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 				<span epub:type="z3998:roman">I</span>
 				<span epub:type="subtitle">Variation Under Domestication</span>
 			</h2>
-			<p epub:type="bridgehead">Causes of Variability⁠—Effects of Habit and the Use and Disuse of Parts⁠—Correlated Variation⁠—Inheritance⁠—Character of Domestic Varieties⁠—Difficulty of Distinguishing Between Varieties and Species⁠—Origin of Domestic Varieties from One or More Species⁠—Domestic Pigeons, Their Differences and Origin⁠—Principles of Selection, Anciently Followed, Their Effects⁠—Methodical and Unconscious Selection⁠—Unknown Origin of Our Domestic Productions⁠—Circumstances Favourable to Man's Power of Selection.</p>
+			<p epub:type="bridgehead">Causes of Variability⁠—Effects of Habit and the Use and Disuse of Parts⁠—Correlated Variation⁠—Inheritance⁠—Character of Domestic Varieties⁠—Difficulty of Distinguishing Between Varieties and Species⁠—Origin of Domestic Varieties from One or More Species⁠—Domestic Pigeons, Their Differences and Origin⁠—Principles of Selection, Anciently Followed, Their Effects⁠—Methodical and Unconscious Selection⁠—Unknown Origin of Our Domestic Productions⁠—Circumstances Favourable to Man’s Power of Selection.</p>
 			<section id="chapter-1-1" epub:type="z3998:subchapter">
 				<h3 epub:type="title">Causes of Variability</h3>
 				<p>When we compare the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us is, that they generally differ more from each other than do the individuals of any one species or variety in a state of nature. And if we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, we are driven to conclude that this great variability is due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent species had been exposed under nature. There is, also, some probability in the view propounded by Andrew Knight, that this variability may be partly connected with excess of food. It seems clear that organic beings must be exposed during several generations to new conditions to cause any great amount of variation; and that, when the organisation has once begun to vary, it generally continues varying for many generations. No case is on record of a variable organism ceasing to vary under cultivation. Our oldest cultivated plants, such as wheat, still yield new varieties: our oldest domesticated animals are still capable of rapid improvement or modification.</p>

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 					</tr>
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 						<td>Tertiary strata</td>
-						<td>2,240⁠</td>
+						<td>2,240</td>
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 			<h3 epub:type="title">On the Sudden Appearance of Whole Groups of Allied Species</h3>
 			<p>The abrupt manner in which whole groups of species suddenly appear in certain formations, has been urged by several palaeontologists⁠—for instance, by Agassiz, Pictet, and Sedgwick, as a fatal objection to the belief in the transmutation of species. If numerous species, belonging to the same genera or families, have really started into life at once, the fact would be fatal to the theory of evolution through natural selection. For the development by this means of a group of forms, all of which are descended from some one progenitor, must have been an extremely slow process; and the progenitors must have lived long before their modified descendants. But we continually overrate the perfection of the geological record, and falsely infer, because certain genera or families have not been found beneath a certain stage, that they did not exist before that stage. In all cases positive palaeontological evidence may be implicitly trusted; negative evidence is worthless, as experience has so often shown. We continually forget how large the world is, compared with the area over which our geological formations have been carefully examined; we forget that groups of species may elsewhere have long existed, and have slowly multiplied, before they invaded the ancient archipelagoes of Europe and the United States. We do not make due allowance for the enormous intervals of time which have elapsed between our consecutive formations, longer perhaps in many cases than the time required for the accumulation of each formation. These intervals will have given time for the multiplication of species from some one parent-form: and in the succeeding formation, such groups or species will appear as if suddenly created.</p>
 			<p>I may here recall a remark formerly made, namely, that it might require a long succession of ages to adapt an organism to some new and peculiar line of life, for instance, to fly through the air; and consequently that the transitional forms would often long remain confined to some one region; but that, when this adaptation had once been effected, and a few species had thus acquired a great advantage over other organisms, a comparatively short time would be necessary to produce many divergent forms, which would spread rapidly and widely throughout the world. Professor Pictet, in his excellent <i epub:type="se:name.publication.book">Review</i> of this work, in commenting on early transitional forms, and taking birds as an illustration, cannot see how the successive modifications of the anterior limbs of a supposed prototype could possibly have been of any advantage. But look at the penguins of the Southern Ocean; have not these birds their front limbs in this precise intermediate state of “neither true arms nor true wings?” Yet these birds hold their place victoriously in the battle for life; for they exist in infinite numbers and of many kinds. I do not suppose that we here see the real transitional grades through which the wings of birds have passed; but what special difficulty is there in believing that it might profit the modified descendants of the penguin, first to become enabled to flap along the surface of the sea like the logger-headed duck, and ultimately to rise from its surface and glide through the air?</p>
-			<p>I will now give a few examples to illustrate the foregoing remarks, and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. Even in so short an interval as that between the first and second editions of Pictet’s great work on <i epub:type="se:name.publication.book">Palaeontology</i>, published in <time datetime="1844">1844</time>⁠–⁠<time datetime="1846">46</time> and in <time datetime="1853">1853</time>⁠–<time datetime="1857">⁠57</time>, the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified; and a third edition would require still further changes. I may recall the well-known fact that in geological treatises, published not many years ago, mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series; and true mammals have been discovered in the new red sandstone at nearly the commencement of this great series. Cuvier used to urge that no monkey occurred in any tertiary stratum; but now extinct species have been discovered in India, South America and in Europe, as far back as the miocene stage. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States, who would have ventured to suppose that no less than at least thirty different bird-like animals, some of gigantic size, existed during that period? Not a fragment of bone has been discovered in these beds. Not long ago, palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period; but now we know, on the authority of Professor Owen, that a bird certainly lived during the deposition of the upper greensand; and still more recently, that strange bird, the Archeopteryx, with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws, has been discovered in the oolitic slates of Solenhofen. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world.</p>
+			<p>I will now give a few examples to illustrate the foregoing remarks, and to show how liable we are to error in supposing that whole groups of species have suddenly been produced. Even in so short an interval as that between the first and second editions of Pictet’s great work on <i epub:type="se:name.publication.book">Palaeontology</i>, published in <time datetime="1844">1844</time>⁠–⁠<time datetime="1846">46</time> and in <time datetime="1853">1853</time>⁠–⁠<time datetime="1857">57</time>, the conclusions on the first appearance and disappearance of several groups of animals have been considerably modified; and a third edition would require still further changes. I may recall the well-known fact that in geological treatises, published not many years ago, mammals were always spoken of as having abruptly come in at the commencement of the tertiary series. And now one of the richest known accumulations of fossil mammals belongs to the middle of the secondary series; and true mammals have been discovered in the new red sandstone at nearly the commencement of this great series. Cuvier used to urge that no monkey occurred in any tertiary stratum; but now extinct species have been discovered in India, South America and in Europe, as far back as the miocene stage. Had it not been for the rare accident of the preservation of footsteps in the new red sandstone of the United States, who would have ventured to suppose that no less than at least thirty different bird-like animals, some of gigantic size, existed during that period? Not a fragment of bone has been discovered in these beds. Not long ago, palaeontologists maintained that the whole class of birds came suddenly into existence during the eocene period; but now we know, on the authority of Professor Owen, that a bird certainly lived during the deposition of the upper greensand; and still more recently, that strange bird, the Archeopteryx, with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws, has been discovered in the oolitic slates of Solenhofen. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world.</p>
 			<p>I may give another instance, which, from having passed under my own eyes has much struck me. In a memoir on <i epub:type="se:name.publication.book">Fossil Sessile Cirripedes</i>, I stated that, from the large number of existing and extinct tertiary species; from the extraordinary abundance of the individuals of many species all over the world, from the Arctic regions to the equator, inhabiting various zones of depths, from the upper tidal limits to fifty fathoms; from the perfect manner in which specimens are preserved in the oldest tertiary beds; from the ease with which even a fragment of a valve can be recognised; from all these circumstances, I inferred that, had sessile cirripedes existed during the secondary periods, they would certainly have been preserved and discovered; and as not one species had then been discovered in beds of this age, I concluded that this great group had been suddenly developed at the commencement of the tertiary series. This was a sore trouble to me, adding, as I then thought, one more instance of the abrupt appearance of a great group of species. But my work had hardly been published, when a skilful palaeontologist, <abbr class="name">M.</abbr> Bosquet, sent me a drawing of a perfect specimen of an unmistakable sessile cirripede, which he had himself extracted from the chalk of Belgium. And, as if to make the case as striking as possible, this cirripede was a Chthamalus, a very common, large, and ubiquitous genus, of which not one species has as yet been found even in any tertiary stratum. Still more recently, a Pyrgoma, a member of a distinct subfamily of sessile cirripedes, has been discovered by <abbr>Mr.</abbr> Woodward in the upper chalk; so that we now have abundant evidence of the existence of this group of animals during the secondary period.</p>
 			<p>The case most frequently insisted on by palaeontologists of the apparently sudden appearance of a whole group of species, is that of the teleostean fishes, low down, according to Agassiz, in the Chalk period. This group includes the large majority of existing species. But certain Jurassic and Triassic forms are now commonly admitted to be teleostean; and even some palaeozoic forms have thus been classed by one high authority. If the teleosteans had really appeared suddenly in the northern hemisphere at the commencement of the chalk formation, the fact would have been highly remarkable; but it would not have formed an insuperable difficulty, unless it could likewise have been shown that at the same period the species were suddenly and simultaneously developed in other quarters of the world. It is almost superfluous to remark that hardly any fossil-fish are known from south of the equator; and by running through Pictet’s <i epub:type="se:name.publication.book">Palaeontology</i> it will be seen that very few species are known from several formations in Europe. Some few families of fish now have a confined range; the teleostean fishes might formerly have had a similarly confined range, and after having been largely developed in some one sea, have spread widely. Nor have we any right to suppose that the seas of the world have always been so freely open from south to north as they are at present. Even at this day, if the Malay Archipelago were converted into land, the tropical parts of the Indian Ocean would form a large and perfectly enclosed basin, in which any great group of marine animals might be multiplied; and here they would remain confined, until some of the species became adapted to a cooler climate, and were enabled to double the southern capes of Africa or Australia, and thus reach other and distant seas.</p>
 			<p>From these considerations, from our ignorance of the geology of other countries beyond the confines of Europe and the United States, and from the revolution in our palaeontological knowledge effected by the discoveries of the last dozen years, it seems to me to be about as rash to dogmatize on the succession of organic forms throughout the world, as it would be for a naturalist to land for five minutes on a barren point in Australia, and then to discuss the number and range of its productions.</p>

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 		<section id="introduction" epub:type="introduction">
 			<h2 epub:type="title">Introduction</h2>
 			<p>When on board <abbr class="initialism">HMS</abbr> <i epub:type="se:name.vessel.ship">Beagle</i>, as naturalist, I was much struck with certain facts in the distribution of the organic beings inhabiting South America, and in the geological relations of the present to the past inhabitants of that continent. These facts, as will be seen in the latter chapters of this volume, seemed to throw some light on the origin of species⁠—that mystery of mysteries, as it has been called by one of our greatest philosophers. On my return home, it occurred to me, in <time datetime="1837">1837</time>, that something might perhaps be made out on this question by patiently accumulating and reflecting on all sorts of facts which could possibly have any bearing on it. After five years’ work I allowed myself to speculate on the subject, and drew up some short notes; these I enlarged in <time datetime="1844">1844</time> into a sketch of the conclusions, which then seemed to me probable: from that period to the present day I have steadily pursued the same object. I hope that I may be excused for entering on these personal details, as I give them to show that I have not been hasty in coming to a decision.</p>
-			<p>My work is now (<time datetime="1859">1859</time>) nearly finished; but as it will take me many more years to complete it, and as my health is far from strong, I have been urged to publish this abstract. I have more especially been induced to do this, as <abbr>Mr.</abbr> Wallace, who is now studying the natural history of the Malay Archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. In <time datetime="1858">1858</time> he sent me a memoir on this subject, with a request that I would forward it to Sir Charles Lyell, who sent it to the Linnean Society, and it is published in the third volume of the journal of that society. Sir <abbr class="name">C.</abbr> Lyell and <abbr>Dr.</abbr> Hooker, who both knew of my work⁠—the latter having read my sketch of <time datetime="1844">1844⁠</time>—honoured me by thinking it advisable to publish, with <abbr>Mr.</abbr> Wallace’s excellent memoir, some brief extracts from my manuscripts.</p>
+			<p>My work is now (<time datetime="1859">1859</time>) nearly finished; but as it will take me many more years to complete it, and as my health is far from strong, I have been urged to publish this abstract. I have more especially been induced to do this, as <abbr>Mr.</abbr> Wallace, who is now studying the natural history of the Malay Archipelago, has arrived at almost exactly the same general conclusions that I have on the origin of species. In <time datetime="1858">1858</time> he sent me a memoir on this subject, with a request that I would forward it to Sir Charles Lyell, who sent it to the Linnean Society, and it is published in the third volume of the journal of that society. Sir <abbr class="name">C.</abbr> Lyell and <abbr>Dr.</abbr> Hooker, who both knew of my work⁠—the latter having read my sketch of <time datetime="1844">1844</time>⁠—honoured me by thinking it advisable to publish, with <abbr>Mr.</abbr> Wallace’s excellent memoir, some brief extracts from my manuscripts.</p>
 			<p>This abstract, which I now publish, must necessarily be imperfect. I cannot here give references and authorities for my several statements; and I must trust to the reader reposing some confidence in my accuracy. No doubt errors may have crept in, though I hope I have always been cautious in trusting to good authorities alone. I can here give only the general conclusions at which I have arrived, with a few facts in illustration, but which, I hope, in most cases will suffice. No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts, with references, on which my conclusions have been grounded; and I hope in a future work to do this. For I am well aware that scarcely a single point is discussed in this volume on which facts cannot be adduced, often apparently leading to conclusions directly opposite to those at which I have arrived. A fair result can be obtained only by fully stating and balancing the facts and arguments on both sides of each question; and this is here impossible.</p>
 			<p>I much regret that want of space prevents my having the satisfaction of acknowledging the generous assistance which I have received from very many naturalists, some of them personally unknown to me. I cannot, however, let this opportunity pass without expressing my deep obligations to <abbr>Dr.</abbr> Hooker, who, for the last fifteen years, has aided me in every possible way by his large stores of knowledge and his excellent judgment.</p>
 			<p>In considering the origin of species, it is quite conceivable that a naturalist, reflecting on the mutual affinities of organic beings, on their embryological relations, their geographical distribution, geological succession, and other such facts, might come to the conclusion that species had not been independently created, but had descended, like varieties, from other species. Nevertheless, such a conclusion, even if well founded, would be unsatisfactory, until it could be shown how the innumerable species inhabiting this world have been modified, so as to acquire that perfection of structure and coadaptation which justly excites our admiration. Naturalists continually refer to external conditions, such as climate, food, <abbr>etc.</abbr>, as the only possible cause of variation. In one limited sense, as we shall hereafter see, this may be true; but it is preposterous to attribute to mere external conditions, the structure, for instance, of the woodpecker, with its feet, tail, beak, and tongue, so admirably adapted to catch insects under the bark of trees. In the case of the mistletoe, which draws its nourishment from certain trees, which has seeds that must be transported by certain birds, and which has flowers with separate sexes absolutely requiring the agency of certain insects to bring pollen from one flower to the other, it is equally preposterous to account for the structure of this parasite, with its relations to several distinct organic beings, by the effects of external conditions, or of habit, or of the volition of the plant itself.</p>