this rule seems to apply to coloration of -all kinds—Protective—Warning—Epigamic. - -The tendency to develop brilliant colours is associated with some -physiological diathesis with which we are not yet acquainted. But -once having started, this tendency gathers force with each succeeding -generation and continues to exhibit an almost kaleidoscopic capacity -for change, unless, and until, checked by Natural Selection, whereby -its further progress in any given direction may be barred, or some -other element or aspect of the coloration may be introduced. - -Given this controlling factor, all the various types of coloration -would seem to be interpretable. By almost common consent, however, -the resplendent coloration of the males among many species of birds, -a coloration often apparent only during the reproductive period, and -the more conspicuous ornamentation of the males of many other groups, -higher and lower in the scale of organization, are supposed to be -governed by an entirely different factor—female choice, or preference. -The exercise of this, it is contended, has gone on for countless -generations, and the tendency has ever been to heighten the intensity -of the ornament by the rejection of the less favoured suitors in favour -of their more resplendent rivals. Birds and Butterflies alike are -supposed to be swayed by the same irresistible desire to mate, and mate -only with what we may call the smartest and best—groomed of their many -suitors; and these, of course, being the most vigorous, most virile, -sustain the stamina of the race and so attain Nature’s end. - -So long as attention was focused alone, or mainly, on birds conspicuous -for the highly ornamental character of their plumage, this theory -seemed reasonable and probable enough, for one may admit in their -courtships an element, at least, of intelligence and keenness of -perception. But it has now been abundantly demonstrated that the -animated displays so characteristic of these gaily-bedecked gallants, -are enacted with no less persistence and vim by species which exhibit -a Quaker-like soberness of dress. Thus, then, the champions of the -Sexual Selection theory have been dazzled by the tinsel, and have -missed the essential elements—the physical and psychological side of -the display—the contortions, prancings, and so on, and they have missed -the even more important element, the preliminary struggle for territory. - -In this new light, the gaily-bedizened individuals of the Insect world -may be surveyed afresh. The explanation of such of their features as -are commonly attributed to Sexual Selection in terms of female choice, -whereby only the most favoured from among a crowd of suitors could -hope to succeed, may now be replaced by that which obtains also in -the case of the higher animals. It seems to fit the facts better. One -cannot understand, for example, how, on the interpretation of Sexual -Selection, the extraordinary disparity in numbers between the sexes -of some species of Butterflies came about. Thus in that marvellously -beautiful genus Ornithoptera there is one species (_O. brookiana_) in -which the females are excessively rare; so much so that the collector -Kunstler could only obtain fifteen females to one thousand males. -Though the males, among the Butterflies, are commonly much more -numerous than the females, the disparity is rarely so great as with -this species; but there are many in which the proportion of males to -females is as fifty to one. As with the higher vertebrates selection -affords no explanation of this curious disproportion. Though according -to Weismann it fulfills “the first postulate in ‘Sexual Selection’ -namely, that there be an unequal number of individuals in the two -sexes.” But Sexual Selection here has a little over-reached itself, -for surely one hundred suitors seems an embarrassing number for an -inexperienced female to have to choose from! To say nothing of the -ninety-nine males doomed to perish without leaving offspring. - -That the beauty of colour and form which the Lepidoptera, and -especially the diurnal Lepidoptera, or Butterflies, exhibit is due to -the choice by the females—albeit an unconscious choice—of the most -resplendent of her suitors, that is, in other words, that she yields at -last to the most ravishing member of the crowd—there is no evidence to -show. There would seem to be no possibility of a differential selection -from among a number of males, for there is no “display” comparable to -that, say, of birds. And what is more, it is unlikely that, if there -were, she would find anything to choose between them, for the range -of variation in, say, one hundred males of any given species is very -slight. Finally we have no trustworthy evidence to show that the eyes -of Butterflies and Moths are sufficiently good to enable them to make -nice distinctions between slightly different males. We have no evidence -that the eyes of Insects are capable of discriminating the details of -the often intricate patterns which their own wings, and those of their -suitors, exhibit. - -In the matter of “Secondary Sexual Characters,” indeed, the Lepidoptera -exhibit very little difference between the sexes. As a rule the females -are larger, often strikingly so, but in the matter of coloration they -show far less disparity. But there are exceptions to every rule. A -striking illustration of this is afforded by the genus Ornithoptera. -The butterflies of this superb group are of huge size, and the females -are larger than their consorts, and commonly are extremely different -therefrom both in coloration and habits. In _Ornithoptera paradisea_ -this disparity attains its maximum. The female, remarks Mr. David -Sharp, “is a large, sombre creature of black, white and grey colours, -but the male is brilliant with gold and green, and is made additionally -remarkable by a long tail of unusual form on each wing.” But a glance -at the two sexes will show that the female, though less gorgeously -arrayed, still disports a livery which is of a highly specialized -or elaborated character. How are we to account for her differences -in shape, size and coloration on the older interpretation of Sexual -Selection? The perceptual powers, the mentality, of a Butterfly are -surely of a far lower grade than those of a bird, or even a fish. Here, -therefore, we cannot attribute the same possibilities of response to -form and colour which we can ascribe with tolerable safety to the -vertebrates. Yet the Sexual Selection theory as generally understood -demands this. - -So far so good. And now as to the part played by Sexual Selection among -the Lepidoptera. Darwin, in formulating this, found its application -to the Lepidoptera a very disconcerting problem, being naturally -disposed to regard the extraordinary wealth of colour which these -insects exhibit as the outcome of a process of female selection, in -every way comparable to that which he held to obtain among the birds. -He did not postulate a conscious, deliberate, selection; but a final -abandonment on the part of the female to the male which, by his beauty -and demonstrativeness, pleased her most. He assumed that at this -critical time she would always be surrounded by rival suitors, offering -varying if slight degrees of difference: and, indeed, in many cases she -is thus surrounded. He remarks, in discussing the case of Butterflies: -“The males sometimes fight together in rivalry, and many may be seen -pursuing or crowding round the same female.” But in the case of the -Silk-moths—and here is another illustration of the merciless criticism -to which he submitted his own theories—he remarks: “The females appear -not to evince the least choice in regard to their partners.” This fact, -which is certainly true in the case of both Butterflies and Moths, and -these gorgeous hues, disconcerted him, as is shown in the passage: -“Unless the female prefer one male to another, the pairing must be left -to mere chance, and this does not appear probable.” - -The facts which have come to light in regard to the “Courtship” of -Butterflies since Darwin wrote are meagre enough, but such as have been -recorded give no support to the supposition that the females are really -influenced by, or even perceive the colours of, their mates. Just on -five-and-twenty years ago the naturalist Skertchly published some -observations on the Courtship of that magnificent Bornean Butterfly -_Ornithoptera brookiana_. He one day came on a male sipping honey from -the flowers of a tree, vibrating its wings with the rapidity of a -Hawk-moth, and the vivid green of the wings flashing in the sunlight, -though the crimson areas thereof were invisible. The female came “and -did all the wooing.” They circled about in flight with the female -above and somewhat behind, so that she could see, we are told, the -emerald markings; but there was no real evidence here that she was -really influenced by his coloration, and if this really were the case -then the coloration of the female equally demands an explanation, -for this, though less gorgeous than that of the male, is far from a -primitive type; on the contrary, it is of a highly differentiated -character. Furthermore, in this genus, as has already been remarked, -the males outnumber the females by, roughly, one hundred to one. Again, -Moseley, the naturalist on the Memorable Voyage of the Challenger in -1872, when in the Aru Islands, was once “lucky enough to find a flock -of about a dozen males fluttering round and mobbing a single female. -They were then hovering slowly, quite close to the ground, and were -easily caught.” But he was by no means convinced that any choice was -exerted. And he suggests “a series of experiments, in which, in the -case of highly-coloured and decorated Butterflies, the colours should -be rubbed off the wings of a few among a number of males, or painted -over of a black or brown colour. It might be tested whether the females -would always prefer the highly-coloured ones.” Such experiments are -foredoomed to failure, for the removal of the scales would remove the -only source of communication between the sexes. - -Wallace, always a strenuous opponent of the Sexual Selection theory, -found in the behaviour of Butterflies and Moths when mate-hunting a -particularly powerful countervailing weapon. He assumes that Darwin -postulated a conscious selection on the part of the female, and with -some show of reason, though it is probable that Wallace was mistaken -in this. “The weakness of the evidence for conscious selection among -these insects,” he remarks, “is so palpable, that Mr. Darwin is obliged -to supplement it by the singularly inconclusive argument, ‘Unless -the female prefer one male to another the pairing must be left to -mere chance, and this does not appear probable’ But he has just said, -‘The males sometimes fight together in rivalry, and many may be seen -pursuing or crowding round the same female’ While in the case of the -Silk-moths—‘the females appear not to evince the least choice in -regard to their partners.’ Surely the plain inference from all this -is, that the males fight and struggle for the almost passive female, -and that the most vigorous and energetic, the strongest-winged or the -most persevering wins her. How can there be chance in this? Natural -Selection would here act, as in birds, in perpetuating the strongest -and most vigorous males; and as these would usually be the more highly -coloured of their race, the same results would be produced as regards -the intensification and variation of colour in the one case as the -other.” - -Commenting on Darwin’s interpretation of those cases wherein the -females are more brilliantly coloured than the males, he insists that -on his (Darwin’s) theory “throughout the whole animal kingdom the males -are usually so ardent that they will accept any female, while the -females are coy, and choose the handsomest males, whence it is believed -the general brilliancy of males as compared with females has arisen.” - -“Mr. Darwin admits,” he continues, “that these bright colours have -been acquired for protection [because they resemble those of species -which from their disagreeable taste are avoided by birds and other -insect-eating enemies]; but as there is no apparent cause for the -strict limitation of the colour to the female, he believes that it -has been kept down in the male by its being unattractive to her. This -appears to me to be a supposition opposed to the whole theory of -Sexual Selection itself. For this theory is, that minute variations -of colour in the male are attractive to the female, have always been -selected, and that thus the brilliant male colours have been produced. -But in this case he thinks that the female Butterfly had a constant -aversion to every trace of colour, even when we must suppose it was -constantly recurring during the successive variations which resulted -in such a marvellous change in herself. But the case admits of a -much more simple interpretation. For if we consider the fact that -the females frequent the forests where the Heliconidæ abound [the -distasteful species already referred to] while the males fly much in -the open and assemble in great numbers with other white and yellow -Butterflies on the banks of rivers, may it not be possible that the -appearance of orange-stripes or patches would be as injurious to the -male as it was useful to the female, by making him a more easy mark -for insectivorous birds among his white companions? This seems a more -probable supposition than the altogether hypothetical choice of the -female, sometimes exercised in favour of, and sometimes against, every -new variety of colour in her partner.” - -Wallace’s arguments are not so crushing as he supposed them to be, -and they contribute nothing towards the solution of the problem to be -faced. But if colour played the part which Darwin believed, and colour -alone be concerned, it is curious that the males should recognize -their mates in a guise so unlike their own. How is it that they do not -pass them by as members of the totally different distasteful species? -Whenever, indeed, the female is more or less brightly liveried than -the male, how do the sexes recognize one another, and how, when they -live in environments so different as those referred to by Wallace, -do they find one another when possessed by the insistent demands of -the “sex-hunger” which is the all-essential stimulant to secure the -continuation of the race? - -The factors which assure the satisfaction of this hunger differ in -some important features from those which obtain among the higher -animals—birds, for example. In the first place there is no necessity to -find and hold territory, which is an imperative necessity where there -are eggs to be brooded and young to be fed. In the second, the males, -as has just been remarked, must search for the females, often indeed, -in the case of many Moths, because they are wingless. - -This search is conducted by the sense of smell. This fact, familiar -enough to-day to the entomologist and the student of Evolution, -was unknown to the earlier naturalists. Neither Darwin nor Wallace -suspected it. It would have been wonderful if they had, for there is -nothing in the general appearance of these insects which suggests an -organ of smell, nor is there anything in the structure of the nervous -system which would indicate this subtle sense. During recent years, -however, the number of workers engaged on the investigation of the -senses of animals has increased immensely, and great strides have been -made in perfecting instruments of research. To the efforts of these -workers we owe the discovery of the seat of the scent-detecting organs -and the source of the scent. The former are furnished by the antennæ, -which lodge also the senses of taste and touch. - -Among the Lepidoptera these constitute important secondary sexual -characters, the antennæ, among the Moths at any rate, presenting -striking differences in male and female. The scent-producing organs -are very elusive structures, and so far have been definitely traced, -among Butterflies, only in the males, where they are formed by certain -peculiarly modified scales known as “androconia.” They may be either -irregularly scattered over the wing, or may form complex structures. -Sometimes they are arranged in the form of brightly-coloured, -bristle-like tufts on the hind-wings, sometimes in a fringe along the -edge of the hind-wing. In some of the Moths they are arranged to form -a thick, glistening white felt, which fills a folded-over portion -of the edge of the hind-wing, and in many cases “the perfume can be -retained,” Weismann remarks, “and then, by a sudden turning out of the -wing-fold, be allowed to stream forth.” In the Ghost-moth (_Hepialus -humuli_), the hind-legs of the male have become pressed into service -and have become transformed into scent-bottles, since they are swollen -and filled with glands for the manufacture of odorous matter. - -The naturalist Fritz Müller discovered the fact that some of the -Butterflies which haunted his Brazilian garden exhaled a flower-like -fragrance. Anyone can test this curious trait for himself who will take -the trouble to brush his finger over the wing of a newly-caught male -Garden-White Butterfly (_Pieris napi_). The white powder which will -adhere to the finger will be found to be made up of the wing-scales, -which will exhale a delicate perfume of lemon or balsam! Among the -Moths the strong odour of musk is exhaled by the Convolvulus Hawk-moth -(_Sphinx convolvuli_). - -It is, however, only in the males that these odours can be detected, -and, though palpable enough to human nostrils, their power of -diffusion is apparently extremely limited. They would seem to serve as -aphrodisiacs for the stimulation of the female, and, as a consequence, -there is no need that they should start into activity until the male -has arrived at the immediate neighbourhood of his prospective mate. - -[Illustration: Plate 31. - -BRIGHT COLOURS WHICH CANNOT UK ATTRIBUTED TO “SEXUAL SELECTION.” - -1. Eyed Hawk-moth, under the influence of excitement. - -2, A Butterfly, Zeuxidia horsfieldi, Feld, showing tufts of -scent-diffusing scales on the hind-wings. - -Face page 200.] - -With the females of the Moths, however, matters are otherwise. For the -most part Moths are nocturnal, and hence could not distinguish one -another when on the search for mates, and in many species the females -are wingless, and consequently are unable to move from the immediate -neighbourhood in which they emerged from the pupal stage. In either -case some means of informing the males of the presence of females is -an imperative necessity for the continuation of the race. This is -provided by means of a subtle odour exhaled by the females which, -though imperceptible to human nostrils, must possess an extraordinarily -penetrating power. Weismann gives an instance of this in the case of -the nocturnal Eyed Hawk-moth (_Smerinthus ocellatus_). He placed some -females, without any special intention, in a covered vessel near an -open window. “The very next morning several males had gathered, and -were sitting on the window-sill, or on the wall of the room close to -the vessel, and by continuing the experiment I caught, in the course of -nine nights, no fewer than forty-two males of this species, which I had -never believed to be so numerous in the gardens of the town....” To -this power of exhaling odours we may attribute the wingless condition -of many Moths, for otherwise the loss of flight would have brought -about extinction long before any perceptible reduction in the wings had -taken place. The odour which such prisoners emit seems to possess an -irresistible attractiveness, and this fact is commonly taken advantage -of by entomologists. The Common Vapourer Moth (_Orgyia antiqua_) -affords a good illustration of this. The female is wingless, and little -more than a pouch for eggs, but in certain seasons it is very abundant, -even in the midst of London. That experienced entomologist Prof. Selwyn -Image, in a letter to my friend Mr. John Cooke, remarks, on this theme, -that the Caterpillars may be seen crawling by hundreds in and around -the squares, while the males may be seen flying up and down New Oxford -Street or Tottenham Court Road. If a virgin female be put in a box -placed outside the window, within a very short space of time, often not -more than a few minutes, several males will be fluttering round her. -This device for attracting males is commonly known as “assembling.” - -More striking is the case of the Oak-eggar Moth (_Lasiocampa quercus_). -Mr. Richard South, in his most useful “Moths of the British Isles,” -relates that on one occasion he had a number of pupæ in a cage in -a cottage on the edge of a moor near Lynton, North Devon, and these -attracted quite a number of males into the room containing the precious -casket, and he was enabled to capture several. The next day he placed a -female which had meanwhile emerged, in a “roomy chip-box, and carried -it, in a satchel, to the moor, where it was placed on the ground; the -males began to arrive soon afterwards, and some fine examples were -secured.” But the sequel is even more remarkable; for, he remarks: -“Although the female was taken on the moor only on one occasion, that -satchel continued to be an object of interest to the male Eggars -for several days afterwards.” That this scent is capable of being -transferred to foreign objects, and of retaining its power for several -days, is a striking proof of its pungency, yet it is quite impalpable -to human nostrils! The Kentish Glory Moth (_Endromis versicolor_) -affords yet another instance of this curious attraction by scent, the -effectiveness of which is not even lessened by exhalations of the -human body, for if a virgin female be placed in a box, and this be -placed in one’s pocket, the males will often swarm round one and even -endeavour to gain access to the box. In all such cases the females, -even when capable of flight—the female Vapourer is wingless—never fly -until after impregnation has taken place. Hence males with defective -scent—detecting powers inevitably fail to leave offspring. - -Selection, then, here lies between males of the most active -scent-detecting powers, and not between those of the most brilliant -colours. Nevertheless, both males and females—where the females -are winged—exhibit a remarkably beautiful coloration, and this is -especially true of the Kentish Glory, wherein both sexes wear a -resplendent dress. That of the male—which is much smaller than the -female—differs in that the fore-wings are darker, but bear the same -pattern as in the female, while the hind-wings are chestnut-red instead -of cream colour as in the female. If this scent-factor has replaced -colour as an inciting agent to pairing, then these Moths should be of -sombre hues. That such is not the case seems sufficient to show that -the colour is not due to Sexual Selection, for it is highly improbable -that scent and colour are both of equal importance, and this being so, -one would expect to find the negligible factor eliminated. - -The existence, then, of bright colours in this and other species -in like case, seems to show that it has nothing to do with Sexual -Selection, directly at any rate. The males having assembled, their -presence is probably communicated to the female by the characteristic -male odour, which is never of the same penetrating quality as that -of the female. There is no need that it should possess this, for the -females never seek their mates. The successful male, where several -rivals are competing, is probably not simply the strongest, but he -who also disperses the right odour necessary to provoke the pairing -response. These illustrations furnished by the scent-hunting, -scent-dispersing males and females are of the highest importance -to students of the Sexual Selection theory, for they seem to show -conclusively that coloration plays at any rate but a minor part -therein. The importance of the scent-detecting organs is shown in -the very different types of antennæ which obtain between male and -female Moths, those of the male taking the form of huge feather-like -structures, as in some Saturniidæ, and far exceeding those of the -female in size. - -The methods of pairing which obtain among Butterflies and Moths, it is -not surprising to find, are very different; for whereas in the former -it takes place on the wing, in the latter the female is always in a -resting position. Where the females are winged, long flights are often -taken for the purpose of depositing and distributing the eggs: the -flightless forms make no such excursions. A few, as in the case of some -of the _Psychidæ_ are not only wingless, but limbless and maggot-like. -They never leave the chrysalis case, but deposit their eggs inside it. -Though there is undoubtedly much that is wonderful about the mating -of these scent-distributing species, the history of the Moths of the -genus Acentrophus is more wonderful and more mysterious still. For -the females are aquatic. The males may sometimes be found in crowds -fluttering over the surface of large but shallow sheets of water. The -females, which are wingless, come to the surface and, like sirens, draw -the males under water, where coupling takes place; after which they -probably immediately die. But how do they discover their submerged -mates? The escape from the water of any odour which the females may -possess seems well nigh impossible. - -Whether display, such as birds appear to delight in ever takes place -among the Lepidoptera seems doubtful Nevertheless, something closely -akin thereto seems to have been found in the case of certain species -of Butterflies (_Heliconius melpomene_ and _H. rhea_), which have been -seen dancing in the air like gnats, and when some of them withdrew -others took their places. Again, having regard to the fact that birds, -when alarmed or excited, will perform the display which is more or less -characteristic of periods of sexual excitement, it is possible that the -position of alarm assumed by some of the Hawk Moths may also be used in -Courtship (Fig. 1, Plate 31). But we have no evidence on this point, -and from the part played by scent in the mating of Butterflies it seems -improbable that such displays take place. - -A serious attempt to test the Sexual Selection theory by experiment—to -test the extent, if any, of female choice in mating—was made some -years ago by Mayer, an American naturalist, on the large Bombycid Moth -(_Callosamia promethea_). This species exhibits striking dissimilarity -between the sexes in regard to colour and pattern. “The females,” -remarks Professor Kellog, “are reddish brown in ground colour, while -the males are blackish, and in the two sexes the pattern is distinctly -different....” Mayer took four hundred and forty-nine pupæ, in -cocoons, of this moth and endeavoured to discover, first of all, -whether the males found the females by sight or smell. Enclosing -females in jars, some of which were covered and some of which were -uncovered, he found that males paid no attention to females enclosed in -transparent jars so closed as to prevent the escape of odours, while -such as were enclosed in boxes or wrapped in cotton-wool, so as to be -invisible, but yet capable of exhaling odour, were besieged by males. -To locate the organs of scent in the female he cut off the abdomen of -several and placed the abdomens and their late owners at some distance -apart. Males came to the abdomens and not to the thorax and wings. -Males whose antennæ were covered with shellac, photographic paste, -glue, paraffin, etc., showed no response to the female exhalations, -until the covering medium was removed. - -Mayer next tested the selective action of the females. He began by -removing their wings and affixing to the stumps the wings of males. -The males mated with the females quite as readily as under normal -conditions, though the most conspicuous female characters had been -exchanged for those of the male. After this he affixed female wings -upon the males, but mating took place as usual. The females did not -seem to detect anything unusual in their suitors, nor did normal -males attempt to pair with males bearing female wings. Later he tried -the experiment of dyeing the wings of three hundred males scarlet -or green, and matched these against three hundred which were left -untouched. The disguised, dyed males succeeded in pairing as easily -as their normally-coloured brethren. The females exhibited no choice -whatever. Hence, then, we have further reason to believe that with -the Lepidoptera scent, not sight, is the channel by which mates are -found. So far as the evidence goes, it seems to show conclusively that -in all that concerns sexual relationships, scent is the guiding and -determining factor. By scent the females attract the males, and by -scent of another kind the males sharpen the procreative appetites of -the females. - -If the interpretation adopted in these pages is correct, these -manifestations and emanations of colour and scent are readily -accounted for; for they are manifestations of inherent growth changes -which, having started, are free to go on increasing in amplitude -unless, and until, checked by natural selection. There is nothing -unreasonable or improbable in this interpretation; on the contrary, -it embraces also many other features hitherto ignored, but no less -demanding an explanation. Such, for example, as the infinite variety -of form and sculpture which the scales of the wings and the eggs -display. These are details visible only by the aid of the microscope, -but they demand explanation as much as the more obvious characters. -Moreover they have the advantage of belonging to a set of characters -which cannot in any way influence the choice, if choice there be, in -the selection of mates, nor are they of a nature likely to affect the -results of the struggle for existence. Of these characters, then—the -sculpturing of the egg-shell and of the scales, the “nervation” of -the wings, and coloration—we can say no more than that they are -idiosyncrasies of growth, free to develop in any direction unless, and -until, checked by natural selection, which will speedily eliminate -disharmonies with the environment. - - - - -CHAPTER XI - -BEETLES THAT “BLUFF” - - -The Coloration, and other Forms of Ornament in Beetles, and the -Significance thereof in regard to the Sexual Selection Theory—The -Courtship of Grasshoppers and their Kin—The Remarkable Ears of Locusts -and Grasshoppers—The Field-cricket and the Katydid as Troubadours—The -Wonderful Performances of the Cicadas—The Duels of Long-horned -Locusts—Dragon-flies—The May-flies’ “Dance of Death”—The Jaws of -the Giant Alder-fly and their Strange Use—Some Curious Facts about -Stone-flies. - -In these pages it is contended that neither brilliant coloration nor -any other form of ornamentation is to be ascribed to the direct action -of “Sexual Selection.” That is to say, such conspicuous features have -not been dependent on the action of female choice for their survival -and development, but are rather the “expression points” of the -internal, inherent growth variations, which, not being inimical to the -welfare of the species, have been free to pursue their development in -any direction which apparent chance may dictate. - -The Butterflies and Moths well illustrate this in regard to coloration, -for scent, not colour, would seem to be their principal source of -information as to the outer world. The Beetles are no less instructive; -for these creatures, though they contain numerous highly-coloured -and some exquisitely beautiful species, are more remarkable for -their bizarre shapes, and it seems impossible to regard these as the -products of sexual selection. Yet this is the interpretation of their -origin which, in the judgment of Darwin, we must adopt. He evidently -had misgivings as to the correctness of this view; but it must be -remembered that in reviewing the facts relating to these lower orders -of Creation he was biased by the evidence which he had brought together -in regard to the behaviour of the higher groups under the stimulus of -sexual emotion. Convinced that female choice obtained here, he was -but following the logical result of such conclusions in postulating -the same factor wherever it could conceivably be applied. The most -formidable critic of the Darwinian theory of Sexual Selection was -Darwin himself. The dominant ambition in all his work was to explain -his facts, not to establish his theory; and he was convinced that his -theory of Sexual Selection did achieve that end; though there were -cases where the evidence he was analysing seemed less clear than in -others. That the Beetles presented difficulties is evident from his -comments thereon. He was puzzled by the vivid coloration which some -species present. “They may serve,” he remarks, “as a warning or means -of recognition ... as with Beetles the colours of the two sexes are -generally alike, we have no evidence that they have been gained through -sexual selection; but this is at least possible, for they may have been -developed in one sex and then transferred to the other; and this view -is even in some degree probable in those groups which possess other -well-marked secondary sexual characters.... - -“Some Longicorns, especially certain Prionidæ, offer an exception to -the rule that the sexes of Beetles do not differ in colour. Most of -these insects are large and splendidly coloured. The males of the genus -Pyrodes ... are generally redder but rather duller than the females, -the latter being coloured of a more or less splendid golden-green. On -the other hand, in one species the male is golden-green, the female -being tinted with red and purple. In the genus Esmeralda the sexes -differ so greatly in colour that they have been ranked as distinct -species: in one species both are of a beautiful shining green, but -the male has a red thorax. On the whole, as far as I could judge, the -females of those Prionidæ in which the sexes differ are coloured more -richly than the males, and this does not accord with the common rule in -regard to colour when acquired through sexual selection.” - -While there is nothing very remarkable in the two sexes being coloured -alike, it is certainly strange to find the female more brilliantly -coloured than the male. And this because among the higher vertebrates, -as among the birds, the female exceeds in brilliance only where she -also plays the rôle of wooer instead of wooed; leaving to the male the -whole responsibility of rearing the family. With the Beetles the family -has to rear itself, parental care being limited to the right disposal -of the eggs. By some change in the character of the germ-plasm the -females may have, in these cases, acquired more “maleness,” more of the -qualities which are answerable for the secondary sexual characters of -the male, or, what seems rather to be the case here, a result like that -which has been reached in certain of the Pigeons and Parrots has been -arrived at. That is to say, the tendency to intensification of pigment -in the female struck out a new line, instead of following that of the -male. This rather rare form of sexual dimorphism is also met with, it -will be remembered, among the Butterflies and Moths. - -While brilliant colour is the more usual form of ornament among the -Beetles, there are many species wherein the males have developed -enormous horns, or have greatly exaggerated the length of the jaws; -and these outgrowths give the impression of a formidable armature, -but so far as the evidence goes this is by no means the case. They -must therefore be relegated to the category of “ornaments,” though the -term “excrescences” would more fittingly apply to them, for they are -“ornaments” only from a human standpoint. At any rate, there is no -evidence whatever that they serve to enhance their possessors in the -eyes of the females. - -In relation to the Sexual Selection theory these excrescences are -of quite exceptional interest, for they throw a strong light on the -meaning of ornament, such as obtains among birds, which seem to show a -consciousness of its existence and effectiveness. Darwin argued from -the birds to the Beetles. Convinced that the gorgeous crests and trains -and vivid colours were appreciated by the females of the former, he was -impelled to believe that the ornaments of the latter had developed in -like case by the fostering influences of the females. Similarly, from -the evidence as to the use of horns in the case of mammals, and spurs -in the case of birds, he was induced to believe that the horn-like -outgrowths of Beetles had been attained by like influences. But in -both kinds of cases, he could only infer their action, for he could -discover no really decisive instances of conquest either by display or -by battle, such as he was able to produce in the case of the higher -animals. Had chance directed his attention in the beginning either -to the Warblers among the birds, or the beetles among the insects, -his interpretation of the action of sexual selection, it is more than -probable, would have been materially different from that developed in -the “Descent of Man.” No additions of any importance have been added to -the facts he so laboriously collected. - -As touching the “horns” it should be remarked that these may arise -either from the head or from the thorax, or from both, and sometimes -even from the under surface of the body. - -One of the most remarkable instances of these singular outgrowths is -that of the Hercules Beetle (_Dynastes hercules_) of the West Indies -and tropical America. Herein the roof of the head is prolonged into -a great upturned beam bearing tooth-like prominences, and the top -of this is opposed to a still more massive beam, whose base covers -the whole roof of the thorax, and whose tip extends far beyond that -projecting from the head. A pair of “teeth” point downwards from the -middle of this beam, whose under surface is thickly covered with short -chestnut-coloured hairs forming a brush-like surface. In another, -_Copris isidis_, the head bears two short, rhinoceros-like horns, -and the thorax a short, triangular overhanging ledge: in _Phanœus -jaunus_ there is a single horn on the head, and the thorax bears -two short, forwardly-projecting blades, one on each side; while in -_Onthophagus rangifer_—the Reindeer Beetle—the head bears a pair of -horns curiously like the antlers of a deer. One might cite many such -instances, all varying in detail, but these will suffice. - -Darwin, in commenting on these structures, remarked: “The -extraordinary size of the horns and their widely different structure -in closely-allied forms indicate that they have been formed for -some purpose; but their excessive variability in the males of the -same species leads to the inference that this purpose cannot be of a -definite nature. The horns do not show marks of friction, as if used -for any ordinary work. Some authors suppose that as the males wander -about much more than the females, they require horns as a defence -against their enemies; but as the horns are often blunt, they do not -seem well adapted for defence. The most obvious conjecture is that -they are used by the males for fighting together, but the males have -never been observed to fight, nor could Mr. Bates, after a careful -examination of numerous species, find any sufficient evidence, in their -mutilated or broken condition, of their having been thus used. If -the males had been habitual fighters, the size of their bodies would -probably have been increased through sexual selection, so as to have -exceeded that of the females; but Mr. Bates, after comparing the two -sexes in above a hundred species of the Copridæ, did not find any -marked difference in this respect amongst well-developed individuals. -In Lethrus, moreover, a Beetle belonging to the same great division of -the Lamellicorns, the males are known to fight, but are not provided -with horns, though their mandibles are much larger than those of the -female. - -“The conclusion that horns have been acquired as ornaments is that -which best agrees with the fact of their having been so immensely, -yet not fixedly, developed—as shown by their extreme variability in -the same species This view will at first appear extremely improbable, -but we shall ... find with many animals standing much higher in the -scale, namely, fishes, amphibians, reptiles and birds, that various -kinds of crests, knobs, horns and combs have been developed apparently -for this sole purpose.” - -The assumption that these “animals standing much higher in the scale” -owe their weapons to the selective action of the females forms the crux -of the whole Sexual Selection theory in regard to the significance -of ornament. The evidence that the intensification of pigment and -the eccentricities of growth in the shape of crests and frills have -a fascinating effect on the female is more than under suspicion; it -is discredited by the facts which have come to light in regard to -behaviour during the periods of sexual exaltation. And there is a -growing conviction that this is so. No better proof could be found that -“ornaments” can, and do, exist in spite of, rather than because of, the -action of “sexual selection.” They are the accidents of this selection, -not a part of its machinery. - -Incipient horns are found in not a few cases among the females of -these insects, while in others, as in the case of the Reindeer Beetle, -they are almost as well developed as in the males. This is what one -would expect to find if these outgrowths were the result of inherent -variations restrained as to their size by natural selection, which -eliminates only when this growth penalizes, by increasing the struggle -for existence. - -As to the actual behaviour of Beetles when sexually excited but very -little information is obtainable; but there are records of species the -males of which fight with rivals for the possession of females. Wallace -saw two males of _Leptorhynchus augustatus_, a Beetle with no name in -common speech and a long beak, “fighting for a female, who stood close -by busy at her boring. They pushed at each other with their rostra, and -clawed and thumped in the greatest rage.” The smaller male, however, -“soon ran away, acknowledging himself vanquished.” In this case, it -is to be noted, the combatants lacked weapons. With the Stag Beetle it -is otherwise, and this species is said to engage in fierce conflicts. -Darwin cites an instance where two males were enclosed with one female -in a box, when the larger severely pinched the smaller one, until he -resigned his pretensions. This being so, it is curious to find that the -female, which makes no display of pugnacity, has the stronger jaws. -The fact that there are so few records of fighting among male Beetles, -and the absence of injury to the highly-polished surfaces of the horns -or jaws where these are conspicuously large, seem to indicate that at -most no more than a semblance of fighting ever takes place. In a North -American Stag Beetle (_Lucanus elaphus_) the jaws, which are greatly -developed, are used, Darwin tells us, for seizing the female, but they -do not appear to be employed for this purpose in any other species. -It might be held that they play the part of terrifying agents, as the -eye-spots of Caterpillars and adult Lepidoptera are believed to do. -At any rate, they seem to be so used in the case of a Beetle of South -Chile (_Chiasmognathus grantii_) wherein the jaws are of great size -and have their inner edges toothed. When threatened “he faces round, -opens his great jaws, and at the same time stridulates freely.” But -this parade of force is evidently no more than “bluffing,” for Darwin, -who describes this behaviour, remarks, “the mandibles were not strong -enough to pinch my finger so as to cause actual pain.” In the female, -it may be remarked, the jaws are quite small. - -That too much stress has been laid upon the significance of the -enlarged jaws and other hypertrophied developments in the Beetles -seems to be shown by the case of certain carnivorous Beetles, of which -one species (_Taphroderes distortus_) may serve as an example. Herein -the left jaw takes the form of a long, crooked strap-shaped outgrowth, -whose purpose cannot even be conjectured. And in this connection one -may cite the case of certain species of Homoptera—Bugs—which occur in -tropical South America. Here, in both sexes, as may be seen in Plate -40, Fig. 4, the neck-shield is produced backwards far beyond the body, -to form a most elaborate superstructure which appears to confound the -most ingenious attempts at interpretation. - -It is to be noted that wherever special structures are necessary for -the performance of specific acts such as are of vital importance to -the well-being of the race, they are developed to perfection: there -is little or no variation in their size, and no doubt as to their -purpose. Thus in many species means are necessary to enable the male -to seize and hold the female during the sexual embrace. In the Water -Beetle of our ponds and ditches (_Dytiscus marginalis_) the male bears -a very remarkable sucker on each fore-leg, the adhesive surface of -which, under the microscope, reveals an extraordinary complexity and -wondrous beauty. This sucker forms a very conspicuous “secondary sexual -character,” and is used in embracing his mate, whose back is deeply -grooved to enhance the hold of the suckers. In some species punctures -take the place of grooves. Suckers, like those of _Dytiscus_ are met with -again in a Wasp (_Crabro cribrarius_). In another genus of Beetles -(_Penthe_) cited by Darwin, the antennæ of the male have a few of the -middle joints dilated and their under surfaces furnished with a cushion -of hairs to aid in the sexual embrace. - -Beetles are creatures of solitary habits; how, then, do they find their -mates when, by the insistence of the reproductive desires, they are -driven forth to begin the search? Though we have no direct evidence, it -seems more than probable that, as with the Butterflies and Moths, scent -furnishes their most reliable guide. At any rate, in a large number -of species, as among the Lamellicornia, the antennæ bear leaf-like -plates, which are much more developed in the males, in which they -probably serve as scent-detecting organs. - -In some species stridulating organs occur such as are met with in even -greater perfection among the Crickets and Grasshoppers, and among the -Spiders and Scorpions. That these “musical-boxes” provide a means of -communication between the sexes there can be no doubt, even if, as -some contend, they are commonly used only to frighten enemies. This -purpose may well be the explanation of their presence in the larval -Stag Beetle, for it cannot be claimed that they have any relation to -the acts of courtship at this stage of development. - -Stridulating organs, wherever they are met with, are fashioned on -the same principle. The mechanism for sound-production differs -conspicuously from that which produces the voice in the vertebrates. -For where there are no lungs or breathing apparatus, comparable to -that of birds and beasts, there can be no internal voice-mechanism. -Instead, the skeleton which in these creatures forms the external -surface of the body—that is to say, it encloses the muscles, whereas -in the vertebrates it is internal and overlain by the muscles—produces -the necessary sounds. And this by means of rubbing two opposed surfaces -against one another, one of which is ridged, the other toothed. In -the details both of position and structure a wonderful variety will be -discovered when all the known types are surveyed, and it is possible -that they perform different functions in different groups. - -The Locusts and Grasshoppers are among the finest performers of -these “harpists,” and it would seem that in this group, at any -rate, the music they make is of an erotic character. In one of our -native Grasshoppers (_Stenobothrus melanopterus_) these high-pitched -and somewhat strident notes are produced by rubbing the roughened -inner surface of the hindmost thigh, which forms the base of the -great leaping leg, against one of the libs of the wing-case which -is specially enlarged and has a sharp edge. Thereby the wing is -thrown into a state of vibration and the musical sound produced. The -roughening of the inner surface of the thigh just referred to is -produced by a row of bead-like projections whose appearance under the -microscope is depicted in the adjoining illustration. This apparatus -is well developed in the males, and but feebly, or not at all, in the -females. That the music it produces is appreciated by the performers -and their mates there can be no doubt, for they are provided with -a special apparatus which fulfils the purpose of an ear. In the -short-horned Grasshoppers (_Acridiidæ_) this is placed in the middle -of the body just above the base of the great thigh. It differs in the -details of its construction. In some cases it is formed by a delicate -sheet of membrane surrounded by a rim, in others the membrane may be -slightly depressed, and in some very much so, the rim closing up to -form a broad slit. Such ears, it is to be noted, exist in both sexes, -while the stridulating organs do not. That such sound-producing organs -serve as stimulants to the sexual passions of the females is but a -natural inference. Some authorities, however, regard this as doubtful, -since there are species which appear to lack these stridulating -instruments, though possessing ears. But closer observation will -probably show that these apparently dumb species are not really so, as -Dr. David Sharp, commenting on this fact remarks: “It is well known -that sounds inaudible to some human ears are perfectly distinct to -others. Tyndall, in his work on Sound, has illustrated this by a fact -that is of special interest from our present point of view. ‘Crossing -the Wengern Alp with a friend’ he says, ‘the grass on each side of the -path swarmed with insects which, to me, rent the air with their shrill -chirruping. My friend heard nothing of this, the Insect world lying -beyond his limit of audition!’ If human ears are so different in their -capacity for perceiving vibrations, it of course becomes more than -probable that auditory organs so differently constituted as are those -of insects from our own may hear sounds when the best human ear can -detect nothing audible. On the whole, therefore, it would appear most -probable that the Orthoptera provided with acoustic organs, and which -we consider dumb, are not really so, but produce sounds which we cannot -hear, and do so in some manner unknown to us. If this be the case, it -is probable that these ears are special organs for hearing particular -sounds.” - -[Illustration: Plate 32. - -STRIDULATING ORGANS. ETC. - -1. The stridulating mechanism of the Red Ocypode Crab. - -2. The stridulating apparatus of a Grasshopper—highly magnified. - -3. The head of a Gnat with the compound eyes split up. - -4. The “ear” of a Grasshopper. - -[Face page 218.] - -Certain of the Grasshoppers of Africa, known to entomologists as -Pneumorides, have undergone a most extraordinary transformation -of their bodily shape, as if in response to the demands of these -musical performances. They have entirely lost the power of leaping, -and the abdomen, in the male, has become transformed into a huge, -pellucid, inflated bag or bladder, apparently to serve the purpose -of a resonator, increasing the volume of the sound produced by the -stridulating organ, which consists of a series of ridges placed on -each side. The noise which this mechanism produces is, as might be -supposed, considerable. It is curious to remark that in this group -the females are more vividly coloured than the males. In the case of -one South African species—_Pneumora scutellaris_—this coloration is -so extravagant that she has been said to look as if “got up” for a -fancy-dress ball (Plate 33, Fig. 1). Her ground colour is of a light -green, with pearly-white markings, surrounded by an edging of magenta; -the white areas are very numerous. The face has magenta patches, and -numerous, tiny, pearly-white tubercles, each of which, when placed on a -green part, is surrounded by a ring of mauve. This scheme of coloration -distinguishes her as one of the most remarkably coloured of insects. -But to what are we to attribute these hues? Sexual Selection will -not explain them, and it seems unreasonable to regard them either as -affording a protective or a warning coloration. They may then, perhaps, -be allowed to rank as another instance of unchecked variation in the -direction of vivid colouring, such as has been already described as -occurring both among birds and other animals lower in the scale. - -[Illustration: Plate 33. - -CRICKETS AND MAY FLIES. - -1 and 2 afford illustrations of the excessive development of “ornament -“; fig. 3 of devices for seizing the female; figs. 4 and 5 of -unaccountable differences in the development of wings. - -1. The Pneumatic Cricket (_Pneumora scutellaris_), showing the strange -markings on the female. - -2. The Cleft-footed Burrowing Cricket (_Schizodactylus monstrosus_). - -3. The Giant Alder-fly (_Corydalis crassicornis_), with its huge jaws -for grasping the female. - -4. The Stone-fly (_Perla maxima_), the large-winged Continental form. - -5. Loch Tanna Stone-fly (_Isogenus nubecula_), male, with vistigial -wings. - -[Face page 220.] - -In the Locustidæ the ear is placed on the side of the front leg and -the rim surrounding it may either take an oval shape or close up to -form a slit. The air necessary for the efficient action of the acoustic -apparatus is admitted through a gaping hole in the side of the body, -above the base of the leg, an arrangement not met with among any other -insects. The musical apparatus of these insects differs from that of -the Acridiidæ, for it is formed only by the wing-cases, and not by -the wing case and the leg. One of the wings bears a file on its -inner surface, the other, the right wing, is furnished with a sharp -edge placed on a prominent part of its inner margin. By slightly -tilting the fore-wings, or wing-cases, and vibrating them rapidly, -the edge passes under the file and a musical sound is produced. By -this means one of our native long-horned Grasshoppers (_Locusta -viridissima_) produces a shrill, but not unpleasant, sound, capable of -being sustained continuously for a quarter of an hour. But a species -encountered by Bates during his travels in the Amazons is a much more -efficient performer. Known by the name of Tanana by the natives, it -is so much admired by them for its singing powers that it is kept in -little cages as we keep Canaries. That these organs are of importance -to the species may be gathered from the case of a Bulgarian long-horned -or Green Grasshopper (_Poecilimon affinis_), wherein the wings have so -degenerated as to be useless in flight, but in the male they have been -retained solely as musical instruments. In some species both sexes have -a music-producing apparatus, but as a rule this is present only in the -male. - -That these curious and complex stridulating organs do indeed primarily -act as aphrodisiacs seems to have been clearly demonstrated by the -naturalist Bates, who, in speaking of the European Field Cricket -remarks: “The male has been observed to place himself at the entrance -to his burrow, and stridulate until a female approaches, when the -louder notes are succeeded by a more subdued tone, whilst the -successful musician caresses with his antennæ the mate he has won.” - -Among the most efficient and most celebrated performers of all on these -instruments of percussion are the “Katydids” of North America. The -sounds they produce are said to form the words “Katy-did, O-she-did, -Katy-did-she-did.” The first of these extraordinary concerts is heard -about mid-July; by mid-August they are in full song. By others the -sounds have been likened to those produced by the slow turning of a -child’s rattle, ending in a sudden jerk; and this prolonged rattling, -which is peculiar to the male, is always answered by a single, sharp -“chirp” or “tschick” from one or more females, who produce the sound by -a sudden upward jerk of the wings. - -Pride of place, however, among insect performances of this kind must -surely be awarded to the Cicadas, which are notoriously the noisiest -members of the Insect world, far eclipsing the shrill calls of the -Grasshoppers and even of the Crickets. Darwin remarks that the noise -they made could “be plainly heard on board the _Beagle_ when anchored -off Brazil at a quarter of a mile from the shore; and Captain Hancock -says it can be heard at a distance of a mile. The Greeks formerly kept, -and the Chinese now keep, these insects in cages for the sake of their -song, so that it must be pleasing to the ears of some men.” Only the -males sing, the females being completely dumb, and this prompted the -Greek poet Xenarchus to make the now famous remark, “Happy the Cicadas’ -lives, for they have voiceless wives.” Another naturalist, Riley, says: -“The general noise, on approaching the infested woods, is a combination -of that of a distant threshing-machine and a distant frog-pond.” -Another species, _Tympanoterpes gigas_, also Brazilian, is said to make -a noise equal to the whistle of a locomotive: recalling that of a nest -of young snakes, or young birds, when disturbed—a sort of scream. They -can also produce a chirp like that of a Cricket and a very loud, shrill -screech prolonged for fifteen or twenty seconds, gradually increasing -and decreasing in force. - -Curiously enough, no special auditory organs have yet been discovered, -and it has been suggested that these insects do not hear in our sense -of the word, but feel rhythmical vibrations. But whether the males -“sing in rivalry,” as Dr. David Sharp suggests, is another matter. -The purpose of the “song” in the first place is no doubt intended as -a guide to the females seeking mates. But it is quite conceivable -that the call of one male may stimulate every other male in the -neighbourhood. Darwin, commenting on this aspect of the music, gives a -quotation from Dr. Hertman, who says: “The drums are now ... heard -in all directions. This I believe to be the marital summons from the -males. Standing in thick chestnut sprouts about as high as my head, -where hundreds were around me, I observed the females coming around the -drumming males.... This season a dwarf pear-tree in my garden produced -about fifty larvæ of _Cicada pruinosa_; and I several times noticed -the females alight near a male while he was uttering his clanging -notes.” - -The structures, he remarks, from which these sounds proceed, “must be -ranked amongst the most remarkable voice-organs in the animal kingdom. -They are totally different from the stridulating organs that are found -in many other insects.... Some difference of opinion has existed as to -the manner in which the structures act, but the account given by Carlet -... will, we believe, be found to be essentially correct.” They are -partly thoracic and partly abdominal. On examining a male Cicada there -will be seen, on the under surface, two plates, meeting in the middle -line of the body and overlapping the base of the abdomen. They can -be slightly moved away from the abdomen, and thereby a wide fissure -will be produced, displaying the mechanism beneath. If the whole -operculum be removed, three membranes will be seen, an external, called -the “timbal,” an anterior folded and soft membrane, and a posterior -“mirror.” This last is a very beautiful object, tensely stretched and -pellucid, yet reflecting all the hues of the rainbow. The sound is -primarily produced by the vibrations of the timbal, to which a muscle -is attached; the other membranes are probably also set vibrating, and -the whole skeleton helps to increase or modify the sound, which is -probably also influenced by the position of the operculum, which varies -in different species. A further control of the tension of the air is -exerted by “stigmata” or pores, and certain air-chambers connected -therewith. - -Throughout these pages comment has been made on the apparently -“fortuitous” character of complex patterns and structures. The -“musical-box” of the Cicada affords yet another instance. Nevertheless -there is an impressive harmony between the several parts; an -interdependence which is not fortuitous. There is obviously a nexus of -growth-controlling factors preserving harmony between each separate -part which as yet has escaped all endeavour to discover. - -While it is difficult to picture the initial stages of growth of so -complex an organ as that of the Cicada, the beginnings of simpler -structures such as the stridulating organs of Beetles and Grasshoppers -are more easily discernible. “It is probable,” remarks Darwin, “that -the two sexes of many kinds of Beetles were at first enabled to find -each other by the slight shuffling noise produced by the rubbing -together of the adjoining hard parts of their bodies; and that as -those males or females which made the greatest noise succeeded best -in finding partners, rugosities on various parts of their bodies were -gradually developed by means of sexual selection into true stridulating -organs.” - -Structures to which we can ascribe a use are commonly supposed to have -been evolved for the purpose which we assign to them. The “horns” -of Beetles afford a case in point; but there are many other equally -remarkable and extravagant developments among the insects which seem to -defy explanation. And they will continue to do so until it is realized -that they are but exaggerations of the normal processes of growth, -which is not limited to definite areas but may produce extensions -and excrescences of an almost infinitely varied character. The only -controlling factor is that imposed by Natural Selection when these -growth-changes tend to impair the well-being of the organism as a -whole. Often such changes confer benefits, giving rise to new organs, -and in this case Natural Selection encourages the new departure. -Nothing, indeed, “succeeds like success.” New departures in one -direction may be promptly suppressed, in another they spell fortune: -there is no “Socialism” in Nature. Often these “new departures” neither -help nor hinder, and instances of this kind are commonly afforded -by “ornaments.” One of the most singular illustrations of this kind -is furnished by that extraordinary Long-horned Grasshopper of India -(_Schizodactylus monstrosus_), wherein the wings, when at rest, have -their tips coiled up like a watch-spring, while the appendages to the -legs are scarcely less remarkable. It is a burrower, driving long -tunnels in the banks of rivers. But little is known of its habits, -save that it does not emerge from its burrow till night, when it takes -long flights. This being so, the bizarre character of its wings and -legs is the more difficult to explain on any Sexual Selection theory. -But regarded as spontaneous variations which have not fallen under the -ban of Natural Selection, they are somewhat less puzzling, though, -having regard to the extraordinary transformation which the burrowing -Mole-cricket and the allied Cylindrodes have undergone, in adaptation -to fossorial habits, the legs of this insect are remarkable indeed. - -While there can be no doubt that the musical performances of the -Crickets and Locusts play an important part in courtship, in some of -the Long-horned Locusts, at any rate, the males fight furiously when -mate-hunting, and to this end the head and jaws are greatly enlarged. -During the progress of the duel the wings are extended and held erect, -which is hardly what one would have expected, since in this position -they would seem to be more exposed to danger. - -All the insects so far surveyed have been more or less conspicuous -for their vivid hues, yet in none of these have elaborate “displays” -been recorded. To find demonstrativeness of this kind one must turn -apparently to a group of minute, lowly organized, dull-coloured, -wingless insects with ugly misshapen heads and bodies. The sexes do -not differ in appearance, but they are interesting on account of the -sedulous court which the males pay to the females. The late Lord -Avebury, in a Communication to the Linnean Society, remarked of them: -“It is very amusing to see these little creatures (_Smynthurus luteus_) -coquetting together. The male, which is much smaller than the female, -runs round her, and they butt one another, standing face to face -and moving backwards and forwards like two playful lambs. Then the -female pretends to run away and the male runs after her. With a queer -appearance of anger, he gets in front and stands facing her again; then -she turns coyly round, but he, quicker and more active, scuttles round -too, and seems to whip her with his antennæ; then for a bit they stand -face to face, play with their antennas and seem to be all in all to one -another.” - -The Dragon-flies are among the most beautiful of insects; they are -also relatively long-lived, and they are conspicuous. Yet this beauty -must be attributed to some inherent inward grace rather than to the -æsthetic instincts of the female. Moreover, in the matter of size -and beauty there is little to choose between the sexes; where any -difference occurs the males have the advantage. Though the mode of -copulation is well known, nothing has been discovered as to the means -whereby male and female discover one another. It is doubtful whether -this can be done by sight, for with all the beauty of their shimmering -suits of mail and gauzy wings, their vision is limited to a field -of a few inches. Possibly scent is their guide; at any rate, dead -Dragon-flies have a vile odour. - -It is worth noting that there are no wingless Dragon-flies, and that -none have developed unnecessary ornament in the form of spines, horns, -or frills of any kind, such as are so commonly met with among groups of -more sedentary habits like the Phasmidæ and the Beetles, for example. -In other words, there is clearly a direct relation between ornament -and the mode of life. It is also clear that some modes of subsistence -are very inelastic, allowing of no more than very slight structural -variations, for the Dragon-flies are an extremely ancient group. Fossil -species of large size are known from the Lower Lias, and the remains -of a giant measuring two feet in expanse of wings has been found in -the Carboniferous. This species, however, seems to have stood near the -parting of the ways between the May-flies and the Dragon-flies. But -be this as it may, undoubtedly Dragon-flies hovered over the backs of -sleeping Ichthyosaurs and furnished food for Pterodactyles millions of -years ago, as they now hover over lazy kine for the sake of the flies -forgathered there, or dodge to avoid the stoop of the Hobby, and in all -this vast space of time they have not appreciably changed. - -And what is true of the Dragon-fly is true also of the May-fly, for it -is clear that they are of the same stock. It is true at any rate in so -far as the conformation of the body is concerned. The possibility that -it may be equally true in regard to the details of their life-history -almost staggers one, because these are, in many respects, of a quite -remarkable character. As with the Dragon-fly, there is a prolonged -period of larval life, lasting from one to two years, which time is -passed in streams and pools where a luxuriant vegetation ensures a -plentiful supply of food. Some are carnivorous, but in the majority of -species minute plants only are eaten. More than forty species are to -be reckoned as natives of the British Islands, the commonest being the -“Green Drake” and “Grey Drake,” beloved by the fisherman. These names -are applied, it may be mentioned, to the phase known as the sub-imago -which precedes the fully-adult stage, of _Ephemera vulgata_ and _E. -danica_. - -Save that it is curious that while the larvæ of some species are -carnivorous those of others are vegetarians, there is nothing very -remarkable about what may be called the infantile period. But when -this is ended the span of life remaining to them as adults is brief -indeed. Instinctively realizing that the time of transfiguration is at -hand, the erstwhile crawling grub rises to the surface of the stream, -and almost in the twinkling of an eye it mounts into the air on gauzy -wings, there for a brief space to execute an aerial dance which in its -every phase is amazing. Some species never see the sun. They emerge as -the sky begins to redden, and as its glory fades they, too, expire. -This brief space is all that Nature has allowed them in which to fulfil -her behest to all living things—to increase and multiply. And myriads -die without even a chance to effect this consummation of existence. - -The dance is a Dance of Death, and it is performed by a host so vast -as to surpass the bounds of belief save to those who have had the good -fortune to witness a scene so amazing. - -D’Albertis tells of a gathering which he witnessed on the Fly River, -New Guinea—for these insects have a world-wide distribution—wherein -countless myriads were assembled. “For miles the surface of the river, -from side to side, was white with them as they hung over it on gauzy -wings; at certain moments, obeying some mysterious signal, they would -rise in the air and then sink down anew like a fall of snow.” And in -this assemblage he estimated that there was but one female to every -five or six thousand males. It is during this flight that the act of -mating is performed. The fortunate male from the host of rivals, in -this mid-air embrace is aided by the foremost pair of legs, which are -especially curved to effect this purpose. The embrace is momentary. -Thereafter he dies; to the female a somewhat longer span of life -remains, for she has yet to deposit her eggs, and this being done _en -masse_, she, too, expires. - -It is curious that these creatures, which in their winged state have -never seen the sun, should be attracted by light. But such is the -case. I well remember witnessing an instance of this years ago, while -staying, one August, at Bingen on the Rhine. Dinner was served in the -open air, and just as the soup was served May-flies in myriads swarmed -round the lamps and fell on the tables as thick as snow-flakes. Some -of these were in copula, and I succeeded in bottling a few specimens -for the British Museum, where they still remain to remind me of this -amazing scene. - -About three hundred species of May-flies are known, and some enjoy a -somewhat longer span of life than others. In no case, however, do they -emerge till just before sunset; but in some species it is believed life -may be prolonged for as much as three or four days, or even longer, if -the weather be cold and wet, so as to keep them in a state of enforced -rest, which amounts to a state of coma. - -That their hold on life during this final stage of existence is -brief there can be no gainsaying, for it is passed fasting. Jaws are -wanting, and the whole alimentary canal has been transformed into -one long air-chamber. Its walls are now of extreme tenuity, and by -changes in the interior of the tube, valves are formed which convert -the stomach into a capacious air-sac.” When movements,” remarks Dr. -David Sharp, “tend to increase the capacity of the body cavity then air -enters into the stomachic sac by the mouth orifice, but when muscular -contractions result in pressure on the sac they close the orifices of -its extremities by the valve-like structures just referred to; the -result is, that as the complex movements of the body are made the -stomach becomes more and more distended by air.” It was known even to -the old naturalists that the dancing May-fly is a sort of balloon, -but they were not acquainted with the exact mode of inflation. Palmen -says that in addition to the valve-like arrangements we have described, -the entry into the canal is controlled by a circular muscle with which -are connected radiating muscles attached to the walls of the head. The -canal thus strangely transformed performs the functions of a balloon, -and at the same time aids the functions of the reproductive organs. - -Where vast numbers of individuals set out simultaneously to achieve -their nuptials there would seem to be no need for special devices -on the part of either sex to call attention to their whereabouts. -Nevertheless, it is highly probable that the female exhales some -distinctive odour; otherwise, having regard to the fact that she -is overwhelmingly outnumbered by suitors, her discovery in such a -crowd would be impossible, and it is of vital importance that no -time should be lost in effecting conjugation, for the time for its -accomplishment is perilously short. But there is another possible means -of discrimination—the males may distinguish the females by the very -different appearance of the head in the latter. At any rate, this may -be true of some species wherein the males have no less than seven eyes, -and these of three different kinds! The compound eyes, characteristic -of insects, are, in these, divided, one half being set upon the summit -of a pillar raised high above the level of the head, the other part -remaining in its normal place at the side of the head; and in front of -these, on what may be called the forehead, are three separate simple -eyes, or “ocelli.” A reference to Plate 32, Fig. 3, will make this -clear. - -That the history of the later life of the May-fly is remarkable -no one will deny: in many respects it is unique. Yet for all its -strangeness it enables us to set our compasses, so to speak, in -regard to the phenomena of sex in other groups. The extraordinary -disparity in the proportions between male and female, for example, is -full of significance, for it shows, as has been suggested more than -once in these pages, that, in the case of polygamous species, we are -probably in error in supposing that the excess of females is due to -the reduction in the number of males by reason of the elimination -of males by fighting. The excess of males, or females, as the case -may be, is due to an inherent quality in the germ-plasm. The May-fly -might be regarded as an excessively polyandrous species if the number -of males in relation to females alone be regarded: but actually it -is monogamous. After a prosaic infancy they are suddenly transformed -into gay lovers, dancing a marriage-dance. But for them is no marriage -feast, nor any later sequence of domesticity. One in ten thousand -may find a mate, and only in this is he more fortunate than his -neighbours, for, like them, he too must die before the dawn. Theirs is -not even a sleep and a forgetting, but “one splendid hour of Life, and -then—oblivion.” It may be urged that even these which might seem to -have been fooled, have not really lived in vain, for hosts of animals -feast upon their bodies. Myriads, indeed, are snapped up by fishes even -before they have opened their wings, while birds rudely invade the -swarms as they dance in mid-air, feasting on these fasting ones. But -this is, after all, an inglorious end, and leaves us still asking _Cui -bono_? - -Is this amazing life-history a thing of yesterday, a new phase, or an -order of things as old as the origin of the species, dating back some -millions of years? - -So far as one can profitably speculate on such a theme it would seem -more likely to be a relatively recent innovation. The nearly-related -Alder-flies (_Sialidæ_), so well known to anglers, seem like to meet -with a similar fate, for the female lives but for a few days only -and the male has an even briefer existence as a winged insect. The -family to which the Alder-flies belong contains a few species which -attain gigantic proportions, as, for example, in the case of the North -American members of the genus _Corydalis_, which are giants. The males -thereof are remarkable for the fact that they are armed with enormous -jaws, which may be likened to a pair of callipers whose limbs have been -crossed. These weapons serve as claspers, enabling the males to seize -and hold the females during the act of mating. But even here the same -brief span of life has to suffice them, for death follows swiftly on -the fulfilment of the nuptial rites. - -The Perlidæ, or Stone-flies, which, like the Sialidæ, are aquatic -Neuroptera, the larval stages being passed in streams, present very -puzzling features in regard to the adult males which, so far, have -baffled all attempts at solution; yet they seem to have a very -important bearing on the all-important work of reproduction. They -are among the earliest insects to appear in spring, and possess an -extraordinary power of resisting cold. One species, _Capina vernalis_, -common in the Albany River, in Canada, frequently comes up through -cracks in the ice and casts its skin there! Another, _Nemoura -glacialis_, which appears at about the same time, actually performs -the nuptial rites in crevices in the dissolving ice! Happily reason -is denied them, or they would find life a mockery indeed; for having -attained their final development, when the joyous and exhilarating -exercise of flight should be theirs, they are compelled forthwith -to fulfil their reproductive functions and die—in an ice-chamber! -The males have wings which are rarely or never unfurled; as a rule -they present nothing more than a crumpled mass of gauzy tissue, as if -glued together. Such species as attain to flight are most indifferent -performers, travelling but slowly, with laboured movements and settling -after a few yards have been traversed. - -As a rule, among insects, where there is a difference in the power of -flight, it is the male which is superior. The case of _Nemoura_, just -referred to, affords an instance where the contrary is the case, and -Mr. J. J. Lister records the case of one of these flies—_Isogenus -nubecula_—taken at Loch Tanna in Arran, wherein the wings of the female -were greatly reduced, while those of the male were so much so as to be -mere useless vestiges. Similar facts have been recorded of more than -one species in Scotland, but in all such cases the phenomenon seems to -be associated with the appearance of the insect in very early spring. -In another species—_Nemoura trifasciata_—only the front wings are -reduced, the hind pair being large enough to cover the body. In male -specimens of _Perla maxima_ taken in Scotland, the wings are so short -as to be useless for the purposes of flight, yet, in the same species -taken in Central Europe, they are of ample proportions. - -These facts are puzzling indeed, but they seem to show that flight is -not essential to attain the ends of reproduction. As to whether these -flies secure their mates by any kind of “courtship,” or how they find -one another, seems not to be known. But the female is remarkable for -the fact that she carries her eggs about with her, to the number of -five or six thousand, attached to the end of the abdomen. - -Having regard to the fact that three thousand species of Perlidæ are -known, and that they have a wide distribution over the earth’s surface, -one might have expected that more would be known of their singular -life-history. They are, however, flies of very unattractive appearance -and great frailty, hence, save to anglers, by whom they are esteemed as -bait for trout, they attract but little attention. - - - - -CHAPTER XII - -SCORPIONS, SPIDERS AND CRABS - - -Musical Lovers among Spiders and Scorpions—Colour among Spiders, and -its uses—The Spiders’ Dance of Death—Spiders and Conjugal Bliss—How -Pairing is accomplished—Scorpions in Love—Musical Crabs—Quarrelsome -Fiddler-crabs—Crabs and Courtship in the Deep Sea—Amazons among -Prawns—Brine-shrimps and Water-fleas—“Natural” _v._ “Sexual” Selection. - -It is a curious and significant fact that in the most brilliantly -coloured of the Invertebrates—the Butterflies and Moths—” courtship” -in the sense of “wooing” is extremely rarely met with; and this is -quite contrary to what the Sexual Selection theory of Darwin demands, -for, according to this, the colours are the result of that selection. -On the other hand, Spiders, which are for the most part dull-coloured -creatures, and the Scorpions, which are also dull-coloured, are -commonly extraordinarily demonstrative during the early stages of -“mate-hunger.” Some practise the form of instrumental music known as -“stridulation,” others dance and indulge in other forms of posturing. - -In the Spiders the stridulating apparatus is formed either between -the limb-bearing portions of the body, or “cephalothorax,” and the -abdomen; between the palps or leg-like feelers, and the jaws; or -between these feelers and the front legs. But the construction is -similar in all. In some Spiders the abdomen bears a horny collar, which -is toothed, and these teeth, as the abdomen is raised and depressed, -scrape against a number of delicate ridges on the thorax, or “chest,” -which form a surface recalling that of a file. The grating of these -opposing surfaces against one another produces shrill rasping or -chirping sounds, which, in some cases at any rate, seem to be designed -to inform the female of the presence of a suitor. Those who will, -may examine this strange instrument for themselves if they will take -the trouble to seek for it in one of our commonest English Spiders -(_Steatoda bipunctata_). That it serves as a sexual excitant, or as -an aid to mate-hunting, is indicated by the fact that it is found in -males only, or in a very rudimentary condition in the female. There is -a large Spider in Assam (_Chilobrachys stridulatus_) which produces -a sound like the drawing of the back of a knife along the edge of a -strong comb; and there are others which, by the friction of the feelers -against the jaws, produce sounds like the buzzing of bees. One of the -Wolf-spiders (_Lycosa kochy_) is known as the “purring” or “drumming” -Spider from its custom, at mating-time, of rapidly drumming on dead -leaves with its feelers. It is a wood-haunting species, and runs hither -and thither over the ground as if searching for something, and pausing -frequently to “purr.” This singular method of producing sound recalls -that of the drumming of Woodpeckers on the hollow branches of trees, -and similarly is produced without any special mechanism. - -That the Scorpions should possess similar stridulating organs is only -what we should expect, having regard to their kinship with the Spiders. -In the great Rock scorpions of India and Africa the stridulating -apparatus lies between the basal segment of the pincers and that of -the first pair of legs, and consists of a set of tubercles and a -cluster of curved, hair-tipped spines. During moments of excitement the -pincers are waved up and down so that the spinules scrape against the -tubercles, emitting a rustling sound, which has been compared to that -produced by rubbing a stiff tooth-brush with one’s finger-nails. In -the South African _Opisthophthalmus_ the mechanism differs, consisting -of leaf-like hairs placed on the inner surface of the jaws. But since -both sexes possess these strange sound-producing mechanisms it has -been suggested that their main, if not their only purpose, is to serve -as a warning to enemies to keep their distance. Some of the great -bird-eating Spiders (_Aviculariidæ_) produce a kind of whistle; others, -sounds like the dropping of shot upon a plate. - -These stridulating contrivances present some curious and puzzling -features. In the first place the sounds they produce are never loud -to human ears; therein they differ from the shrill piercing sounds -produced by like mechanism by the Crickets and Grasshoppers, though -even with some of these the notes are, to us, inaudible. In the second, -it has been suggested that where both sexes possess a stridulating -apparatus its purpose is solely to warn off enemies, and this because -the performers have no sense of hearing, and are thus, we presume, -unaware of the sounds they produce. There is something unsatisfactory -about this line of argument. There seems to be no evidence either that -the sounds produced are loud enough to terrify an enemy, or that the -performers are really deaf. - -In cases where the males alone stridulate it is always supposed that -this “music” serves the purpose of a lure, or acts as an excitant, to -the female, even though inaudible to human ears. But there are many -people who are unable to hear the shrill squeal of our native bats. Yet -no one doubts but that all bats hear it. The argument as to the absence -of any sense of hearing in certain Spiders is based on their failure -to respond to the vibrations of a tuning-fork, but this evidence is -not conclusive. Neither is it safe to infer that the presence of -stridulating organs in the adult and immature stages of both sexes, -in some species, precludes their recognition as secondary sexual -characters. They may serve the double purpose of sexual excitants and -terrifying enemies, their motive being expressed by the quality of the -sound as certainly as the timbre of the human voice may express rage or -pleasure. - -Neither Spiders nor Scorpions exhibit any very striking secondary -sexual characters. As a rule the female is the larger, often -strikingly so. Bright colours are rare, and are met with only among -the Spiders, wherein sometimes the male, sometimes the female, is the -more resplendent; where bright colours—apple-green, red and yellow—do -occur, they seem rather to be of the type known as Anti-cryptic, -or aggressive resemblance colours. That is to say, they are hues -developed to deceive prey by reason of the likeness they afford the -wearer to its surroundings. Thus, for example, one of our native -Spiders (_Tibellus oblongus_) is straw-coloured, and has an elongated -body, which is therefore seen with difficulty amid dry grass and -rushes which are the haunts of the species. _Misumena vatia_, one of -the Crab-spiders, resembles the flowers on which it is accustomed -to lurk for its prey. It is of a variable hue, commonly yellow or -pink, and a favoured lurking-place is near the blooms of the great -mullein (_Verbascum thapsus_), where it seizes upon bees coming for -honey. Exotic relatives of this species afford far more striking -illustrations of this kind. One has a pink, three-lobed body which -bears a striking likeness to a withered flower, and it exhales a sweet -odour of jasmine. Insects attracted by the smell are thus readily -pounced upon. Dr. Trimen, of Cape Town, describes a rose-red species -which exactly matches an oleander flower, and to complete the deception -the abdomen is marked with white. The same observer, approaching a -bush of the yellow-flowered _Senecio pubigera_, noticed that two of -the numerous butterflies settled upon it did not fly away with their -companions. Each of these he found to be in the clutches of a spider -whose remarkable resemblance to the flower lay not only in its colour, -but in the attitude it assumed. “Holding on to the flower-stalk by -the two hinder pairs of legs, it extended the two long front pairs -upwards and laterally. In this position it was scarcely possible to -believe that it was not a flower seen in profile, the rounded abdomen -representing the central mass of florets, and the extended legs the ray -florets; while to complete the illusion the femora of the front pair -of legs, addressed to the thorax, have each a longitudinal red stripe -which represents the ferruginous stripe on the sepals of the flower.” -But more remarkable still is the case cited by my friend Dr. H. O. -Forbes. This came under his notice while butterfly hunting in Java. The -butterflies of the family _Hesperidæ_ have a habit of settling on the -excreta of birds. Forbes noticed one on a leaf apparently enjoying a -feast. Creeping up, he seized hold of this victim of a depraved taste -and found it mysteriously held down. On further examination of this -“excreta” he found that it was really a spider! Later, when in Sumatra, -the same species once more in like manner deceived him. The deception -is more than usually remarkable, for it is not due to the coloration -of the body, but to what may almost be described as a diabolically -ingenious display of intelligence. For the creature weaves upon a leaf -a small white patch of web exactly resembling the fluid excrement of -a bird sliding down the smooth surface of the leaf. Having completed -this, the weaver lies on its back in the middle of the web holding on -by the spines with which the legs are furnished. It then awaits its -victim with the disengaged portions of the legs ready to close in a -deadly embrace the moment the lure has done its work. Though somewhat -in the nature of a digression, these facts show that colour often -plays a vital part in well-being; though in the matter of courtship -its rôle has probably been overestimated. Colour as an aid to “mate -hunting “probably nowhere plays so important a part as was at one time -believed. The Warblers among the birds, and the Spiders among more -lowly animals, seem to demonstrate this fact. - -The actual mating of Spiders, the act of coition, is peculiar, and -demands notice, for the orgasm is not accomplished at the moment of the -ejection of the sexual products. The male discharges the seminal fluid -upon a small web woven for the purpose, and the liberated spermatozoa -are then sucked up into a tube—the _receptaculum seminis_—which lies -coiled up within a hollow bulb attached to the base of the last joint -of the leg-like feeler, or “pedipalp” at the base of the head. The -precious fluid is there stored and retained until the moment arrives -when these palps can be thrust into die genital aperture of the -female, and their contents discharged for the second and last time. -This is the critical moment of the Spider’s life, and it is noteworthy -that it should occur now, instead of at the moment of the discharge -from the body. The ejection from the palpal organ is effected by means -of a fibro-elastic bag, in its normal, collapsed, state spirally -disposed round the base of the bulb which contains the sperm tube. -Immediately preceding copulation this elastic bag or “hæmatodocha” -becomes turgid with blood, and it is probably the pressure thus exerted -on its base which affords the final fury of desire without which, -indeed, one might well imagine the necessary courage for copulation -would never be raised, at any rate, in the case of some species. - -Strange as these facts are, the nice adjustment of the instincts for -their effectual performance is, by comparison, stranger still. By what -subtle sense is the male Spider informed of the importance of the -fertilizing fluid which escapes his body? What prompts him before its -escape to prepare a web for its reception? What prompts him after its -deposition to collect it within the palp till it shall be needed? The -least defect in the instincts appertaining to these vitally important -acts would mean the extinction of the race. We cannot suppose that the -nature of their performance is in any way realized by the performer, -and this makes their orderly execution the more wonderful. - -[Illustration: Plate 34. - -MALE ASTIA DISPLAYING BEFORE THE LESS BRILLIANT FEMALE.] - -[Illustration: - -From drawings, T. Carreras, in “Marvels of the Universe.” - -MALE ICIUS DISPLAYING. - -The “courtship” of the male spider takes the form of a “display” -recalling that of birds. He commonly ends in being eaten by his mate. - -Face page 242.] - -Our knowledge of Spiders under the afflatus of sexual desire has been -immensely increased by the long and patient observations of Mr. and -Mrs. Peckham. The fact that their investigations were carried on with -captive specimens, and therefore under artificial conditions both as -to environment and the number of individuals placed together at one -time, must not be lost sight of; nor must we forget that they worked -under the firm conviction that the Sexual Selection theory of choice -by the females was an indisputable fact. Wherever colour was present -they looked for, and saw, evidence that the female appreciated such -hues, though from their observations it would seem that dull-coloured -species behaved as though they were suffused with resplendent hues. -In the course of their studies the courtship of several species was -investigated, but a summary of their results is all that can be given -here. _Saitis pulex_ formed the subject of one of their experiments. -A male was placed in a box containing a mature female. “He saw her as -she stood perfectly still, twelve inches away; the glance seemed to -excite him and he moved towards her; when some four inches from her he -stood still, and then began the most remarkable performance that an -amorous male could offer to an admiring female. She eyed him eagerly, -changing her position from time to time so that he might be always in -view. He, raising his whole body on one side by straightening out the -legs, and lowering it on the other by folding the first two pairs of -legs up and under, leans so far over as to be in danger of losing his -balance, which he only maintains by sidling rapidly towards the lowered -side. The palpus, too, on this side was turned back to correspond to -the direction of the legs nearest to it. He moved in a semicircle for -about two inches, and then instantly reversed the position of the legs -and circled in the opposite direction, gradually approaching nearer and -nearer to the female. Now she dashes towards him, while he, raising his -first pair of legs, extends them upwards and forwards as if to hold her -off, but withal slowly retreats. Again and again he circles from side -to side, she gazing towards him in a softer mood, evidently admiring -the grace of his antics. This is repeated until we have counted one -hundred and eleven circles made by the ardent little male. Now he -approaches nearer and nearer, and when almost within reach whirls madly -around and around her, she joining and whirling with him in a giddy -maze. Again he falls back and resumes his semicircular motions with his -body tilted over; she, all excitement, lowers her head and raises her -body so that it is almost vertical; both draw nearer; she moves slowly -under him, he crawling over her head, and the mating is accomplished. -After they have paired once the preliminary courtship is not so long. -On one occasion a female was the more eager of the two, but this is -evidently very exceptional. The female always watches the antics of the -male intently, but often refuses him in the end, even after dancing -before her for a long time.” - -Of another species—_Epiblemum scenicum_—they write: “The females seemed -to have some difficulty in choosing from among the males, but after a -decision has been reached and a male accepted, there appeared to be -complete agreement.” A species of the genus Iritis, which seems to have -baffled identification, was watched for hours under natural conditions -as well as in confinement.” A dozen or more males, and about half as -many females, were assembled together within the length of one of the -rails. The males were rushing hither and thither, dancing opposite now -one female, now another; often two males met each other, when a short -passage of arms followed. They waved their first legs, sidled back and -forth, and then rushed together and clinched, but quickly separated, -neither being hurt, only to run off in search of fairer foes.” - -These most patient observers seem to have been convinced that whenever -Spiders possess vividly coloured areas on their bodies they are not -only conscious of this fact, but desire to make the most of such -splendour during the period of love-making. Thus they interpret the -behaviour of a curiously ant-like Spider—_Synageles picata_—which -has the first pair of legs especially thickened, flattened on the -anterior surface, and of a highly iridescent steel-blue colour. As -he approaches the female he pauses “every few moments to rock from -side to side, and to bend his brilliant legs so that she may look -full at them; ... he could not have chosen a better position than -the one he took to make a display.” And similarly they interpret the -movements of another species—_Dendryphantes capitatus_—which has a -bronze-brown face, rendered conspicuous by snow-white bands. The -attitude he assumes when sexually excited is one which seems, to them -at any rate, to serve admirably to expose this feature to the watchful -female. But he has other charms, and his “antics are repeated for a -very long time, often for hours; when at last, the female, either won -by his beauty or worn out by his persistence, accepts his addresses.” -_Habrocestum splendens_—unhappily these creatures have no names in -common speech—possesses an abdomen of a magnificent purplish red, and -the attitude which he assumes at courtship they regard as one designed -to display this to the full. Another case of quite remarkable interest -is that of _Astia vittata_, because the males appear to be dimorphic. -That is to say, they appear under two quite distinct forms, the one -red, like the female; and the other black, with three tufts of hair -just behind the head. The attitudes and the movements of courtship, it -is significant to remark, are entirely different in the two varieties: -the black form, assumed to be the later development, “is much the more -lively of the two, and whenever the varieties were seen to compete for -a female, the black one was successful.” - -Professor Poulton, commenting on this particular case, contends that -“it must be admitted that these facts afford the _strongest support_ to -the theory of Sexual Selection.” But do they? A further examination of -the facts will probably show that the red “form” is but an immature -example, and this being so, the difference in performance and the -invariable success of Othello is at once accounted for. The fact that -the “two forms pass into each other” and that the “tufts only occur in -the fully developed _niger_ form” is an additional reason for regarding -the red form as immature. - -Professor Poulton remarks: “When the males possess any special -adornment they make a point of displaying them as fully as possible.” -If this be so it seems to be a very foolhardy proceeding, akin to -holding the proverbial “red rag” to a bull: for it is well known that -the male Spider seeking a mate carries his life in his hands, at any -rate in the case of many species. Mr. and Mrs. Peckham observed several -instances of this remarkable sequel to Love’s embraces. In describing -the female of _Phidippus morsitans_ they remark that she was “a savage -monster. The two males we provided for her had offered her only the -merest civilities, when she leaped upon them and killed them.” The -first pair of legs in the males of this species possess “special -adornments” in the shape of long white hairs, and it was “while one -of the males was waving these handsome legs over his head that he was -seized by his mate and devoured.” Again, in the case of a male of -_Phidippus rufus_, the display of his “ornaments” was his undoing, -for he was “caught and eaten when he insisted upon showing off his -fine points too persistently.” Thus the females seem to “select” the -more resplendent males as much for eating as for mating! The ogre-like -habits of the females in this regard, indeed, are almost without -parallel in the animal kingdom. - -Anyone who cares to take the trouble to watch the web of the large -Garden Spider (_Epeira diademata_) may witness one of these connubial -tragedies. In this species, the males are conspicuously smaller than -the females, and it is possible that this disparity has been brought -about by Sexual Selection, the largest and least active males having -been exterminated. In some species the discrepancy in size is most -striking, as for example in _Nephila chrysogaster_, the female of which -measures two inches in length, the male not more than one-tenth of an -inch, and less than one one-thousand-three-hundredth part of her weight. - -The males, apparently, fully realize the perils which their amours may -lead them into. They haunt the borders of the webs of unmated females, -but exhibit a hesitating, irresolute manner. For hours they will linger -near her, feeling the silken carpet cautiously with their legs, and -apparently trying to ascertain the nature of the welcome likely to be -extended to them. The odds are against them: for even if allowed to -mate, unless they are extraordinarily agile in slipping away the moment -they have attained their object, the chances are they will be slain and -eaten! - -Among some species, however, matters are otherwise: for the males of -the genus _Linyphia_, for example, are generally to be found living -peacefully with their consorts. - -More rarely the male weaves a small nuptial tent, into which he -partly leads and partly drives the female: though the “driving” would -appear to be merely for form’s sake! The habits of the Cellar Spider -(_Tegenaria parietina_), a long-legged species fairly common in the -South of England, affords a yet further interesting and instructive -contrast with the foregoing accounts. The pairing habits of this -species have been studied by many observers, but perhaps the best -account is that of Mr. F. M. Campbell. He found, to begin with, that in -this species the tender ties of mating are at any rate rarely violated -by the horrible aftermath of cannibalism so common a feature with so -many other Spiders. - -One or two illustrations from Mr. Campbell’s work must suffice. On -one occasion he placed together a male and a female. For four days -they took no notice of one another; then the female cast what proved -to be her last skin, and within three hours after, the male began -to show signs of interest in her presence—which is a fact of some -significance, for not till then had she attained maturity. “After a -few convulsive twitchings of the legs, the male pressed forwards, -moving his palpi”—the leg like “feelers” on each side of the head -which form the genital organs (page 241)—“up and down, when, as they -touched the palpi of the female, the pair played with these organs -like two friendly bees with their antennæ. After a few minutes the -female raised herself, leaning a little on her left side, and the male -crept forward until his head was under the breast of his mate, while -his first pair of legs were resting upon hers. He now advanced his -right palpus, leaning a little to the left and using the left palpus -as part of his support. The right palpus was slightly twisted so as -to bring the surface (containing the fertilizing germs) opposite the -sexual organs of the female.... He now rapidly raised his palpus up and -down for four or more seconds, and with such energy as to compel her -to assume a vertical position. He then retired and again approached -her, repeating the movements ... occasionally pausing before he -withdrew his palpus.... At times he would leave the female for five -minutes, and strut with straightened legs round the vase, wagging his -abdomen. Now and then he would remain perfectly still with the palpus -withdrawn, or play with the palpi of the female, while she seemed in -a comatose state. He would then renew the union with undiminished -vigour, appearing on each occasion less desirous of changing his -position. I left them at 12-30 a.m. and returned at 7 a.m. The male -was still using his right palpus. I saw no application of the left -palpus, but have no doubt that it was employed during the night, as -in other cases. I have not observed the pairing ever interrupted by a -fresh collection of semen, although there is no reason to think this -may not occur. The duration of pairing is long; but I am inclined to -think it is more dependent on the difficulty in inserting the palpus -than on sexual endurance. The impregnation appeared to take place -when the male retained his palpi in front of the bursa copulatrices -for about thirty seconds, which was frequently the case.” There -are occasions, however, when a very different sequel attends this -dalliance. In one instance, for example, Mr. Campbell placed a pair -together, and at once the male began to pay his addresses. “Shortly -afterwards he rapidly applied one of his palpi to the female ... -apparently with her consent.” Five hours later “he charged her, tore -away two legs ... and began to suck one, using the mandibles to hold -the limb as a human being would a stick of asparagus.” It is not -surprising to find she died an hour afterwards. An examination of her -remains brought to light the fact that she was not mature. But this -does not apparently explain the ferocity of her partner, for this -investigator on two other occasions saw males similarly dismember -their spouses an hour or so after impregnation. This horrid feast -cannot have been prompted by hunger, for one of these males had, but -a few hours previously, eaten a daddy-long-legs and two blow-flies. -Only twice did this investigator see a female of this species drive -away a male, and in each case immediately after union. “On the other -hand,” he says, “I have kept an adult pair together from the 22nd of -August to the 28th of October, and they lived in perfect amity. The -male never ceased paying unrequited attentions except to feed.” It will -have been remarked that the behaviour of this species in regard to -mating differs conspicuously from the accounts of observations on other -species, wherein the aggressive instincts are displayed by the female. -Mr. Campbell, commenting on these facts, remarks that such conduct is -just what one would expect from creatures which lead solitary lives, -and must have “come to regard weaker forms of animal life as food, or -as an inconvenience, if we except its young or its mate when in the -act of pairing.” Instincts which are habitually practised throughout -the greater portion of the life of the species, and on which existence -depends, would scarcely be suspended for a longer period than necessary -for sexual union. Spiders frequently eat one another, and such an -occurrence after pairing is only curious if considered apart from their -habits. When the sexual desire is satisfied, their actions would again -be directed by the dominant instinct of destruction. - -It is to be noticed that the attack, when made by a female, often -immediately follows the sexual union, while in the case where males -assume the aggressive it takes place some time afterwards. Mr. Campbell -explains this by the supposition that the action of the female, when -satiated, would be precipitated by the threatened and unacceptable -continued application of the hard, spiny palpus, while the more lasting -desire of the male would have to subside before he became directed by -another instinct. By that time, other attractions, if not his wandering -disposition, would take him away from the web. - -The fact that male Spiders are comparatively rare is perhaps explained -by the fact that they are very short-lived; they probably die soon -after pairing—even if they are not eaten! The snares they spin, it is -to be noted, are very imperfect, though curiously enough, when young -they make perfect snares on a small scale. - -It will have been noticed, in the course of the foregoing descriptions, -that Spiders display a more or less conspicuous wariness, a cool, -deliberate “counting on the cost” in their matrimonial ventures that is -often wanting in such matters in the human race. But, then, the risks -involved are more patent, more imminent. Mr. Campbell comments on this -intelligent behaviour in the case of the Cellar Spider, remarking that -they measure “each other’s strength when on the same web by the tension -and motion of the threads.” - -A word as to the Scorpions. These creatures are near relations of the -Spiders, and in many things resemble them, notably in regard to their -ferocity. One does not meet here, however, with the same disparity in -size between the sexes, nor are vivid colours ever developed. This, -according to some, would be accounted for by the fact that though these -creatures possess numerous eyes they are practically blind, and depend -for their information as to what is going on around them by their sense -of touch, which is excessively delicate. They are morose in disposition -and always solitary. It has been said that if two are found under the -same stone—a favourite lurking-place—one is engaged in eating the -other! Nevertheless, they are of abstemious habits, for the naturalist -Fabre found that from October to March they last, though throughout -this time they remain alert, and always ready to resent disturbance. In -April they exhibit more activity, though even then they eat but little. -But now they begin to wander in search of mates. - -Fabre’s observations on their mating habits are exceedingly -interesting, and they have brought to light some very extraordinary -phenomena. His notes were made on the species common in the South -of France—_Buthus occitans_. Mr. Cecil Warburton, referring to the -distinguished Frenchman’s work, quotes the following noteworthy passage -in the Cambridge Natural History: “After some very curious antics, in -which the animals stood face to face with raised tails, which they -intertwined ... they always indulged in what Fabre calls a ‘promenade -à deux’ hand in hand, so to speak, the male seizing the chelæ of the -female with his own and walking backwards, while the female followed, -usually without any reluctance. This promenade occupied an hour or -more, during which the animals turned several times. At length, if -in the neighbourhood of a suitable stone, the male would dig a hole, -without for a moment entirely quitting his hold of the female, and -presently both would disappear into the newly-formed retreat.” - -[Illustration: Plate 35. - -Photo by P. H. Fabre. - -SCORPIONS. - -The early stages in the courtship of the scorpions are full of romance. -The two prospective partners for life engaging in a kind of waltz, -holding each other’s “hands.” - -Face page 252.] - -After the mating, as with the Spiders, the male is often devoured by -the female. After any combat with an enemy, such as a _Lycosa_ or a -_Scolopendra_, it seems to be _de rigueur_ to eat the vanquished. - -If the mating period in the case of the higher animals rouses the -males to the pitch of frenzy, that frenzy is dangerous only to -possible rivals. With the more lowly Spiders and Scorpions ferocity of -disposition is a normal feature, and one which can with difficulty be -held in check long enough to permit the all-important act of mating -to take place. In how far this is accounted for by the extremely -deficient senses of sight and hearing, which are such marked features -in these animals, it would be difficult to estimate. But that the -manner of their display is governed by these deficiencies there can be -no doubt. The Spider, having a more or less efficient vision at short -range, executes more or less elaborate antics in front of the female, -designed, as in the case of the birds, to serve as excitants to fan -sexual desire, already smouldering, to a flame. With the purblind -Scorpion the Spider-antics are useless; he must proclaim his desire -by a pressure of the hand, and by intertwining his tail with that -of his prospective mate as they “walk out” together. But Scorpions -at one time were credited with a very acute sense of hearing; later -investigations, however, fail to yield any evidence whatever that they -possess this sense, though experiment has proved that their sense of -touch is excessively delicate and seems to reside in the hairs which -are thickly distributed over the legs and body. Now, hearing and touch -are senses near akin, and the vibrations produced by stridulation may -be, and probably are, received by, and interpreted through, the medium -of these hairs. For though the Scorpion may not respond to sounds made -by curious investigators, it may be that they can perceive notes of a -low pitch imperceptible to our ears, such as are made by stridulating -organs, as in the case of the Spiders. - -Perchance certain comb-like structures known as the “pectines” may -play a part in mate-hunting. These are placed on either side of the -under-surface of the body between the last two pairs of legs. The fact -that they are larger in the male, and sometimes strangely modified in -the female, seems to show that they have some function in relation to -sex. They also appear to serve as sources of information as to the -nature of the ground traversed by the animal, since they are long in -species which walk with the body raised high off the ground and short -in such as adopt a more grovelling posture. That the Scorpions possess -but a very limited means of gleaning information of the outer world -there can be no doubt. How, then, do they find one another when that -insistent desire to mate begins to make itself felt? Are the “pectines” -their informants through the sense of smell? Do the hairs scattered -over the body act as sound-collectors responding to the notes emitted -by the stridulating organs? These are points on which information is -much to be desired. - -[Illustration: Plate 36. - -Photo by P. H. Fabre. - -DEATH OF THE MALE SCORPION. - -But by the time the nuptial rites have been performed the female has -thrown off her “sweetness,” and ends by eating her lover! - -Face page 254.] - -[Illustration: Plate 37. - -Photo by Paul H. Fabre. - -THE FEMALE MANTIS DEVOURING HER MATE. - -With these insects, as with the spiders and scorpions, the male is -often eaten by the female.] - -Our survey of the “Arthropoda,” as those limb-bearing jointed animals -invested in a horny, or, more exactly, a “chitinous” external skeleton -are called has so far been confined to such as, during adult -life, at least, are land-dwellers. But the aquatic types known as the -“Crustacea” furnish some extremely interesting facts in regard to the -problems of sex. In the first place, they too possess a stridulating -apparatus. This is curious, but not surprising, because, although -the skeleton of such creatures is of a harder and almost stone-like -character, the development of roughened surfaces working in opposition -to one another might well have been foretold to occur, at least in -some individuals. Colonel Alcock—a naturalist who has contributed -largely to our knowledge of marine animals by his researches in the -Indian Ocean—in his most delightful book “A Naturalist in Indian Seas,” -describes what he calls a “musical crab.” This is the great-horned -Coromandel Strand Crab (_Ocypoda macrocera_). In both sexes of this -remarkable genus he says, “the nippers, or chelipeds, are singularly -unequal in size, and in all the species but one there is present on -the inner surface of the ‘hand’ of the larger cheliped a transverse -row of five teeth, which, when the cheliped is flexed, can be made to -play against a ridge or another row of teeth on its ‘arm’ ... much as a -man might rub one side of his chest with the palm of the corresponding -hand. The whole mechanism, except that it is on a larger scale and has -a more finished appearance, is very much like that by means of which -crickets and grasshoppers produce their shrill music, and no one has -ever doubted that it is used for the same purpose, though very few -people have actually heard it in action. I myself ... was beginning -to think that the structure must, after all, have some quite other -function, when one morning ... on the sandy wastes of the Godavari -delta, I at last, like Ancient Pistol, heard with ears that which I -had been so long waiting for. That is to say, I heard a noise very -much like that which an angry squirrel makes, and discovered that it -came from a red ocypode crab into whose burrow another individual had -trespassed. - -“In order to understand the matter it should be known that these -crabs ... are gregarious, and that each one has a burrow of its own. -Though they may be seen marching in battalions across the sand, yet as -a rule they stay close to their burrows, methodically searching and -sifting the surrounding sand for any food that may have been thrown -up by the tide, and flying to their burrows with headlong speed when -alarmed. At first sight one does not understand the necessity for so -much wariness, and for such a deep system of entrenchment, for the -creatures seem to hold undisputed possession of the whole shore; but as -a matter of fact they are preyed upon all day long by Brahminy kites, -and when the jackals come out in the evening, by them. Now, although -each crab may on ordinary peaceful occasions know its own home, yet -when a crowd of them are running for their lives they may sometimes, -one would think, act on the devil take the hindmost principle and -try to squeeze into the nearest burrow. But as ancient philosophers -do report, things may be done upon occasion which it is inexpedient -to make a habit of doing, and this seems to be one of those things; -for if many Crabs made a practice of crowding into one small burrow -they would certainly run the risk of being suffocated, if not crushed -to death outright. It seems probable, therefore, that it would be -advantageous to the species as a whole if the rights of property in -burrows were rigidly respected, and if each individual member possessed -some means of giving notice that its burrow was occupied ... and -I think that this consideration gives us a clue to the use of the -stridulating mechanism. At any rate, I was often able, after my first -accidental discovery, to elicit the sound, by catching one of these -crabs and forcing it into a burrow which I knew was already occupied: -the intruder would never go far in, but would crouch just inside the -mouth of the burrow, and if it were made to travel deeper, then the -voice of the rightful owner would be heard in indignant remonstrance -from the depths.” Another species, the Grey Ocypode Crab (_Ocypoda -ceratophthalmus_), possesses a similar instrument, and makes therewith -a loud, croaking noise. But it does not often burrow deeply. Colonel -Alcock therefore suggests that in this case it may be used for scaring -enemies. - -That these curious musical instruments may also be used in mate-hunting -seems highly probable. If the stridulation is produced on one occasion -to announce the fact that callers are not desired, it may on another -signify an equally emphatic invitation to enter, the mood of the -occupant being expressed by the character of the sounds emitted. It is -significant, at any rate, that there are no external sexual differences -in these species; hence the probability that it is by stridulation that -the sexes distinguish one another. - -This view seems to obtain confirmation from the fact that the Crabs of -the genus _Gelasimus_, or “Fiddler-crabs,” which are near relations of -the ocypode Crabs, and, like them, live in burrows in large companies, -and are exposed to the same enemies, which they avoid in the same -way by burrowing, have no stridulating mechanism, but the sexes are -strikingly different. This is especially so in the case of the nippers, -or chelipeds. These, in the female, are slender and much shorter than -the legs, being used mainly for feeding. In the adult male one of these -“hands” is often twice as big as the body itself! “Many uses,” remarks -Colonel Alcock, “have been assigned to this enormous, lop-sided organ: -some say that it is used as a stopper to barricade the mouth of the -burrow, others that it is a sort of cradle or bridal-couch upon which -the female reclines—the male, in this case, literally bestowing his -hand upon the female; but from observations of _Gelasimus annulipes_, the -species which most frequents the Godavari mud-flats, I believe that it -primarily serves as a war-club, for the males indulge in interminable -tournaments for the hand of the female; and secondarily, for it is of a -most beautiful cherry-red colour, as an ornament to attract and delight -the latter capricious sex. - -“Landing one afternoon in March upon a cheerful mud-flat of the -Godavari sea-face, I was bewildered by the sight of a multitude -of small pink objects twinkling in the sun, and always, like -will-o’-the-wisps, disappearing as I came near to them, but flashing -brightly on ahead as far as the eye could reach. It was not until I -stayed perfectly quiet that I discovered that these twinkling gems were -the brandished nippers of a host of the males of _Gelasimus annulipes_. -By long watching, I found out that the little creatures were waving -their nippers with a purpose—the purpose apparently being to attract -the attention of an occasional infrequent female, who, uncertain, -coy, and hard to please, might be seen unconcernedly sifting the sand -at the mouth of her burrow. If this demure little flirt happened -to creep near the burrow of one of the males, then that favoured -individual became frantic with excitement, dancing round his domain -on tip-toe and waving his great cherry hand as if demented. Then, if -another male, burning with jealousy, showed a desire to interfere, the -two puny little suitors would make savage back-handed swipes at one -another, wielding their cumbrous hands as if they had no weight at all. -Unfortunately, though I spent many a precious hour on the watch from -time to time, I could never see that these combats came to anything; -the males seemed always to be in a state of passionate excitement, and -the females to be always indifferent and unconcerned; and though the -dismembered chelipeds of vanquished males could often be seen lying -on the battle-field, I never had the satisfaction of beholding a good -stand-up fight, fought out to the sweet end, or a female rewarding a -successful champion with her heartless person.” - -[Illustration: Plate 38. - -THE “FIDDLER-CRAB” AMONG MANGOE ROOTS. - -This species is remarkable for the enormous size of the right “arm,” -which exceeds that of the body.] - -[Illustration: _Photos by W. Saville-Kent._ - -THE “FIDDLER-CRAB.” - -This “strong right arm” is used in conflicts with rivals for the -possession of the females. - -[Face page 258.] - -The fascinating tale of Colonel Alcock’s observations does not end -here, however, for he has brought to light some extremely interesting -facts in regard to the sexual aspect of Crustacean life in the deep -sea; information gathered during his exploration work on board the -Investigatory much of which was done to enlarge our knowledge of the -abysses of the ocean where the light of day never penetrates. Here, he -remarks, the conditions of life might seem to be reduced to a minimum -of simplicity, yet evidences are not wanting that, among the higher -Crustacea, they are complicated, much as they are everywhere else, by -the play of the sexual instincts. - -In these awful depths, where reigns eternal night, most of the -inhabitants, of whatever kind, from fishes downwards, are blind and -eyeless, or they possess enormous eyes and a purblind vision responding -to the only light these regions display, that of phosphorescence, -which is generated by so large a number of those creatures which are -condemned by Fate to live this sunless life. - -“It is written,” he remarks, “that the male must exert himself to -find a mate, and where sight cannot help him in his search, a kind of -blind-man’s buff is the only alternative. In this serious game many -deep-sea Crustacea, especially those of the Shrimp-tribes, trust to the -sense of smell, as the greatly developed outer, or olfactory, branch of -the first pair of antennæ bears witness. These antennæ, again, seem -to be used by the males of some species for catching their partners, -and in _Parapeneus rectacutus_ ... they are turned into a sort of crook -for this purpose. This has long been thought to be their function in -the Prawns of the oceanic genus _Sergestes_.” In the male of certain -other deep-sea Prawns, the hind pair of foot-jaws are modified in a way -which can only mean that they are used for hooking on to a partner of -the opposite sex. In the deep-sea Hermit-lobsters of the genus _Munida_ -the nippers are greatly enlarged, as in many Shore-crabs, for the -purpose of subjugating rivals and embracing the females; and in all -such cases these are much smaller in the female and immature male. - -Mention of numerous cases has already been made where the female is -larger than the male, and is the more pugnacious, and in such cases -the females are generally more numerous than the males. Some of -the deep-sea Prawns exhibit the same peculiarity. And in these the -sword-like forward prolongation of the head-shield is far larger than -in the male. Now this rostrum is the most formidable weapon which the -Prawn possesses, so that we may, with tolerable certainty, conclude -that the females fight their rivals for the possession of the males, -which are, in these species, far less numerous than the females. - -Among the lower Crustacea, such as the “Fairy-shrimps,” -“Brine-shrimps,” the “Water-fleas,” and the “Copepoda,” which play so -important a part in furnishing food for many of the fishes which in -turn feed us, secondary sexual characters of an extremely interesting -kind are met with. These, however, are never such as appeal to the eye, -for the vision in these creatures is but feebly developed. Scent, as -is usual where sight is defective, plays an important part in enabling -the sexes to discover one another. Selection here secures success only -to such as have the proper odour and the most sensitive organs of -smell. In these creatures, as with the butterflies and moths, the odour -emanating from the female is most powerful, while the sense of smell -is most developed in the male. One of the most striking illustrations -of these facts is furnished by that very beautiful species _Leptodora -hyalina_—a veritable giant among these small Crustacea—wherein the -antennæ of the male are produced into enormously elongated comb-like -structures, the teeth of the comb being formed by delicate olfactory -filaments. In the female these antennæ are extremely short and their -olfactory filaments are limited to a small terminal tuft to the -antennæ, answering to the larger tuft at the base of the comb of the -male. - -To the majority of species, however, delicate odours seem to make -little or no appeal, since excessive development of the olfactory -apparatus, such as is seen in the aberrant Water-flea (_Leptodora_), -is rare. This is perhaps explained by the fact that Leptodora is a -species which does not herd together in vast numbers, hence, probably, -the need of some exceptional means whereby the males may discover -the whereabouts of the females, while in the case of the swarming -hosts formed by Water-fleas and Brine-shrimps, for example, no such -highly specialized aid is necessary. Instead, the males have developed -powerful arms for the capture and retention of the females. In the case -of the Brine-shrimp these arms are of quite formidable proportions. The -males of the Copepoda, remarks Weismann, “possess on their anterior -antennæ an arrangement which enables them to throw a long, whip-like -structure like a lasso round the head of the female as she rapidly -swims away. The antennæ of the male Daphnids, too, are in one genus -(_Moina_) developed into a grasping apparatus; ... the first antennæ -... are not only much longer and stronger than those of the female, -but they are also armed with claws at the end, so that the males can -catch their mates as with a fork, and hold them fast. And even that -was not enough, for, in addition, the males of most Daphnids possess a -sickle-shaped but blunt claw on the first pair of legs, which enables -them to cling to the smooth shell of the female, and to clamber up on -to it to get into the proper position for copulation. - -[Illustration: Plate 40. - -SOME REMARKABLE DEVICES. - -1. A Water-flea (_Moina rectirostris_): male showing the claspers-the -front pair of “legs,” for grasping the female. - -2. The female of the same, in which the “claspers” appear as mere -stumps. - -3. The aberrant Water-flea (_Leptodora kindlii_): the male showing the -long comb-like antenna for the discovery of the female (the left only -is drawn), and the female, just beneath, lacking this olfactory organ. - -4. An extraordinary species of Bug in which the upper surface of the -thorax has been produced backwards to form an overhanging pent-house, -of unknown function, and illustrating the theory of “Hypertely.” - -Face page 262.] - -“If we inquire into the manner of the origin of secondary sexual -characters of this kind, we shall find that both may have been -increased by sexual selection, for a male with a better sickle will -succeed more quickly in getting into the proper position for copulation -than one with a less perfect mechanism. This assumption does not rest -on mere theory, for I was once able ... to observe for a considerable -time, under the microscope, a female to whose shell two males were -clinging, each trying to push the other off. Nevertheless, it seems -to me very questionable whether the origin of this sickle-claw -can be referred to sexual selection, for without this clamping-organ -copulation in most Daphnids would not be possible. It was thus not as -an advantage which one male had over another that the clamping-sickle -evolved, but rather as a necessary acquisition of the whole family, -which must have developed in all the species at the same time as the -other peculiarities, and notably those of the shell. The competition of -the males among themselves is thus in this case simply an expression -of the struggle for existence on the part of the species as such, and -it is not a question merely of a character which makes it easier for -the males to gain possession of the females, but of one which had -necessarily to arise lest the species should become extinct. In other -words, in this case Natural Selection and Sexual Selection coincide. - -“The case of the antennæ of _Moina_, which have been modified into -grasping organs is quite different; these owe their origin, not to -natural selection, but to sexual selection, for antennæ of that kind -are not indispensable to the existence of the species, as we can see -from the closely related genera, Daphnia and Simocephalus, where the -males have quite short, stump-like antennæ, furnished with olfactory -filaments not much more numerous than the females possess. Just as -these supernumerary olfactory filaments were produced by sexual -selection and not by the ordinary natural selection, because those -males with the more acute sense of smell had an advantage over those in -which it was blunted, so the males of the genus Moina which could grasp -most securely had an advantage over those that gripped less firmly, and -thus arose these two different kinds of male characteristics. Neither -of them is of advantage to the species as such, but only to the males -in their competition for the possession of the females.” - -Much uncertainty would seem to exist in regard to two very -extraordinary marine species of Copepoda. In one, _Calocalanus pavo_ -the male possesses enormous antennæ, and a remarkable development of -iridescent feather-like structures at the end of the body, arranged -in a sort of open fan-work; the female has what may be called “normal” -antennæ, and a brush-like tuft at the end of the body. In the other -species—_Calocalanus plumulosus_—of which the female only is known, -there is a similar arrangement of plume-like structures at the end of -the body, but all but one are extremely small; the single plume differs -from the rest in being of enormous length. Commonly these structures -are regarded as mechanisms to reduce the expenditure of energy -necessary to keep at the surface of the water, for these creatures -inhabit the surface-waters of the open ocean. Many larval Crustacea -inhabiting similar areas are in like manner kept afloat, or at any rate -aided in keeping afloat, by the excessive development of spines. But -if this be the purpose of these strange excrescences of _Calocalanus_ -it seems curious that the female of C. pavo should not be similarly -provided. If they are to be regarded as secondary sexual characters it -is curious that the females of _C. pavo_ and _C. plumulosus_ should be -so utterly dissimilar. The male of _C. plumulosus_ is unknown. On the -whole, it seems more reasonable to regard these strange structures as -mechanical aids to swimming rather than as secondary sexual characters. - - - - -CHAPTER XIII - -SOME STRANGE MARRIAGE-CUSTOMS: AND VIRGIN BIRTHS - - -The Courtship of the Cuttle-fish—The Sumptuous Cradle of the -Argonaut—The Love-darts of the Snail—Hermaphrodites and the Dangers -of Self-fertilization—Oysters and Beauty—Sex reduced to its Lowest -Terms—Parthenogenesis and Virgin Birth—The Story of the Hive-bee—The -Departure of the Queen—The New Queen and her Marriage-flight—The -Celebration of the Nuptials and its Surprising Sequel—The Widowed Queen -turns Executioner—The Queen as Mother—The Queen’s Daughters—Nursemaids’ -Duties—Change of Work—The Drones and their Career—Food and Sex—The -Bumble-bee and its Life-story. - -That the psychical emotions sway the goad of sexual instincts in the -higher animals there can be no doubt; and there can be as little -uncertainty that this stimulating and controlling factor gradually -loses force as we descend in the scale of animal life. Just where -it ceases it is impossible to say. A vague, nebulous intelligence -doubtless persists after these more subtle emotions have ceased, and -this, probably, in turn, gives place to purely instinctive behaviour. -These various phases of the sexual problem grade one into the other. -But they are all parts of a continuous sequence, beginning, apparently, -in relatively simple responses to chemical interactions of the kind -known as chemotaxis and ending with the passion which, in the human -race, may become a consuming fire, purifying and ennobling, or exactly -the reverse—according to the nature of the inflammable material. That -is to say, in the phenomena of sex one sees emotions in the making. The -begetting of children becomes the underlying goal of life, the hidden -heart and soul of animated nature. - -This being so, one cannot but feel surprised at the discovery that, -in certain groups of the animal kingdom one meets with a strange -exception to this great rule. And this is furnished by the phenomenon -of parthenogenesis, wherein sexual desire has been dethroned. -Offspring result from Virgin births: parental care is non-existent. -This anomalous condition must be regarded as an offshoot of the -normal course of events traced in these pages, and not as a primitive -condition. This interpretation seems to be shown clearly enough in -that almost every case where parthenogenesis obtains, males, sooner -or later, make their appearance—periodically or sporadically. Every -stage between the normal, seasonal appearance of males and their entire -suppression can be traced, and an analysis of these cases demonstrates -unequivocally the uplifting character of the bi-sexual state, if only -by the fact that the uni-sexual condition makes no demands on the -parent, and does nothing to foster the growth of the higher emotions. - -No attempt need now be made to discover the origin of parthenogenesis. -Let it be assumed, for the moment, that it is a condition derived from -hermaphroditism, wherein each individual is monœcious or bi-sexual. -In all diœcious or uni-sexual animals, that is to say, where the -individuals composing the species are either male or female, each -contains a leaven of the opposite sex, even when adult. It is still -a moot point whether, in the earlier stages of development, chance -decides whether the sex shall be male or female, or, at any rate, -whether the growing body is potentially male or female, till the die is -cast by some as yet undiscovered factor; or whether this is determined -from the very beginning of germinal life. In many of the lower animals, -as among the Mollusca and some of the insects, each individual is as -much male as female, and it is from a condition such as this that -parthenogenesis probably had its rise. - -These two groups are selected here because they, more than any others -in like case, afford some extremely interesting gradations in this -strange phenomena of what is to be regarded as the degeneration of -sexual individuality, for each contains some members wherein the sexes -are separate, and in these cases sexual desire is present in varying -degrees. In some it is associated with very remarkable phenomena. - -Among the Mollusca the Octopuses afford one of the most striking -illustrations of such phenomena. In these creatures one of the -sucker-bearing arms is more or less completely transformed to subserve -the ends of sexual congress. Without entering into the technical -details of the changes, it will suffice to remark that it is modified -in such a way as to allow the transference of the spermatozoa from -the body cavity wherein they are formed, to the arm near, or at, the -tip of which they are stored in a special sac or “spermatophore,” and -such modified arms are said to be “hectocotylized.” This extraordinary -modification attains its maximum development in the celebrated -Argonaut, and one or two of the more typical Octopuses. In the Argonaut -this arm does not make its appearance until sexual maturity has been -attained, when a large more or less globular swelling appears, -enclosing the third arm of the left side, coiled upon itself. Having -attained its full development the sac bursts and releases the arm. The -folds which formed the sac now bend back to form a new receptacle into -which the spermatophore is passed. But this is not all. The tip of the -newly released arm bears another sac, which sooner or later bursts, -forming a long, slender penis, and along the central tube of this the -spermatozoa pass from the spermatophore to their destination. Their -conveyance thereto forms the last and most amazing feature of this -strange history. The male, eager with pent-up desire, and glowing with -all the colours of the rainbow, gradually approaches the female of his -choice, who apparently awaits him with no little palpitation, and then, -with a sudden rush flings himself upon her, and apparently thrusts -the penis into her mantle cavity, when at once the whole arm breaks -off from his body and remains attached to her person, retaining its -vitality, strange as it may seem, for some considerable time, during -which, no doubt, the spermatozoa are slowly making their way out of -the spermatophore and along the channel prepared for their reception. -That the Cuttle-fish are polyandrous there seems to be little room for -doubt, inasmuch as no less than four such detached arms have been found -beneath the mantle of one female. With the majority of the Cuttle-fish -and Octopus tribe the arm is not detached, but when it is so, and this -occurs in all the species belonging to three different genera, a new -arm is grown. - -[Illustration: Plate 41. - -SOME REMARKABLE METHODS OF “COURTSHIP.” - -1. The female Argonaut and her egg-casket. - -2 and 3. The male Argonaut and his “hectocotylized” arm. - -4. A Cuttle-fish (_Ocyhöe catenulata ♂_), showing the “hectocotylized” -arm described in the text, and the “spermatophore” at the base of the -long filament. - -Face page 268.] - -As a rule, among these animals the males are smaller than the females. -In the case of the Argonaut there is a yet more striking difference, -for the female possesses a very beautiful shell in which she carries -her eggs. This remarkable cradle, translucent and beautifully -sculptured, she attaches to her person by means of a pair of arms which -are expanded to form great lobes, almost but not quite completely -covering the shell. The earlier naturalists believed that this shell -served as a boat, and that the lobated arms were spread as sails! This -supposed fact naturally caught the fancy of the poets, who seized upon -it to point a moral and adorn a tale. Byron celebrated these imaginary -feats of seamanship in the familiar lines: - - The tender Nautilus who steers his prow, - The sea-born sailor of his shell-canoe. - -and Pope bids us: - - Learn of the little Nautilus to sail, - Spread the thin oar, and catch the driving gale. - -Sir Richard Owen years ago, however, dispelled these pretty fancies, -though the facts are surely as wonderful as the fables they have -replaced. They afford, too, one of the most striking secondary sexual -characters to be met with among the Mollusca; nowhere else, indeed, -among the members of this group is so strange a cradle to be met with. - -But little, unfortunately, is known of the behaviour of these animals, -which are by far the most active of the Mollusca, and which also -display no small degree of intelligence. Their eyes, which are of great -size and complex structure, are undoubtedly far more effective organs -of vision than are possessed by any other Molluscs. It is possible, -therefore, that the sexes discover one another by sight; and it is -certain that something in the nature of a “Courtship” takes place. -The majority of the species, also, possess the most extraordinary -powers of changing their coloration, especially during moments of great -excitement. The magnificence of the hues which succeed one another, -like a series of variegated blushes suffusing the whole body, may be -one of the weapons in the armoury of Cuttle-fish love-making. In how -far the “courtship” of the Cuttle-fish resembles that of terrestrial -animals, however, is a matter on which at present nothing is really -known. That even the comparatively sedentary species, like the Octopus, -seize upon and hold territory is very improbable, for there is no -need of such landed estates, inasmuch as the offspring are not tended -and fed by the parents—this would indeed be a laborious task in the -case of some of the “Squids” which lay between thirty thousand and -forty thousand eggs! Having regard to the fact that the records of -the reproductive habits of the Octopus tribe date back to the time of -Aristotle, more than two thousand two hundred years ago—for he first -drew attention to the hectocotylized arm—it is curious that so little -has been gleaned during this vast space of time. - -There are facts in regard to the sexual relationships of some of the -Snails that are in nowise less remarkable than those just related of -the Octopus tribe. Unlike the Octopuses, the Snails are hermaphrodite, -nevertheless sexual congress takes place as with unisexual species: -the eggs of the one being fertilized by the spermatozoa of the other. -During this process the orgasm of the sexual act appears to be brought -about by stabbing one another by means of a little dart formed of -carbonate of lime, the dart burying itself in the flesh and apparently -promoting a pleasurable, tingling sensation in the course of its -journey. Speedily, no doubt, it becomes absorbed, the material being -then available for the formation of a new dart. - -This remarkable instrument, which is known as a “Love-dart,” or -_Spiculum amoris_, assumes a different form in each species in which -it occurs. In some the shaft is ridged like a bayonet, as in the case -of the Garden Snail, in others the form assumed is that of an awl. -These darts are formed within a special receptacle, or “dart-sac,” but -so far no explanation as to the origin of these remarkable structures -has even been hinted at. They do not seem to have been derived by the -modification of some pre-existing organ serving a different function, -as wings, for example, are derived from walking limbs, or as lungs are -derived from air-sacs. Their origin is as mysterious as their use: -for they are not found in all Snails, though they occur in one or two -Slugs—which are degenerate Snails. But no other Molluscs save the -Snails and one or two of their immediate allies are so armed. - -The hermaphrodite conditions of these animals, as with other Mollusca -in like case, present some knotty points for consideration, and -especially in regard to the problem of sex-attraction. Where each -individual is as much male as female, which is the dominating factor in -desire, the maleness or the femaleness? Though each individual contains -both ova and sperm cells, probably these ripen at different times, to -avoid danger of self-fertilization. In this case the sex impulses are -on the same footing as in the case of animals wherein the sexes are -not thus combined. That is to say, the individual which is for the -moment only potentially male mates with another for the moment only -potentially female. But this being so, how does each discover the -condition of the other? - -Many of the Snails, like _Helix nemoralis_, are gaily coloured. Are -these hues, these bands of black and yellow, the product of “sexual -selection”—the outcome of a process of selection from among the most -conspicuously coloured individuals as postulated by the Darwinian -theory of Sexual Selection? If so, then this choice must be regarded -as a periodic recurrence coinciding with the period during which the -individual is dominated by its female attributes. In due course it -becomes, for the time, a male, and may find itself rejected, owing -to a lack of intensity in its coloration, or, on the other hand, it -may vanquish a rival by its very splendour. Each, in short, would -help materially in this process of beautification. If the choice of -mating for it is this rather than a choice of mates—proceeds on these -lines, the bright coloration of the members of this species becomes -easy to understand. But does it? It is more than doubtful whether the -eyes of Snails are sufficiently good to distinguish the coloration -of their neighbours’ shells, or for the matter of that of their own, -for their eyes being carried on long mobile stalks, they should have -no difficulty in contemplating their own charms. And what of Snails -of more sober hues? It seems highly probable that here, as in so -many cases, scent is the selecting factor, and the coloration is an -“accidental” feature. That the colour of the shell plays no such part -as that just postulated may be gathered from the evidence afforded by -many marine species, whose shells, though conspicuously marked, are, -during life, completely enveloped and concealed by the all-investing, -fleshy mantle. In like manner the exquisite beauty in the form and -sculpturing of the shell which so many species exhibit, are characters -which cannot be regarded as due to sexual selection. - -As touching the danger of self-fertilization to which reference has -been made. That this is real is shown by the fact that the ova and -spermatozoa are rarely ripe in one individual at the same time. -However, among the pulmonata, or air-breathing gastropods, it seems to -have been established that self-fertilization can, and does, occur. -That in some species, at any rate, where cross-fertilization, for some -reason, is impossible, the individual thus isolated can store up its -own spermatozoa to be used in fertilizing its own eggs. But the fact -that this rarely happens is testimony enough that such occurrences are -inimical to well-being. - -The Lamellibranch, or bivalve Mollusca, _e.g._, Oyster, Mussel, and -Cockle, afford valuable evidence as to excrescences and extravagances -of growth which appeal to our eyes as ornamental, and therefore likely -to be due to the influence of sexual selection. And this because such -ornamentation is a very conspicuous feature among these animals. Yet, -save in a few cases, locomotion is impossible, and sight is wanting. -Light-distinguishing organs, and therefore eyes, are possessed by some, -but in no case probably are they strong enough to appreciate form. -Even if they did, such revelations of beauty would play no part in -mate selection from among the most ornamental; for these creatures are -commonly fixed throughout life in one position, often, indeed, buried -in mud or sand. Some move laboriously: a few, like the Cockles and -Pectens, swim by rapidly opening and closing the shell. The Pectens -are brilliantly coloured, not only as regards the shell, which is also -beautifully sculptured, but the foot also is of a vivid scarlet, and -the Pecten have numerous minute eyes. But the Cockles and Mussels -possess like attributes as to colour and sculpture, yet they are -blind. More to the point is the fact that these animals do not mate -after the fashion of higher animals, but the males, where the sexes -are distinct, discharge immense quantities of spermatozoa into the -water, and these find their way to the ova of the female through the -action of the inhalent currents set up by the animal for the purpose of -drawing in fresh supplies of water containing food and oxygen. There -are no “secondary sexual characters,” that is to say, that even where -the sexes are separate, and many, like the Oysters, are hermaphrodite, -they are externally indistinguishable. Nevertheless, many, as has been -already remarked, have shells of great beauty. As, for example, the -giant Tridacna and the strangely spinous valves of the “Thorny Oysters” -(_Spondylidæ_). - -The fact that the Lamellibranch, or bivalve molluscs, are far less -numerous in point of species than the univalve tribes is accounted for -by the fact that in the first place they are of necessity aquatic, and -in the second their means of locomotion is extremely limited. Some few -species swim spasmodically: some crawl: many are incapable of movement -when once the motile larva settles down and the shell-bearing adult -stage is attained. Such species can extend their range only by means -of larval wanderings. Enormous numbers, millions, of young have to be -produced and set adrift each year by every adult in the community, and -yet but a few of each brood can ever attain to maturity. Life, for such -species, must be a dull, monotonous business: the only opportunity for -excitement is that which is preliminary to being eaten, and the only -purpose in life is to be eaten. But happily Oysters don’t think. They -and their kind are simply semi-conscious living things, responding -mechanically to stimuli. Any approach, then, to beauty, either of form -or coloration, or both, must be regarded as due to innate, inherent -changes in the germ-plasm affecting the parts so made conspicuous: -the only form of selection to which such “ornaments” can be subjected -is Natural Selection. If, and when, such ornaments penalize their -possessor either by their cumbrousness or their conspicuous characters, -or by increasing the difficulty of feeding or distributing offspring, -then the further development of such excrescences is checked by the -death of all individuals which have passed the bounds of endurance in -this respect. - -Sex, and all that appertains thereto, in short, is in these creatures -reduced to its lowest terms. There are not wanting, to-day, both men -and women, who affect to believe that all would be well for the human -race could a similar slowing-down, or strangulation, of the sexual -instincts be brought about. Such blind leaders might profitably -contemplate the Oyster: but such contemplation, to be profitable, -requires intelligence of a higher order than these protagonists of -folly appear to possess. - -In justice to Darwin it should be remarked that he himself fully -realized, and carefully points out, the inconceivability of the -application of the Sexual Selection theory to the Mollusca. In -commenting on the beauty of colour and shape which many species -display, he remarks: “The colours do not appear in most cases, to be -of any use as a protection; they are probably the direct result, as in -the lowest classes, of _the nature of the tissues_[1]: the patterns and -the sculpture of the shell depending on the manner of growth.” Just -so: and this is surely the fundamental explanation of ornament, using -this term in its widest sense, everywhere in the Animal Kingdom. The -peculiarities and eccentricities of behaviour, which occur among the -higher groups, act as “aphrodisiacs” to hasten reproduction because -this confers an advantage, the earliest to produce offspring—so soon as -the conditions for their nurture are favourable—having the best chance -of survival. Premature sexual activity is checked by the death of the -offspring. - -[1] Italics mine. - -It has been contended that the hermaphrodite condition represents the -primitive mode of reproduction among the multicellular animals—that is -to say, all animals above the level of those whose bodies are composed -of but a single cell, or particle, of protoplasm—but this view is -probably erroneous, and the hermaphrodite state must be regarded as a -secondary condition, a later innovation. - -More remarkable are the facts concerned with that singular form of -reproduction known as parthenogenesis, or the production of offspring -by virgin females. This is undoubtedly a degenerate sexual condition -occurring as a normal mode of reproduction, among the microscopic -“Rotifers,” _e.g._ the “Wheel-animalcule,” Crustacea, and Insects, and -in varying degrees of intensity. - -The most familiar instances of Parthenogenesis are furnished by the -Hymenoptera, and notably by the Bees and the Aphides. - -There are certain cases among the Rotifers where no males have ever -been found, and it is possible that they have become entirely -suppressed, but in every other case the periodical advent of males is -an absolute essential for the continuation of the race. Perhaps the -least degenerate of these types are the Bees, wherein we meet with -well-developed, highly-organized males and females, which, in their -sexual relationships, are perfectly normal. But in the fulfilment -of the mating instincts in these insects, a most amazing sequence -of events is revealed such as are without parallel in the rest of -the Animal Kingdom. The story has been charmingly told already by -Maeterlinck, in his delightful “Life of the Bee,” and it has been -told again by Tickner Edwardes, with less of poetry, perhaps, but -still fascinatingly: and it must be told again now, but in a condensed -fashion. - -Briefly, a community of hive-bees harbours both male and female -individuals only for a very short space. During the greater part of the -year it consists only of a vast concourse of infertile females, the -daughters of one mother; the “queen” of the hive. The males of that -hive are the brothers, not the fathers, of the workers, as some have -supposed, and their sojourn there is brief. To gain a clear idea of the -facts in regard to the life-history of these insects it is necessary -to trace some of the incidents which lead up to the manner in which -the population of the hive is regulated, and its continuance ensured. -These may well begin with the time when the number of the inhabitants -consonant with the well-being of the hive has reached its limit. This -occurs during the early part of June, when the queen leaves the hive, -accompanied by several thousands of her daughters; they settle at -some distance from their late abode in a “swarm” for the purpose of -founding a new colony. Here we may leave them. The house just vacated -is, however, not entirely deserted. A few of the inhabitants, the -infertile sexless workers, degenerate females—degenerate so far as -the power of reproduction is concerned at any rate—are left behind, -and there remain also in their cradles a variable number of unhatched -queens, and drones or males. One of these potential queens and the -males now speedily emerge, and for a day or two remain within the -seclusion of the hive, feeding upon the honey stored in the combs. - -The males are the first to leave, making daily excursions abroad in -the search for mates. They display in this a very leisurely behaviour, -rising late and not venturing out till the day is well aired. Returning -early in the afternoon with sharpened appetites, they feed to repletion -and soon fall asleep. - -In about three days, however, the young queen ventures abroad, timidly -at first, to stretch her wings in the sunshine. She is preparing for -the great moment of her life, the nuptial flight. So far, though drones -may swarm on every side of her, no sign of recognition is given, nor do -the males evince any consciousness of her presence. She behaves warily -and demurely throughout. Her first excursions abroad are very brief; -they are not so much trial flights, apparently, as efforts to locate -the exact position of the hive in relation to the outer world. To this -end the flights are rapidly extended in ever-widening circles, till at -last, with lightning speed, she makes for the blue sky, to return to -the gloom of the hive almost immediately after. During all this time -the stimulus of sexual desire has been gathering force, and now, being -no longer controllable, she darts off, and up into the sky; almost -at once she is recognized by the swarms of males from neighbouring -hives, some thousands in number, which for days have been seeking this -event. Instantly they give eager chase, mounting after her higher and -ever higher. But as they ascend so their numbers decrease. Some, the -feeble, the ill-fed from impoverished hives, are speedily left behind; -many endure to the end, but only one secures the prize, and this great -moment of his life is also his last, for the fact of impregnation is -no sooner completed than Death claims him. He falls earthwards, as if -struck by lightning, and in his fall the intromittent organ is dragged -from his body, to be removed by the survivor of this mad flight, on her -descent. - -She leaves a bride and returns a widow, filled with murderous -intentions. There are captive queens in the hive, and she can -tolerate no rivals. So soon as she has removed from her person the -embarrassing souvenir of her nuptial flight she makes for the Royal -cells. Accompanied by attendant workers she proceeds to tear off their -waxen coverings and put their occupants to death with a thrust of her -stiletto. No sooner is the work of execution over than the dead bodies -are seized by the workers and borne out of the hive. This awful task -is soon over, however, and henceforth for four or five long years she -remains a prisoner within the walls of her own palace. Craving neither -the air nor the light of the sun, she will die without once having -sipped the nectar from a flower. And during all this time, save during -the winter sleep, her sole duty is to produce sons and daughters. In -the prime of her maternity she may lay as many as three thousand eggs -a day. But strangely enough the number of eggs produced is determined -for her by the workers, who are the real rulers in this constitutional -state. By varying the amount and quality of the food they give her they -can increase or check the number of eggs produced; while even the sex -of the resultant larva is apparently also under their control. - -During that brief, weird honeymoon in the clouds she received a store -of spermatozoa, the fertilizing male germs, sufficient for all the eggs -she can ever lay, and they may amount to nigh on a million. Incredible -as this may seem, their purpose is yet more so; for they are destined -to be expended solely in the production of female offspring doomed for -the most part to perpetual spinsterhood. One youngster in ten thousand -may attain to a higher state, may, if Fate wills, become a queen and -mother. And because of this need for mothers to carry on the race, this -extraordinary state of affairs has been brought about. All is under the -control of her daughters—the spinster-workers. As she proceeds on her -rounds of egg-laying an attendant crowd waits upon her, controlling -her actions by gentle caresses. As she passes from cell to cell, the -cradles of the young that are to be, she thrusts down her abdomen and -lays an egg in each. The cells destined to produce the workers are -the smallest, those for drones are larger, and those for queens are -largest of all, and the walls are formed of pure pollen, not of wax as -are those of the workers and drones. But it would seem that she never -lays an egg in any of the last named. The sight of a queen-cell rouses -her to fury. These cells, then, are filled by the workers, who remove -the requisite number of worker—eggs from the cells in which they were -laid and deposit them in the queen-cradles. The larvæ at hatching, -and for the first three days of life, differ in no wise from their -sisters around them. Their Royal state is determined solely by the food -which is administered to them. This consists of “bee-jelly,” which is -furnished in abundance: a white, shining liquid, regurgitated by the -ever-zealous nurse-bees. These superfed babies cease feeding at about -the fifth day, and each spins for herself a silken vestment in which to -undergo the pupal state. This done, the door of each cell is sealed up -with pollen. During the following sixteen days strange transformations -take place: the queen that is to be is taking shape. But the cradle now -becomes a prison, for at the end of the sixteenth day each of the four -or five young queens begins to clamour for release. But this cannot be, -for such as succeeded in emerging would immediately be slain by the -reigning queen. A small hole is bored through the roof of the cell, and -through this each is fed, and a close guard is kept night and day to -ensure that they shall not emerge till the moment is ripe. Soon each -captive begins to gnaw away the roof of her prison chamber, and as -rapidly more material is placed by her guards on the outer surface. Not -until the old queen leaves the hive with thousands of her daughters to -“swarm” and found a new colony will freedom be allowed; and then only -to one. The rest must remain till the new queen either also “swarms,” -or returns from her nuptial flight, and in this case all will be -slaughtered in their cramped quarters, unable to resist. - -But what of the drone? He, as has already been mentioned, is reared -in a larger cradle than that of his sisters—save such as are destined -to be queens—and for the first three days of his life is fed on -“bee-milk” of a special kind and more generous quality than that of his -worker—sisters, the Cinderellas of the hive; but this generous diet -is diminished at the end of three days, when a mixture of honey and -pollen is given him. In about three weeks or rather more he emerges, a -great, lazy drone, and for a fortnight more he wanders about the hive -alternately soliciting bee-milk from his sisters and helping himself -to honey from the comb, and when full to repletion he seeks some -snug corner in which to sleep off his surfeit. In due time, however, -he ventures abroad, his hour is at hand. He takes his daily flights -abroad in search of a mate, returning home early in the afternoon for -his rations, being too indolent or too stupid to draw nectar from the -flowers for himself. Thus for many days he and his brothers disport -themselves in riotous living, till one or other of them attains the -end for which he was born; and after a few delirious moments drops -earthwards a mutilated corpse. - -But so far only a part of the story of the drone’s life-history has -been told. Though the son of a queen, he has never had a father; and -should he ever attain to the dignity of fatherhood his posthumous -children are all daughters, most of whom die spinsters within six or -seven weeks of their birth, worn out by a life of ceaseless toil and -drudgery! - -The queen, it will be remembered, cohabits with the male but once -in her life. The sperm-cells then received are stored in a special -receptacle and are released during the passage of the egg down the -oviduct. In this act of releasing the fertilizing germs a singular -economy is practised. In the case of most other creatures myriads of -sperm-cells are released for the fertilization of a single egg, and -of these but one can possibly attain its goal, the minute aperture or -“micropyle” which is the doorway to the germ liberated, in the form -of an egg, by the female. The rest die. In the case of the queen bee -but one of these precious sperm-cells is liberated at a time. Hence -her prolonged ability to produce fertilized eggs. But eggs destined -to produce males, or drones, are never thus fertilized: they are born -without the intervention of a father. A queen which has never mated -will lay only male-producing eggs. This is an astounding thing, but it -is true. No less remarkable is the fact that the sperm-cells should -survive in their encapsuled state for periods extending over several -years: it seems almost incredible, but it is nevertheless true. - -One cannot suppose that the queen in coming to a drone cell -deliberately withholds the male germ as the egg passes down her -oviduct; some inhibitory factor preventing the release of the -sperm-cell must be brought into play which as yet we have not -discovered. This production of males from unfertilized eggs, or -“parthenogenesis” as it is called, is a common feature among the -hymenoptera, and some other groups of insects, and it occurs also among -other lowly creatures to be described later. - -Having regard to the importance of the workers, a brief summary of -their life-history must be given. These, it has already been indicated, -are all, at any rate till three days old, potential queens. Their -development into, or degradation to, the lower grade is determined, -apparently, solely by the quality of the food, for the fact that -queens are reared only in specially constructed cells of large size -with walls of pollen instead of wax is explained by the larger size -of the queen and the need for a more porous, air-permeated cell-wall -on account of the longer time which must be spent in confinement. -The worker is certainly the most “intellectual” member of the hive, -but this superiority has been gained at a great price. Emerging from -the chrysalis skin at about three weeks from the time that the egg -from which she emerged was laid, she begins forthwith to gnaw her way -through the mass of wax and pollen which forms the door of her prison. -Rather, she eats her way through, for the material removed is swallowed -as it is detached, thus the young bee, as Mr. Tickner Edwardes remarks, -is caused to effect her own release by the promptings of her appetite. -Hunger-strikes in the bee community are unknown. Speedily the youngster -steps out, distinguishable from her elder sisters only by her weak, -grey-hued, flaccid appearance. Her first act on gaining freedom is to -groom herself down, after which she proceeds to explore the gloomy, -busy, crowded thoroughfares of the hive. A day or two is thus passed in -gathering strength. On the second appetite returns, and she proceeds -to help herself from the vats of honey and pollen bins scattered here -and there among the cradles of her sisters yet prisoners. But speedily -she is caught and thrust, so to speak, on to the treadmill of work -which is to know no cessation during her short span of life-some six -or seven weeks. Her first duties are those of nursemaid. Without -instruction, or previous experience, she begins to feed her younger -sisters and brothers yet in the larval stage. But besides, during her -first fortnight, before she is allowed to leave the hive she and her -sisters of the same age have to fulfil a variety of tasks. All the -indoor work of the house falls on these Cinderellas. Not only do they, -and they alone, feed the young, but they have to produce the wax and -build the combs and attend to the sanitary arrangements: “they are the -brewers of the honey and the keepers of the stores; they feed the queen -bee on her ceaseless rounds and give the drones, their brothers, their -daily rations of bee-milk”—what else these lazy creatures need they -take for themselves from the honey-vats. But this is not all. They have -to meet their older sisters returning from the fields and gardens laden -with nectar. This is regurgitated and transferred to the pouches of -the youngsters, by whom it is transformed into honey and stored in the -combs in the upper region of the hive. At the end of about a fortnight -these little drudges are allowed a brief respite, during the heat of -the day, to emerge into the outer air and gather ideas on the world -which is yet to be explored. Soon a measure of freedom is allowed, the -indoor work ceases, and each takes up the new and more agreeable task -of gathering pollen, and after a few days of this the more responsible -task of gathering nectar is undertaken, which is continued till death -ends one of the most crowded, surely, of existences. Such as are born -near “swarming-time” may have the good fortune to take part in the -exodus and the settling down in the new home, and some may taste yet -other moments of excitement, but they are moments only. The worker -bee knows no leisure for the improvement of her mind and morals. She -needs none, for she has neither: she is a creature of routine, a -living automaton apparently. Yet there are incidents in this wonderful -community which seem too complex to be merely the result of instinct -unaided, uninspired, by intelligence albeit of a nebulous kind. - -The worker-bees, it has been remarked, are barren: their reproductive -organs are atrophied, and by the decree, not of the queen-mother of the -hive, nor of the males, but of their own caste. In spite of the fact -that they are incapable of producing offspring, they, and they alone, -determine who shall undertake this task; and they decree the fate that -awaits those thus appointed when they can no longer fulfil this purpose. - -When the queen, waxing old, and waning in fecundity, lays fewer -and fewer eggs, and these only producing males, they take silent -note of the fact, and at the appointed time decree the death of -their Sovereign-mother. Yet they hesitate to lay violent hands on -her. She, as queen, claimed the right in her early youth to slay -her sister-queens, and sped them with a dagger-thrust; now her turn -comes to die. But it must be a bloodless death, carried out with due -ceremonial. So her daughters cluster about her, and in a mock embrace, -that tightens every moment, her breath is squeezed out of her body. -There are no State pensions for those who are past work, but a State -execution instead. This is vastly more economical, and it may yet -commend itself to some would-be social “reformers,” who will doubtless -contrive to make exceptions to the rule! - -The execution of a queen is not an event of common occurrence; but -that of male members of the hive forms part of the ordinary routine, -though coming only within the larger cycle of the year. As the summer -wanes and the harvest of nectar grows perceptibly less, visions of a -possible famine, and its attendant horrors, seem to arise. So heads are -counted and occupations are scrutinized, when it is discovered that -the only members of the community who are contributing nothing to the -general well-being are the males, who are now but useless drains on -the hive. None of the neighbouring hives are now likely to send forth -a virgin queen to her nuptials, to which end each hive is obliged to -contribute—for no hive utilizes the services of its own drones; these -idle fellows, then, are “eating their heads off”—and males, too; perish -the thought! While they had anything to gain from him their motto -was “Feed the brute”; but now, on each, doom is pronounced. It must -be admitted that a live drone at the end of summer is one of life’s -failures. Notoriously unable to feed himself save upon the honey made -by his sisters, and having no function in life to perform save that of -mating, his very existence now is a damning witness against himself. - -When the mother of the hive ceases to maintain the standard of -fertility set by her exacting daughters, she is put to death -stealthily, as if in an excess of devotion: she is smothered under -their embraces. Towards the drones now under sentence no such -consideration is to be shown. When the word goes forth, the slaughter -begins, and it gathers in ferocity. It begins in a massacre of the -innocents—every helpless larval drone is ruthlessly dragged from -its cot and thrown out of the hive to die: there is now no crime in -infanticide, nor in the most gruesome massacre that is presently to -follow. The drones, all unsuspecting, are to be tolerated a brief spell -longer. The cool, calculating spirit of these unsexed ones seems to -realize that there is even yet a remote possibility that the services -of these doomed ones may be wanted. No sooner, however, does it become -clear that this chance is past, than the decree of death is made -absolute, and the poor drones are suddenly and viciously attacked by -half a dozen frenzied spinsters at once. Each tries to bite through -the base of the victim’s wings, and succeeding in this, he is speedily -pushed towards the door of the hive and out into the open, whence -return is impossible, so that nothing is left but death by starvation. -Some of the victims will escape in the _mêlée_, but only for a brief -season. Such as find their way, unmaimed, to the open air, are still -faced by inevitable death. To remain out is to die of starvation or -cold, to return is to fall a prey to the now infuriated guards, who, -strongly reinforced, stand at the doorway of the hive to intercept -and dispatch these unlucky fugitives. It will be remarked that these -executioners make no use of their stings; these they might be unable to -withdraw from their victim’s body, in which case they, too, would die. -But there is no need to run this risk, for the males, their brothers, -whom they so cheerfully slay, are unarmed; they may be attacked without -risk. The dreadful work, however, is soon over, and the survivors, the -queen and her daughters, have the house to themselves to make the final -preparations for the winter sleep, which is apparently undisturbed by -qualms of conscience. - -There are certain structural differences distinguishing the three types -in such a hive—the queen, the drone and the worker—which must now be -referred to. The queen is larger than the worker; she has a larger -and longer abdomen, a longer and much-curved sting, and her eyes have -fewer facets. Only vestiges remain of the wax-secreting organs, and no -trace is to be found of the wonderful pollen-baskets which perform so -important a function in the worker; and finally, her instincts are of a -very different kind. - -The “pollen-basket” of the worker is a strange contrivance. The pollen -is mainly collected by the hairs which clothe the under surface of -the body, from which it is scraped by special brushes of hairs which -clothe the inner surface of the “metatarsus “—the big, flat joint to -which are attached a series of small triangular joints, the last of -which bears the claws. When the brushes are “clogged up,” the legs are -crossed and the pollen is combed out by specially stiff hairs on the -“tibia”—the joint immediately above the metatarsus—and the bolus thus -formed is then transferred to the outer surface of the tibia, which is -trough-shaped, forming the “corbiculum,” or pollen-basket. The next, -or middle, pair of legs are then employed to ram the pollen well into -the basket, for safe conveyance to the hive. On arrival at the combs, -the bee pushes its hind-legs into a cell, or “pollen-tub,” and with a -special spur dislodges the pellet of pollen and lets it fall into the -tub. These are complex movements, performed without instruction and, we -must suppose, without any intelligent conception of their purpose. - -The drone is larger than either queen or worker, and has enormous -eyes, which meet one another over the top of the head; he has no -wax-secreting organs, no pollen-basket, no sting. His antennæ are -longer, his hum is deeper, his sole function is to fertilize a queen, -and this done, he promptly dies. Failing in his first flight, he may -make yet other ventures, but the chances are that he will die without -attaining the only purpose for which he exists. - -The fact that he lives for some days in the hive with the queen, -before her nuptial flight, apparently unaware of her presence, would -seem to indicate some special “trigger” for the release of the sexual -instincts. But it must be remembered that he does not attain to -maturity until after his first flight, and this it is, probably, which -arouses the mate-hunger. More than this, however, it is probable that -coitus is possible only when on the wing, when the air-sacs become -inflated, and exert pressure on the genital organs. How he recognizes -the queen when on her wild flight heavenwards is unknown: possibly by -scent, but more probably by the very different vibrative note of her -wings, that of the male being much stronger and deeper. His continued -return to the hive is a proof of his failure to justify his existence, -for no drone ever experienced Love’s embrace and lived to tell the -tale: hence, when the time comes, he is slain without compunction. - -These differences between the fully-developed male and female present -nothing very striking; but how are the singular peculiarities of -structure and instinct in the “workers” to be accounted for? They are -present in neither queen nor drone, yet by them they are transmitted -to their offspring from one generation to another! It is true that -every worker, for a time, is a potential queen, and every queen, but -for the grace of Chance, might have been a worker. All depends on the -food. It is remarkable, but apparently the fact, that a more generous -diet, or, rather, a more stimulating diet, should so profoundly modify -the organism, but, it is to be noted, this sleight-of-hand is only -successfully practised on a larva during its first three days of -existence. Thus the royal bee jelly stimulates the growth of the sexual -organs and inhibits the development of the structures peculiar to the -worker—the basket, and pollen-hairs, and so on. These structures are -not made by the food; they are simply nourished or inhibited, as the -case may be. Nevertheless, one cannot help being mystified by the fact -that the mere difference in the quality of the food, or, rather, in the -chemical constituents thereof, should cause the inhibition, or, rather, -the suppression, of relatively complex structures like the corbiculum -and the reduction of the number of the facets of the eye. To say that -the structures inhibited, in the case of the queen, are just those -which will be of no service when in her royal state, is by no means to -explain the mystery. And what is true of the physical side is no less -true of the psychical, for with this change of diet the behaviour of -the insect, throughout its whole life, is most profoundly changed. If -the pollen-basket is wanting, no less so are the instinctive actions -associated with its use; if the genital organs are atrophied, so also -are the instinctive acts associated therewith. This nexus between -instinct and structure is not to be lost sight of. - -How—and the question has often been asked—are the experiences of the -infertile females, the workers, transmitted to the germ-plasm? For the -workers, it has been contended, being sterile, are incapable of handing -on such acquirements: this is so. These workers hold the same position -in regard to the species that structures essential to well-being hold -in regard to the individual. These last are not under the control of -the individual, but are determined by a plus or minus quality in its -germ-plasm. The worker-bees are products of the germ-plasm, committed -to the care of the queens. Any strain, so to speak, of that germ-plasm -which gives rise to defective workers brings about its own extinction, -or elimination, sooner or later. Any strain of germ-plasm which -contains, so to speak, a spark of that quality which in the individual -is expressed by intelligent behaviour, will gain advantages in the -struggle for existence. - -The complex, the extraordinarily complex, behaviour of the worker-bees -on any interpretation is still mysterious. This interpretation can be -tested only by a reference to the life-history of other social-bees -which have attained to a less complexity. This shows us that the -sterile worker is not to be regarded as a newly-evolved type so much as -an arrested stage of a more complete ancestral condition, and the fact -that the worker is potentially a queen is further evidence of this. - -A clue to many of the more puzzling features presented by the domestic -economy of the Hive-bee may be obtained by a study of the life-history -of other species of social-bees which have not attained to so high a -degree of specialization. The Bumble-bees afford illustrations of the -stages through which _Apis mellifica_, the Hive-bee, must have passed. - -In the stone Bumble-bee (_Bombus lapidarius_), a queen, who has passed -the winter in blissful sleep, will lay the foundation for a new colony -on some bright May morning by collecting a small quantity of moss. This -done, she starts forth to gather pollen, with which, under cover of the -moss, she forms a waxen cell, mixing the newly-gathered pollen with the -wax so mysteriously formed within her body, as in the case of Hive-bees -of the worker type. Slowly and laboriously this waxen cradle grows. -Fashioned like a globe, its inner surface is lined with pollen soaked -in honey, and with the last pellet of this a number of eggs are laid -arid the nursery is sealed up. By the time these labours are completed -the queen is worn out; she therefore rests awhile, clinging to the -outer wall of this cunningly-wrought cradle. After a few days’ rest she -adds another and commonly yet a third cell to the first, joining each -to the other with wax. But before the third cradle is finished the eggs -in the first have hatched. The youngsters will have consumed the layer -of honey-soaked pollen placed there for this purpose. They therefore -require feeding, and thus the labours of this very industrious queen -are still further increased. Divining the needs of her imprisoned -first-born, she bites a small hole through the nursery wall and pours -in a quantity of honey for their sustenance. In due time they are -“full-fed,” and each spins for itself a silken vestment wherein to -undergo its transformation into a worker-bee. The careful mother, -during this period of transition, now scrapes away an opening through -which the young bees may creep when they awake. This event takes place -in the course of a few days, when her work is materially lightened, for -these newly-hatched workers at once take over the duties of building -nurseries and feeding the further batches of young which, for a time, -follow one another in quick succession. The queen, indeed, has now -nothing else to do but to lay eggs in the nurseries as they are ready. -So far all the children born to her are daughters. The earliest-born, -it is to be noted, were “workers”; those which follow and are tended by -the workers are also females, and supplement their mother’s labours by -producing fertile eggs, though they have never even seen the male of -their own species. Thus, if the queen-mother die her virgin daughters -carry on the colony. But it sometimes happens that she may have left -no descendants capable, for the time, of laying fertile eggs. In this -case, if there be larvæ still in the nursery, the workers feed them -assiduously as if in the hope that some may prove fertile. But if -there be no infants to be fed they apparently abandon work, become -despondent, and spend the greater part of their time sitting at home -by the empty cradles, till at last death comes to their rescue and the -colony is extinct. - -Much that baffles one in the history of the Hive-bee becomes clear in -the light of the facts revealed by the life-story of the Bumble-bee. -In the first place it will be remembered her first eggs produced only -workers, which appeared at a time when her energies were severely -strained, and their food allowance was no more than barely sufficient -to sustain life. The females which appeared later produced fertile -eggs, having been more abundantly fed by their infertile elder sisters. -The number of fertile females which appear at this stage of the colony -seems again to be regulated by the abundance of food, which varies -in amount with fine, or cold, weather. Even among the worker broods -fertile females may appear. They owe their fertility apparently to good -luck, which afforded them the opportunity of securing more food than -their sisters. The birth of young from females about whose virginity -there can be no question is certainly remarkable, but it would seem -that this parthenogenetic state is one of limited endurance, for -towards the end of summer males appear, and these mating with some of -the later-born females, lead again to the appearance of a queen, who, -being fertilized, alone survives the winter to carry on the race with -the succeeding summer. - -Thus, then, the mysterious existence of the workers among the -Hive-bees, displaying structural peculiarities and instincts so -different from those of the queen-mother, is explained. For the queen, -in this case, is evidently the product of a more intensified, more -perfected, social system, relieved, from the first, of the labours -of building and the care of her offspring, duties which the queen -Bumble-bee has at first to perform for herself, because all her -children die at the end of the summer. Among Hive-bees fertile workers -also occasionally occur; they are probably bees which in their larval -state received a more than usually abundant supply of food, or food -approximating to the “bee jelly” which produces young queens. The -difference, then, between the individuals of a colony of Hive-bees and -one of Bumble-bees lies in the greater abundance of fertile workers -and in the fact that the queen of the Hive-bees is relieved of all -work from the first, and so is enabled to devote her whole energies to -the duties of reproduction. She is the descendant of a race of queens -which in earlier times, like the Bumble-bee queen, had to perform -the duties now relegated to her daughters, who inherit not only her -house-building and child-nurturing instincts, but also her potentiality -for child-bearing, though this potentiality is commonly inhibited -by the starvation of the reproductive activities. Selection secures -survival of this state of affairs by the elimination of any tendency -to lose any of these qualities on the part of the queen. The workers -of the Hive-bee, in short, have not evolved their peculiarities of -structure and instinct by some mysterious process of natural selection -confined to the workers individually, for these, being infertile, could -not transmit any of their inherent qualities or tendencies to variation -in the direction of more efficient workers. On the contrary, all that -they possess they inherit from the queen-mother, who transmits to her -offspring the qualities and characteristics her forebears in the female -line possessed in their own person. - - - - -CHAPTER XIV - -PARTHENOGENESIS AND ITS SEQUEL - - -Courtship among the Ants—The Great Renunciation—Maternity carried to -Extremes—Where Males are Superfluous—Degenerate Males—Keeping Death at -Bay—Where Females are Unknown. - -The phenomenon of virgin birth is one of profound mystery. The -existence of so astonishing a mode of reproduction was an established -belief among the ancients, though they could have had no means of -demonstrating the faith that was in them. But these men saw no -difficulty in ascribing to the females of their own race this faculty -of producing offspring without the intervention of a male. One -suspects, indeed, that there was no solid foundation whatever for this -belief in these miraculous powers: they lived in credulous times, and -the recorded occurrences of these, even to them, irregular births are -to be regarded as devised to afford a convenient means of escape from -the consequences of lapses from the path of virtue. Yet, incredible -as it may appear, there are not wanting to-day both men and women who -affect to believe that this mode of reproduction obtains still among -the human race, in certain exceptional cases; and further, they profess -a conviction that in the future it may become the normal mode, males, -in consequence, becoming unnecessary! Such professions of faith are -made only by the ignorant, or by those who trade on human credulity. -Parthenogenesis not only does not occur in the human race, but it does -not even occur in any member of the great group of vertebrates of which -man himself stands at the head, and it never will occur. - -Those near relations of the Bees, the Ants, afford a further insight -into this strange method of reproduction. Each community in the case -of these insects harbours not one, but many queens. The nuptial -flight, like that of the Bees, takes place in mid-air; but myriads -of both sexes participate therein, forming a filmy, ever-shifting -cloud, now rising, now falling, in the shimmering sunlight. At no time -do they seek to attain the altitude, or the privacy, so strenuously -striven for by the Bees. But in the case of the latter there is but -one female, and her life is precious. She must seek sanctuary for the -consummation of her marriage in the highest heavens, beyond the risk -of instant destruction by insect-eating birds; for though thousands of -suitors accompany her, she rises above them all, save one or two, and -hence would form an easy mark. With the Ants there are thousands of -queens, and the destruction of a few hundreds more or less is rather -an advantage to the species than otherwise. On their return to earth -the males die: their life’s work is accomplished. The females, or as we -must call them, the queens, on the other hand, have a long life before -them; far longer than that of the queen bee. But for them the joys of -flight are restricted to this one brief revel, for, no sooner have they -reached terra firma, than they renounce, as it were, the pleasures of -life to devote themselves entirely to the work of reproduction. And as -if to make all regrets vain, to stamp out all possible temptation to -desert their vows, they tear off their gauzy wings, and with them goes -all hope of fertile repentance: for the rest of this life their home is -underground. - -Each queen, on her descent, departs a separate way, and hard is the -road before her. She left the parental nest well-fed, and in good -liking, her body well, stored with food in the shape of fat and the now -useless, bulky, wing-muscles, and with this, her only dowry, she starts -the formation of a new colony out of her own substance. Her first -task is to form a burrow, and at the end of this she fashions a small -chamber. This done, she closes the mouth of the burrow and cuts herself -off from the world. The labour of this burrowing is so severe that it -often wears away her teeth, her only tools, and the hairs from her -body. In this retreat she now waits patiently for the eggs within her -to ripen, which may take months to accomplish: she is still fasting, -or, rather, feeding upon herself. When at last the eggs are laid and -hatched, she feeds her children on saliva, the very juice of her body, -for she is still fasting. Nor is the strain relaxed till the larvæ -undergo their transformation into pupæ, and, after a brief sleep, -emerge as “worker” Ants, puny in stature owing to the poorness of their -food during larval life. In some species this fast may last for ten -long months. So soon, however, as these little workers emerge, like -dutiful daughters they make their way to the outer world, and go forth -in search of food, which they share with their now exhausted mother. -But, besides, they enlarge the original chamber, and drive galleries in -all directions to provide accommodation for the vast population that -is soon to crowd the thoroughfares. Meanwhile the queen resumes her -task of producing more and yet more daughters, in whom she now displays -not the slightest interest. Her elder children now bear away the eggs, -and feed the young as they hatch. In course of time, as with the Bees, -the task of wet-nurse falls on the youngest of the Ants, those who have -just attained to anthood. For ten or fifteen years this queen-mother -may continue her work of reproduction, a slave, indeed, to domesticity, -with monotonous regularity, checked only by the chill of autumn and the -sleep of winter. - -Those among our own race who profess to hail the prospect of a time -when parthenogenesis shall be the normal mode of reproduction may -well take the Ant as an awful warning. Their ambitions may overreach -the mark. The poor queen becomes a slave to reproduction; children -in myriads are born to her; even if she would she could not sustain -her interest in them, she could not even recognize them as the fruit -of her body. Her daughters are born to a lifelong drudgery, her sons -are mere fertilizing agents: for their only purpose in life is to -perpetuate this awful thraldom, this appalling prolificness; and having -accomplished this, they die forthwith. If there be any joy in this life -it is drunk by the males alone. Thus does the female rule overreach -itself. It is well, indeed, that the participants of the joyous nuptial -flights dancing deliriously on gauzy wings in the glare of a summer -day, have no foreknowledge of the long night that is to follow. - -Unlike the Bees, the Ants may produce as many as five grades of -workers, each of which have different duties towards the community. But -the nature of those duties and the manner of the evolution of these -types, are themes foreign to these pages: enough has been said already -to indicate the nature of the problems they present when discussing the -life-history of the Bees. - -The subject of parthenogenesis need be pursued no further in this -volume than is sufficient to bring out its retrograde character. It -is a form of reproduction which may be limited to a small number of -generations, as with the Aphides, or to a single generation alternating -with normal sexual generations, as in many Cynipidæ or Gall-flies, or -it may be the only mode of reproduction, as in some other Gall-flies, -some Saw-flies and some Crustacea, wherein no males have ever been -seen. In some species this form of reproduction gives rise to females -only—the Thelyotokous parthenogenesis of scientific text-books—as in -the Saw-flies and Gall-flies, and the parasitic _Tomognatbous_. In -some other Saw-flies, unfertilized queens and workers of Ants, Bees, -and Wasps, which occasionally produce offspring, the progeny is always -male, and this is known as Arrhenotokous parthenogenesis. In one or two -species of Saw-fly, _e.g._ _Nematus curtispina_, both males and females -may be produced, when the species is said to be Deuterotokous. - -In the case of the Aphides, winged males normally appear in large -numbers at the end of the summer, and these fertilize the females; but -if kept in a warm green-house, parthenogenetic reproduction may be -sustained for as long as four years. Under quite normal circumstances -these tiny insects show a singular range of variability, for egg-laying -and viviparous individuals are met with; while winged and wingless -generations appear sporadically, apparently according to the abundance -of food. The winged form is sometimes so abundant as to float about -in swarms that darken the air. There are at least three kinds of -males-winged males, wingless males with a functional mouth, and small -wingless males which have no mouth, and, one need hardly say, are -very short-lived. The Aphides are a feeble folk, individually, but -collectively a power in the land, causing at times incalculable loss -to the farmer and gardener; but on this head and on the subject of -their strange habits, and sometimes adventurous lives as slaves in the -service of Ants, no more than a hint may be dropped in these pages. -But some such aids to faith seem to be necessary when those who are -not tolerably familiar with these insects are told of their amazing -fertility. Linnæus long since estimated, in regard to one species, -that in the course of one year a single Aphis will give rise to a -quintillion of descendants—all produced without the aid of a male. -Every one of these females begins to reproduce within from ten to -twenty days of her birth, but even this statement does not bring home -the result of such an astounding fecundity like Huxley’s calculation -which was carefully worked out. He estimated that the produce of a -single female would, in the course of ten generations, supposing all -the individuals to survive—and possess the normal fertility of their -race—“contain more ponderable substance than five hundred millions of -stout men: that is, more than the whole population of China.” - -To explain such a riot of reproduction one might almost suppose these -insects to be imbued with a dread of the impending dissolution of -their race, and endowed with the power to avert such a calamity by -these stupendous efforts; for it is evident that parthenogenesis -confers quite extraordinary powers of raising the birth-rate. But then -the normal mode of procreation is capable of achieving results quite -as remarkable. The queen Termite or White Ant, for instance—which, -by the way, is no Ant, but a near relation of the Stone-flies—when -in her prime will lay eggs at the rate of sixty a minute, or eighty -thousand and upwards in the course of a day of twenty-four hours. But -this unenviable mode of breaking the record is attended, surely, with -some little inconvenience; for to attain to such fertility her abdomen -increases until it attains something like two thousand times that of -the workers of the community in which she lives. That the history of -the queen Termite is unique of its kind is not surprising: indeed, such -an amazing story could only be told of creatures which enjoyed the -seclusion of a subterranean existence. Here, on a bare couch, with her -Royal spouse beside her, she lies, a bloated, heaving mass, incapable -of movement, depositing eggs with the rhythm of a machine, the mother -of offspring which she will never see. A more unsightly picture of -maternity it would be impossible to conceive: it is well, indeed, that -it is hidden from the light of day. No such state of affairs could ever -arise among creatures living an outdoor life, with enemies to avoid, -and food to find. - -The instances just surveyed, these extremes of the potentiality of -procreation, are instructive in more ways than one. They are to be -regarded as “excrescences” of reproduction, comparable to those -“excrescences” of individual growth which we call “ornament,” for -example. Individuals on whom this fertility has settled, so to speak, -are the victims of the machinery of sex and reproduction. Their -amazing powers of multiplication are not of their own seeking, they -are inherent manifestations of variations of growth, uncontrollable -save by the machinery of Natural Selection. Incidentally such victims -serve a useful purpose, for their myriad hosts afford food for hordes -of other animals, which in turn are eaten. Little though we realize -it, the well-being of the human race would suffer if these prolific -creatures—the uncomplaining victims of that inexorable law which bids -all living things “increase and multiply” or die—should cease to be; -for with them would disappear a host of animals on whose existence -man’s comfort more or less depends. - -During the millions of years that have rolled by since the first -appearance of life on the earth, who shall count the number of types -which have been exterminated without leaving the faintest trace of -their having ever existed? The survivors which have contrived to -maintain a place in the sun present an infinite range of variation -in colour, size, habit, and structure, as well as in emotions. These -varied aspects are all so many facets of the mysterious phenomenon -we call Life: and they are so many witnesses of the versatility of -Life. Not the least mysterious feature of this Life is its faculty -of reproduction, which expresses itself in an infinite variety of -ways, defying all but the crudest forms of analysis. The evolution of -sex has exercised the speculative ingenuity of some of the acutest -students of Nature from the earliest times, and we are still far from -a satisfactory solution of the problems it presents. Hermaphroditism -and Parthenogenesis are commonly regarded as degenerate forms of -reproduction, but it would probably be more correct to see in them -exceptional modes of adaptation enabling such individuals to occupy -niches in the world untenable to creatures of more conservative habit. -That the peculiar “strains” of animal life have turned into backwaters -which offer no opportunity or possibility of further advancement seems -clear enough, but they are nevertheless interesting and instructive. - -The parthenogenetic Crustacea and the Rotifers afford some good -evidence of this adaptability—of the way in which creatures manage to -cling to the skirts of life by reason of their power to survive the -extremest tests of endurance. And this success has largely been due -to some mysterious property of the germ-plasm enabling reproduction -to take place through the female line alone, or in some cases with -an occasional fillip from the intervention of males. Of the many -marvellous things that could be related of these creatures but few -instances can be cited here. - -The case of the Brine Shrimp (_Artemia salina_) will afford an -exceptionally good illustration because the facts can be tested by -anyone who will take the trouble to make a simple experiment for -himself. Those anxious to do this should dissolve eight ounces of -Tidman’s sea-salt in a glass jar containing five pints of water, -keeping the mixture well stirred till the salt is dissolved. It should -be allowed to stand and be carefully watched. In about three days, with -a pocket-lens, or even without, minute white specks will be seen moving -with a jerky motion up and down the water. These are larval Brine -Shrimps. Now they must be fed. Take a piece of lettuce-leaf or any -green stuff, and pound it up, or grind it up with a knife-blade on a -plate with a little water, till the whole is reduced to the consistency -of green paint; then empty this into the water. This must be done -daily, or at any rate frequently. Quickly these tiny specks will grow -into Brine Shrimps, translucent creatures nearly half an inch long, -swimming about back downwards with a marvellously rhythmical movement -of delicate feet. In all probability no males will be found, but, on -the other hand, both sexes in almost equal numbers may be present. The -males may readily be distinguished by their massive arms immediately -behind the head, for the purpose of embracing the females. - -Whence came these wonderful animals? The mystery is easily explained. -The salt is genuine sea-salt, formed in brine-pans, chiefly in the -Mediterranean. As the water evaporated the Shrimps it contained -gradually died; but the eggs in the females became encapsuled in -the salt-crystals to hatch out long months after. In one of my own -experiments I succeeded with salt that I had kept for more than a -year. Of course, every sample of salt experimented with will not -yield successful results, but failures are not expensive. Now in -this brine-pan there were myriads of other animals which were killed -outright: the Brine Shrimp is at least able to pass on descendants by -reason of the vitality of its eggs. Some near relations of the Brine -Shrimps live in fresh water and possess similar powers of resistance to -adverse conditions. The Fairy Shrimp (_Chirocephalus_) is one of these. -Not unlike its cousin the Brine Shrimp in appearance, it lives in -shallow pools, such as have muddy bottoms and are constantly liable to -dry up. Birds hunting by the margins of the pool where the retreating -water has left a fringe of mud bear away more or less of this on -their feet and transport it to similar pools, or even puddles. Such -transplanted samples may easily contain numbers of eggs of this tiny -creature. Only a year or two ago Fairy Shrimps were found in abundance -in rain pools at Eton, and some, indeed, were discovered swimming gaily -about in a rain-filled cart-rut! - -Another very singular Crustacean, known as _Apus_, bears a curious -superficial likeness to the King Crab (_Limulus_), having a large -back-shield and a long tail. This little creature, a giant compared -with his nearest relations, is an inhabitant of wayside ponds and -ditches. Thousands of females may be taken for years in succession -without the advent of a single male. Then, for some strange reason -which we cannot even guess at, males appear. Like its freshwater -cousin, the Fairy Shrimp, _Apus_ can withstand drought: its favourite -haunts may be transformed into sun-baked hollows, but with a heavy -fall of rain and a few hours’ soaking the eggs left by dead females -develop, and once more the pool and its inhabitants are established -again. Having regard to the extraordinary vitality of these small -creatures, it is curious that they should ever disappear from their -favoured haunts. But they do. Not many years ago _Apus_ could be -found in abundance in many parts of the South of England. It is now -extinct; its last resorts were the ponds at Hampstead: now one may -search in vain for them. “No British specimens,” remarks Dr. Caiman, -a great authority on the Crustacea, “had been recorded for over forty -years, and the species was believed to be extinct in this country, -when it was found in 1907 by Mr. F. Balfour Browne in a brackish marsh -near Southwick, in Kirkcudbrightshire.” These had probably developed -from eggs accidentally transported by some bird from the Continent. -The extinction of the race throughout the British Islands can only -be attributed to the too long absence of males, and the consequent -inability to restore vigour by the more normal method of reproduction -by sexual congress. - -Among the Rotifers the little Wheel-animalcules exhibit an even greater -vitality, for not only can their eggs withstand prolonged desiccation, -but in some the body of the animal survives even harsher treatment. If -specimens be enclosed within a chamber containing a little sand or moss -the contents may be dried over sulphuric acid, or heated up to 200° F., -or left to the neglected dust of years, and will yet revive if a little -fresh water be added to the sand. Males are rare, and when they do -occur are little more than animated receptacles for semen, for they are -incapable of feeding, the gullet and digestive tract being reduced to a -solid cord. A certain amount of nourishment, however, may be absorbed -through the delicate body wall. - -The degeneration of the males in these parthenogenetic species -irresistibly reminds one of the smile of the Cheshire cat; they grow -smaller and smaller, and their functions less and less, till finally -nothing is left. The “complemental males” discovered years ago by -Darwin in the Barnacles well illustrate this process. In dissecting -adult specimens of the stalked Barnacle (_Scalpellum_) he found, just -inside the valves, in a pocket of the mantle, a varying number of -“complemental males,” tiny organisms which Mr. Geoffrey Smith describes -as “little more than bags of spermatozoa,” and they apparently serve -to fertilize the ripe ova of the larger animal—one cannot say of the -female, for Scalpellum, like most of the Barnacles, is hermaphrodite. -But it is believed that these complemental males are really arrested -hermaphrodites. At any rate, if it so be noted that with some of the -Barnacles, as with some other Crustacea, the larvæ are males, but -when adult life is attained female glands appear and hermaphroditism -is established. Such hermaphrodites have the singular distinction of -being males which have acquired female attributes, true females being -unknown among them! - -In one of the parasitic Crustacea (_Chondracanthus_) infesting the -gills of Gurnard, Plaice, Skate and other fish, the adult female is -about half an inch long, and very unlike a Crustacean in appearance; -the male is an extremely minute maggot-like object—a few millimetres -in length—and lives permanently attached to the belly of his mate just -at the base of the egg masses. More remarkable still is the case of -another nearly related parasitic species—_Lernea_—which becomes sexually -mature in its childhood. The males perform their part and die; their -mates arrive at maturity and settle down to a comfortable life as -parasites on fish, reproducing without further mating. - -That Parthenogenesis and Hermaphroditism are but specialized forms of -reproduction, leading sooner or later to degeneration and extinction, -there can be no doubt. They are, so to speak, failures in the evolution -of sex, demonstrating in a very forcible fashion the impossibility -of progress—as we understand it—where the sexual functions are thus -combined. - -To the differentiation of sex, resulting in separate male and female -individuals, we must attribute the marvellous complexity of the pageant -of life which confronts us to-day. The story of the Courtship of -Animals is only one of an infinite number of incidents in this pageant, -and one which is by no means easy of interpretation. - -In these pages an attempt has been made to show that this -differentiation of sex has, throughout, been accompanied by, -and largely moulded by, common instincts and behaviour, and this -interpretation is only to be reached by a study of the phenomena in -their simplest form among the lower grades of animal life. Colour and -the various sexual differences in form have been allowed to dominate -this investigation of the problem of sex, and have diverted attention -from more profitable and fruitful channels. - -The lower we descend in the scale of animal life the less convincing -becomes the argument that the colour, ornament or armature of the -males is the result of sexual selection in the older, Darwinian sense. -The argument of Geddes and Thomson and others that the males are more -“katabolic,” the females more “anabolic,” seems no less unsatisfactory, -for in many cases the female is just as highly ornamented as the -male, and in others she is considerably large. Further, in their less -specialized species the sexes are almost or quite indistinguishable -externally, and are sombrely clad, just as at the opposite extreme we -find them equally ornamented and equally active. - -We shall be nearer the truth if we regard these secondary sexual -characters as expression points of germinal variations. Though we -seem hopelessly ignorant as to the inciting cause of the variations, -at least we seem to be able to lay a finger on the agents by which -they are effected. And these are the hormones of the primary and -secondary sexual glands, whose functions affect more than the merely -sexual side of the organism. They profoundly affect the coloration of -animals, giving rise on the one hand to purely ornamental “secondary -sexual characters,” and on the other to changes of coloration which -achieve the ends of protective resemblance colours, or of “warning -coloration,” as circumstances may demand. There is nothing more -remarkable in this than the control which the pituitary body exercises -over stature, either when in a pathological condition, or when the -controlling action of the other gland secretions is removed, as by -castration. - -Hitherto much has been made of trophic nerves, which control growth; -but it is probable we have overlooked the still more important -action of “trophic” glands, such as the thyroid. This apparently -controls growth in many directions. Adaptations to environment which -are effected by changes in bodily shape-as in the transformation of -land-dwelling mammals into Seals and Whales—are probably largely -controlled by these glands. Their activity is as great as their -manifestation is varied. - -Why their action should be more stimulating in the case of the male, -why he should lead the way in all the new acquirements of the species, -both in non-sexual as well as in sexual characters, is by no means -plain. But the fact remains that this is so. Remove any one of these -glands and the machinery of growth is thrown out of gear; it is not -merely the secondary sexual characters which are affected. - -But these glands are concerned no less intimately with the behaviour of -animals. This is most obvious in all that concerns sexual appetite as -the preceding chapters have already shown. Having regard to the immense -variety of animals concerned, this behaviour presents an underlying -uniformity of expression which must not be lost sight of: and the same -is no less true of what we may call the physical manifestations of -these glandular activities. - -THE END - - - - -INDEX - - - Alcock, Colonel, on courtship of - crabs, 255 - - Alder-flies, claspers of, 233 - - Amorousness, a factor in evolution, 24 - - — power of, 6, 9 - - — where absent, 6 - - Andrews, Dr. C. W., on display - of Frigate-bird, 111 - - Antelopes, battles of, 64 - - — horns of, 63 - - — scent glands of, 67 - - Antennæ, sense of smell in, 199 - - Antlers, branching of, 61 - - — eaten when shed, 58 - - — in female deer, 62 - - — shedding of, 53 - - — types of, 60 - - — use of, 55 - - Ants, nuptial flight of, 297 - - — dismal fate of queen, 298 - - Apes, brilliant colours of, 45 - - — — — use of, 46 - - — family relations of, 43 - - — polygamy in, 48 - - — power of voice in, 42 - - — related to man, 41 - - Aphides, appalling fertility of, 301 - - — parthenogenesis in, 301 - - Argonaut, remarkable egg-cradle of, 269 - - Armature in birds, 117 - - Argus Pheasant, display of, 96 - - — — ocelli of, 97 - - - Baboons, mane of, 44 - - Bailador, dances of, 121 - - Barrett Hamilton, Major, on Fur-seals, 85 - - Bee, Bumble, life of, 292 - - — drone, life of, 281, 289 - - — queen, as executioner, 279 - - — — execution of, 286 - - — — nuptial flight of, 279 - - — worker, evolution of, 290 - - Beetles, fighting between, 214 - - — stridulating organ of, 217 - - — vivid coloration in, 209 - - “Behaviour,” specific character of, 157 - - Birds, secondary sexual characters of, 94 - - Birds-of-Paradise, display of, 101 - - Bower-birds, coloration of, 158 - - — — origin of “bowers,” 160 - - — — singular behaviour of, 157 - - Brine-shrimp, vitality of, 304 - - Bug, extraordinary armature of, 216 - - Bustard, Australian, display of, 108 - - Bustard, Great, display of, 107 - - Butterflies, courtship of, 195 - - — excess of males in, 192 - - — experiments on, 205 - - — and female choice, 193 - - — females larger than males, 193 - - — fighting between, 194 - - — fragrance of, 200 - - — males mobbing females, 196 - - — methods of pairing, 204 - - — scent scales of, 199 - - Butterfly, Small-blue, method of folding wings of, 187 - - - Caiman, Dr. W. T., on A pus, 306 - - Campbell on courtship of spiders, 247 - - Cassowary, roar of, 112 - - Castration, effects of, 144 - - Chamæleons, armature of, 167 - - Cicada, music of, 222 - - Cockles, blindness of, 274 - - Coloration, cause of iridescent, 189 - - — forms of, 186 - - Conger-eel, huge size of females in, 178 - - Cooke, Mr. John, and the behaviour of sheep, 69 - - Courtship, meaning of, 21 - - Crabs, courtship of, 258 - - — seizing mates, 260 - - — stridulating organs of, 255 - - Crane, dances of, 120 - - Crocodile, courtship of, 164 - - Cunningham, J. T., and secondary sexual characters, 14 - - - Dancing in birds, 119 - - Darwin, his theory of Sexual Selection, 12 - - Darwin on coloration of beetles, 210, 211 - - — on coloration of mollusca, 275 - - — on “horns” of beetles, 212 - - — on mane of baboons, 44 - - — on sexual selection in butterflies, 194 - - Deer, antlers of, 53, 62 - - — courtship of, 53 - - — fatal encounters of, 55 - - “Diathetic” types, meaning of, 50 - - “Display,” function of, 183 - - — in birds, need of, 147, 149 - - Double-eyed fish, 176 - - Dragon-flies, antiquity of, 227 - - Dragonet, courtship of, 177 - - - Eland, horns of, 65 - - — strange habits of, 68 - - Elephant, courtship of, 71 - - — remarkable scent glands of, 69 - - — use of tusks in, 70 - - Elephant-seal, courtship of, 87 - - Emotions and human evolution, 32 - - — and sexual selection, 17 - - Emu, air-sacs of, 112 - - Eunuchs, peculiar features of, 145 - - Extinction, causes of, 17 - - - Fabre on courtship of scorpions, 252 - - Fashions among savages, 33 _et seq._ - - Fighting-fish, ferocity of, 182 - - Fish, disparity in size of sexes 178 - - Forbes, Dr. H. O., on deceptive coloration in a spider, 240 - - Frigate-bird, air-sacs in display of, III - - Frilled-lizard, courtship of, 165 - - Frogs, concerts of, 171 - - — courtship of, 169 - - — singing like a Greenfinch, 173 - - Fur-seals, courtship of, 81 - - — polygamy of, 81 - - — precocious sexual instincts in, 86 - - - Germ-plasm, nature of, 11 - - Giraffe, kick of, 73 - - — strong smell of, 68 - - Grasshopper, air-bladder of, 219 - - — stridulating organs of, 218, 221 - - Grebe, Great-crested, courtship of, 151 - - Groos, Professor, on emotions, 18 - - - Hippopotamus, bloody sweat of, 69 - - — teeth of, 72 - - Hooded-seal, air-sac of, 88 - - Hormones, nature of, 16 - - — part played by, 146, 147 - - Horns, evolution of, 51 - - Hottentots, remarkable peculiarities of, 31 - - Howard, H. E., on importance of “territory” 140 - - — on play in warblers, 121 - - — on sexual selection, 138 - - Huxley, Mr. Julian, on the behaviour of Mallard, 149 - - — on the Great-crested Grebe, 151 - - Hypertely, meaning of, 125, 127 - - - Image, Professor Selwyn, on Vapourer Moth, 201 - - Impotency, possible consequences of, 155 - - Ingram, Sir William, on display of King Bird-of-Paradise, 102 - - Insects and sexual selection, 192 - - - Katydid, song of, 222 - - Kinetogenesis, meaning of, 80 - - - Leptodora, olfactory sense of, 262 - - Life, lowest forms of, 3 _et seq._ - - Lion, mane of, 77 - - — polygamy in, 81 - - Lizards, courtship of, 165 _et seq._ - - — fighting among, 167 - - Locusts, ear of, 220 - - — remarkable ornaments of, 225 - - Lucas, Dr. F. A., on sea-lions, 86 - - - Males, degenerate, 307 - - — first to acquire new features, 9 - - — lead in new departures, 190 - - — where superfluous, 307 - - Mallard, remarkable behaviour of, 149 - - Man, evolution of, 22 _et seq._ - - “Mate-hunger,” part played by, 147 - - — power, importance of, 5 - - Mating, preferential, 155 - - May-flies, dance of death of, 229, 232 - - — nuptial flight of, 232 - - — remarkable eyes of male, 231 - - May-flies, remarkable use of stomach in, 230 - - Mayer, Mr., experiments on moths, 205 - - Moina, claspers of, 262 - - Moose, peculiar habits of, 58 - - Morgan, Professor Lloyd, on emotions, 18 - - — on factors in selection, 19, 126 - - Moth, aquatic females in, 204 - - — Eyed-hawk, display of, 201 - - — Ghost, scent-bottle of, 200 - - — Kentish Glory, sexual differences in, 203 - - — Oak-eggar, remarkable sense of smell in, 202 - - — Vapourer, remarkable sense of smell in, 201 - - Moths, bright colours of, 188 - - — diurnal, 189 - - — males, sense of smell of, 199 - - — scent organs of, 200 - - Mouth, brilliant colour in birds, 142 - - Muller, Fritz, on fragrant butterflies, 200 - - Music in birds, 123 _et seq._ - - - Narwhal, tusk of, 90 - - Newts, courtship of, 170 - - — remarkable pairing habits of, 170 - - - Ocelli in birds, nature of, 98 - - Octopus, courtship of, 267 - - — remarkable mating habits of, 268 - - Ornament, controlling factors of, 168 - - Orthogenesis, nature of, 17 - - Osborne, Professor, on factors in evolution, 17 - - Ostrich, roar of, 112 - - Oysters, helpless condition of, 274 - - — and social reformers, 275 - - - Painted Terrapin, remarkable courtship of, 168 - - Parthenogenesis in Crustacea, 304 - - — occurrence of, 276 - - — significance of, 308 - - — types of, 300 - - Peacock, display of, 95 - - Peckham, Mr., on courtship in spiders, 241 - - Pheasants, display of, 100 - - Pigment intensification, cause of, 150 - - Plovers, fatal conflicts among, 117 - - — dances of, 119 - - Plumage, evolution of, 143 - - Polyandry, human, 29 - - — in birds, 134 - - — interpretation of, 136 - - Polygamy, human, 27 - - — in birds, 115, 135 - - — in ungulates, 75 - - — interpretation of, 76, 115, 135, 136 - - Poulton, Professor E. B., on animal coloration, 186 - - — on courtship of spiders, 245 - - — on sexual selection in spiders, 245 - - Prairie-hen, air-sacs in display of, 109 - - Pugnacity in birds, 116 - - - Rays, remarkable teeth of, 176 - - Reproduction, forms of, 10 - - Rhinoceros, horns of, 75 - - Ruff, amorous instincts of, 115 - - — display of, 113 - - — variability of, 114 - - - Sabre-toothed tiger, huge canines of, 80 - - — jaw-flanges of, 80 - - Salmon, coloration of, 180 - - — courtship of, 179 - - Sandpiper, Pectoral, air-sacs in display of, 108 - - Scent glands, 67 - - Scorpions, courtship of, 242 - - Secondary Sexual Characters, meaning of, 13 - - Selection, forms of, 7 - - Selous, Mr. F. C., on habits of moose, 59 - - — on the mane of the lion, 78 - - Sex, beginnings of, 4 - - — birth of, 12 - - Sex-antagonism, 9 - - Sexual instincts, dominance of, 162 - - — grades of, 183 - - — importance of, 27 - - Sexual selection, definition of, 20 - - — and human evolution, 38 - - — in the human race, 32 - - — instance of working of, 147 - - Sexual Selection Theory, modification of, 154 - - Sharp, Dr. David, on stridulating organs, 219 - - Sheep, scent of, 69 - - Skull, human, malformation of, 35 - - Smith Woodward, Dr. A., on factors in evolution, 17 - - Smynthurus, remarkable courtship of, 26 - - Snail, hermaphrodite state of, 271 - - — “love-darts” of, 271 - - — significance of coloration of, 272 - - Somato-plasm, nature of, 11 - - South, Mr., on Oak-eggar moth, 202 - - Spiders, courtship of, 242 _et seq._ - - — dread sequel to nuptial rites, 246 - - — drumming of, 237 - - — nuptial rites in, 248 - - — remarkable coloration of, 239 - - — stridulating organs in, 237 - - Starling, Prof., on Hormones, 14, 16 - - State executions v. State pensions, 286 - - Stickle-back, paternal care in, 181 - - — remarkable nest of, 181 - - Stone-flies, degenerate wings in, 234 - - — nuptial rites performed in ice, 233 - - Stridulating organs of grass hoppers, 218 - - — nature of, 217 - - Sun-bittern, display of, 142 - - Syrinx in birds, 123 - - - Termite, amazing fertility of queen, 302 - - “Territory,” importance of, 154 - - Thomson, Prof. J. A., on evolution of sex, 12, 309 - - Townsend, Mr., on sea-elephants, 87 - - Tragopans, display of, 100 - - Trimen, Dr., on deceptive coloration in a spider, 240 - - Turkey, display of, 99 - - - Use, inherited effects of, 59 - - - Virgin births, belief in, 296 - - Voice in birds, 123 - - Wallace, Alfred Russell, and Sexual Selection, 13, 137, 196 - - Warblers, play in, 121 - - Water-fleas, olfactory organs of, 261 - - Water-fleas, mating apparatus of, 226 - - Weismann, Professor, on coloration of butterflies, 187 - - Whales, armature of, 89 - - — battering-rams of, 179-89 - - Wheel-animalcule, vitality of, 307 - - Windpipes, where coiled, 128 - - - Zebras, fighting among, 73 - - - Printed at The Chapel River Press, Kingston, Surrey. - - -ERRATUM - -For the first line of page 16, instead of “by certain glands of the -ductless glands,” read: “by certain of the ductless glands.” - - - - - -End of Project Gutenberg's The Courtship of Animals, by William Plane Pycraft - -*** END OF THIS PROJECT GUTENBERG EBOOK THE COURTSHIP OF ANIMALS *** - -***** This file should be named 60517-0.txt or 60517-0.zip ***** -This and all associated files of various formats will be found in: - http://www.gutenberg.org/6/0/5/1/60517/ - -Produced by Turgut Dincer (This file was produced from -images made available by The Internet Archive) - -Updated editions will replace the previous one--the old editions will -be renamed. - -Creating the works from print editions not protected by U.S. copyright -law means that no one owns a United States copyright in these works, -so the Foundation (and you!) can copy and distribute it in the United -States without permission and without paying copyright -royalties. 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