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<p>First, for the sterility of species when crossed and of their hybrid offspring. It is impossible to study the several memoirs and works of those two conscientious and admirable observers, Kolreuter and Gartner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility. Kolreuter makes the rule universal; but then he cuts the knot, for in ten cases in which he found two forms, considered by most authors as distinct species, quite fertile together, he unhesitatingly ranks them as varieties. Gartner, also, makes the rule equally universal; and he disputes the entire fertility of Kolreuters ten cases. But in these and in many other cases, Gartner is obliged carefully to count the seeds, in order to show that there is any degree of sterility. He always compares the maximum number of seeds produced by two species when first crossed, and the maximum produced by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature. But causes of serious error here intervene: a plant, to be hybridised, must be castrated, and, what is often more important, must be secluded in order to prevent pollen being brought to it by insects from other plants. Nearly all the plants experimented on by Gartner were potted, and were kept in a chamber in his house. That these processes are often injurious to the fertility of a plant cannot be doubted; for Gartner gives in his table about a score of cases of plants which he castrated, and artificially fertilised with their own pollen, and (excluding all cases such as the Leguminosae, in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. Moreover, as Gartner repeatedly crossed some forms, such as the common red and blue pimpernels (<i epub:type="z3998:taxonomy">Anagallis arvensis</i> and <i epub:type="z3998:taxonomy">coerulea</i>), which the best botanists rank as varieties, and found them absolutely sterile, we may doubt whether many species are really so sterile, when intercrossed, as he believed.</p>
<p>It is certain, on the one hand, that the sterility of various species when crossed is so different in degree and graduates away so insensibly, and, on the other hand, that the fertility of pure species is so easily affected by various circumstances, that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. I think no better evidence of this can be required than that the two most experienced observers who have ever lived, namely Kolreuter and Gartner, arrived at diametrically opposite conclusions in regard to some of the very same forms. It is also most instructive to compare—but I have not space here to enter on details—the evidence advanced by our best botanists on the question whether certain doubtful forms should be ranked as species or varieties, with the evidence from fertility adduced by different hybridisers, or by the same observer from experiments made during different years. It can thus be shown that neither sterility nor fertility affords any certain distinction between species and varieties. The evidence from this source graduates away, and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences.</p>
<p>In regard to the sterility of hybrids in successive generations; though Gartner was enabled to rear some hybrids, carefully guarding them from a cross with either pure parent, for six or seven, and in one case for ten generations, yet he asserts positively that their fertility never increases, but generally decreases greatly and suddenly. With respect to this decrease, it may first be noticed that when any deviation in structure or constitution is common to both parents, this is often transmitted in an augmented degree to the offspring; and both sexual elements in hybrid plants are already affected in some degree. But I believe that their fertility has been diminished in nearly all these cases by an independent cause, namely, by too close interbreeding. I have made so many experiments and collected so many facts, showing on the one hand that an occasional cross with a distinct individual or variety increases the vigour and fertility of the offspring, and on the other hand that very close interbreeding lessens their vigour and fertility, that I cannot doubt the correctness of this conclusion. Hybrids are seldom raised by experimentalists in great numbers; and as the parent-species, or other allied hybrids, generally grow in the same garden, the visits of insects must be carefully prevented during the flowering season: hence hybrids, if left to themselves, will generally be fertilised during each generation by pollen from the same flower; and this would probably be injurious to their fertility, already lessened by their hybrid origin. I am strengthened in this conviction by a remarkable statement repeatedly made by Gartner, namely, that if even the less fertile hybrids be artificially fertilised with hybrid pollen of the same kind, their fertility, notwithstanding the frequent ill effects from manipulation, sometimes decidedly increases, and goes on increasing. Now, in the process of artificial fertilisation, pollen is as often taken by chance (as I know from my own experience) from the anthers of another flower, as from the anthers of the flower itself which is to be fertilised; so that a cross between two flowers, though probably often on the same plant, would be thus effected. Moreover, whenever complicated experiments are in progress, so careful an observer as Gartner would have castrated his hybrids, and this would have insured in each generation a cross with pollen from a distinct flower, either from the same plant or from another plant of the same hybrid nature. And thus, the strange fact of an increase of fertility in the successive generations of <em>artificially fertilised</em> hybrids, in contrast with those spontaneously self-fertilised, may, as I believe, be accounted for by too close interbreeding having been avoided.</p>
<p>Now let us turn to the results arrived at by a third most experienced hybridiser, namely, the <abbr>Hon.</abbr> and <abbr>Rev.</abbr> <abbr class="name">W.</abbr> Herbert. He is as emphatic in his conclusion that some hybrids are perfectly fertile—as fertile as the pure parent-species—as are Kolreuter and Gartner that some degree of sterility between distinct species is a universal law of nature. He experimented on some of the very same species as did Gartner. The difference in their results may, I think, be in part accounted for by Herberts great horticultural skill, and by his having hothouses at his command. Of his many important statements I will here give only a single one as an example, namely, that “every ovule in a pod of <i epub:type="z3998:taxonomy">Crinum capense</i> fertilised by <i epub:type="z3998:taxonomy"><abbr>C.</abbr> revolutum</i> produced a plant, which I never saw to occur in a case of its natural fecundation.” So that here we have perfect, or even more than commonly perfect fertility, in a first cross between two distinct species.</p>
<p>Now let us turn to the results arrived at by a third most experienced hybridiser, namely, the <abbr>Hon.</abbr> and <abbr>Rev.</abbr> <abbr class="name">W.</abbr> Herbert. He is as emphatic in his conclusion that some hybrids are perfectly fertile—as fertile as the pure parent-species—as are Kolreuter and Gartner that some degree of sterility between distinct species is a universal law of nature. He experimented on some of the very same species as did Gartner. The difference in their results may, I think, be in part accounted for by Herberts great horticultural skill, and by his having hothouses at his command. Of his many important statements I will here give only a single one as an example, namely, that “every ovule in a pod of <i epub:type="z3998:taxonomy">Crinum capense</i> fertilised by <i epub:type="z3998:taxonomy"><abbr>C.</abbr> revolutum</i> produced a plant, which I never saw to occur in a case of its natural fecundation.” So that here we have perfect, or even more than commonly perfect fertility, in a first cross between two distinct species.</p>
<p>This case of the <i epub:type="z3998:taxonomy">Crinum</i> leads me to refer to a singular fact, namely, that individual plants of certain species of <i epub:type="z3998:taxonomy">Lobelia</i>, <i epub:type="z3998:taxonomy">Verbascum</i> and <i epub:type="z3998:taxonomy">Passiflora</i>, can easily be fertilised by the pollen from a distinct species, but not by pollen from the same plant, though this pollen can be proved to be perfectly sound by fertilising other plants or species. In the genus <i epub:type="z3998:taxonomy">Hippeastrum</i>, in Corydalis as shown by Professor Hildebrand, in various orchids as shown by <abbr>Mr.</abbr> Scott and Fritz Muller, all the individuals are in this peculiar condition. So that with some species, certain abnormal individuals, and in other species all the individuals, can actually be hybridised much more readily than they can be fertilised by pollen from the same individual plant! To give one instance, a bulb of <i epub:type="z3998:taxonomy">Hippeastrum aulicum</i> produced four flowers; three were fertilised by Herbert with their own pollen, and the fourth was subsequently fertilised by the pollen of a compound hybrid descended from three distinct species: the result was that “the ovaries of the three first flowers soon ceased to grow, and after a few days perished entirely, whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity, and bore good seed, which vegetated freely.” <abbr>Mr.</abbr> Herbert tried similar experiments during many years, and always with the same result. These cases serve to show on what slight and mysterious causes the lesser or greater fertility of a species sometimes depends.</p>
<p>The practical experiments of horticulturists, though not made with scientific precision, deserve some notice. It is notorious in how complicated a manner the species of <i epub:type="z3998:taxonomy">Pelargonium</i>, <i epub:type="z3998:taxonomy">Fuchsia</i>, <i epub:type="z3998:taxonomy">Calceolaria</i>, <i epub:type="z3998:taxonomy">Petunia</i>, <i epub:type="z3998:taxonomy">Rhododendron</i>, <abbr>etc.</abbr>, have been crossed, yet many of these hybrids seed freely. For instance, Herbert asserts that a hybrid from <i epub:type="z3998:taxonomy">Calceolaria integrifolia</i> and <i epub:type="z3998:taxonomy">plantaginea</i>, species most widely dissimilar in general habit, “reproduces itself as perfectly as if it had been a natural species from the mountains of Chile.” I have taken some pains to ascertain the degree of fertility of some of the complex crosses of rhododendrons, and I am assured that many of them are perfectly fertile. <abbr>Mr.</abbr> <abbr class="name">C.</abbr> Noble, for instance, informs me that he raises stocks for grafting from a hybrid between <i epub:type="z3998:taxonomy"><abbr>Rhod.</abbr> ponticum</i> and <i epub:type="z3998:taxonomy">catawbiense</i>, and that this hybrid “seeds as freely as it is possible to imagine.” Had hybrids, when fairly treated, always gone on decreasing in fertility in each successive generation, as Gartner believed to be the case, the fact would have been notorious to nurserymen. Horticulturists raise large beds of the same hybrid, and such alone are fairly treated, for by insect agency the several individuals are allowed to cross freely with each other, and the injurious influence of close interbreeding is thus prevented. Anyone may readily convince himself of the efficiency of insect agency by examining the flowers of the more sterile kinds of hybrid rhododendrons, which produce no pollen, for he will find on their stigmas plenty of pollen brought from other flowers.</p>
<p>In regard to animals, much fewer experiments have been carefully tried than with plants. If our systematic arrangements can be trusted, that is, if the genera of animals are as distinct from each other as are the genera of plants, then we may infer that animals more widely distinct in the scale of nature can be crossed more easily than in the case of plants; but the hybrids themselves are, I think, more sterile. It should, however, be borne in mind that, owing to few animals breeding freely under confinement, few experiments have been fairly tried: for instance, the canary-bird has been crossed with nine distinct species of finches, but, as not one of these breeds freely in confinement, we have no right to expect that the first crosses between them and the canary, or that their hybrids, should be perfectly fertile. Again, with respect to the fertility in successive generations of the more fertile hybrid animals, I hardly know of an instance in which two families of the same hybrid have been raised at the same time from different parents, so as to avoid the ill effects of close interbreeding. On the contrary, brothers and sisters have usually been crossed in each successive generation, in opposition to the constantly repeated admonition of every breeder. And in this case, it is not at all surprising that the inherent sterility in the hybrids should have gone on increasing.</p>

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<p>Lamarck was the first man whose conclusions on the subject excited much attention. This justly celebrated naturalist first published his views in <time datetime="1801">1801</time>; he much enlarged them in <time datetime="1809">1809</time> in his “Philosophie Zoologique,” and subsequently, <time datetime="1815">1815</time>, in the Introduction to his <i epub:type="se:name.publication.book"><abbr>Hist.</abbr> <abbr>Nat.</abbr> des Animaux sans Vertebres</i>. In these works he upholds the doctrine that all species, including man, are descended from other species. He first did the eminent service of arousing attention to the probability of all change in the organic, as well as in the inorganic world, being the result of law, and not of miraculous interposition. Lamarck seems to have been chiefly led to his conclusion on the gradual change of species, by the difficulty of distinguishing species and varieties, by the almost perfect gradation of forms in certain groups, and by the analogy of domestic productions. With respect to the means of modification, he attributed something to the direct action of the physical conditions of life, something to the crossing of already existing forms, and much to use and disuse, that is, to the effects of habit. To this latter agency he seems to attribute all the beautiful adaptations in nature; such as the long neck of the giraffe for browsing on the branches of trees. But he likewise believed in a law of progressive development, and as all the forms of life thus tend to progress, in order to account for the existence at the present day of simple productions, he maintains that such forms are now spontaneously generated.<a href="endnotes.xhtml#note-2" id="noteref-2" epub:type="noteref">2</a></p>
<p>Geoffroy Saint-Hilaire, as is stated in his <i epub:type="se:name.publication.book">Life</i>, written by his son, suspected, as early as <time datetime="1795">1795</time>, that what we call species are various degenerations of the same type. It was not until <time datetime="1828">1828</time> that he published his conviction that the same forms have not been perpetuated since the origin of all things. Geoffroy seems to have relied chiefly on the conditions of life, or the <i xml:lang="fr">monde ambiant</i> as the cause of change. He was cautious in drawing conclusions, and did not believe that existing species are now undergoing modification; and, as his son adds, “<i xml:lang="fr">Cest donc un problème à réserver entièrement à lavenir, supposé même que lavenir doive avoir prise sur lui.</i></p>
<p>In <time datetime="1813">1813</time> <abbr>Dr.</abbr> <abbr class="name">W. C.</abbr> Wells read before the Royal Society “An Account of a White Female, part of whose skin resembles that of a Negro;” but his paper was not published until his famous <i epub:type="se:name.publication.book">Two Essays upon Dew and Single Vision</i> appeared in <time datetime="1818">1818</time>. In this paper he distinctly recognises the principle of natural selection, and this is the first recognition which has been indicated; but he applies it only to the races of man, and to certain characters alone. After remarking that negroes and mulattoes enjoy an immunity from certain tropical diseases, he observes, firstly, that all animals tend to vary in some degree, and, secondly, that agriculturists improve their domesticated animals by selection; and then, he adds, but what is done in this latter case “by art, seems to be done with equal efficacy, though more slowly, by nature, in the formation of varieties of mankind, fitted for the country which they inhabit. Of the accidental varieties of man, which would occur among the first few and scattered inhabitants of the middle regions of Africa, some one would be better fitted than others to bear the diseases of the country. This race would consequently multiply, while the others would decrease; not only from their inability to sustain the attacks of disease, but from their incapacity of contending with their more vigorous neighbours. The colour of this vigorous race I take for granted, from what has been already said, would be dark. But the same disposition to form varieties still existing, a darker and a darker race would in the course of time occur: and as the darkest would be the best fitted for the climate, this would at length become the most prevalent, if not the only race, in the particular country in which it had originated.” He then extends these same views to the white inhabitants of colder climates. I am indebted to <abbr>Mr.</abbr> Rowley, of the United States, for having called my attention, through <abbr>Mr.</abbr> Brace, to the above passage of <abbr>Dr.</abbr> Wells work.</p>
<p>The <abbr>Hon.</abbr> and <abbr>Rev.</abbr> <abbr class="name">W.</abbr> Herbert, afterward Dean of Manchester, in the fourth volume of the <i epub:type="se:name.publication.journal">Horticultural Transactions</i>, <time datetime="1822">1822</time>, and in his work on the <i epub:type="se:name.publication.book">Amaryllidaceae</i> (<time datetime="1837">1837</time>, pages 19, 339), declares that “horticultural experiments have established, beyond the possibility of refutation, that botanical species are only a higher and more permanent class of varieties.” He extends the same view to animals. The dean believes that single species of each genus were created in an originally highly plastic condition, and that these have produced, chiefly by inter-crossing, but likewise by variation, all our existing species.</p>
<p>The <abbr>Hon.</abbr> and <abbr>Rev.</abbr> <abbr class="name">W.</abbr> Herbert, afterward Dean of Manchester, in the fourth volume of the <i epub:type="se:name.publication.journal">Horticultural Transactions</i>, <time datetime="1822">1822</time>, and in his work on the <i epub:type="se:name.publication.book">Amaryllidaceae</i> (<time datetime="1837">1837</time>, pages 19, 339), declares that “horticultural experiments have established, beyond the possibility of refutation, that botanical species are only a higher and more permanent class of varieties.” He extends the same view to animals. The dean believes that single species of each genus were created in an originally highly plastic condition, and that these have produced, chiefly by inter-crossing, but likewise by variation, all our existing species.</p>
<p>In <time datetime="1826">1826</time> Professor Grant, in the concluding paragraph in his well-known paper (<i epub:type="se:name.publication.journal">Edinburgh Philosophical Journal</i>, vol. <span epub:type="z3998:roman">XIV</span>, page 283) on the Spongilla, clearly declares his belief that species are descended from other species, and that they become improved in the course of modification. This same view was given in his Fifty-fifth Lecture, published in the <i epub:type="se:name.publication.journal">Lancet</i> in <time datetime="1834">1834</time>.</p>
<p>In <time datetime="1831">1831</time> <abbr>Mr.</abbr> Patrick Matthew published his work on <i epub:type="se:name.publication.book">Naval Timber and Arboriculture</i>, in which he gives precisely the same view on the origin of species as that (presently to be alluded to) propounded by <abbr>Mr.</abbr> Wallace and myself in the <i epub:type="se:name.publication.journal">Linnean Journal</i>, and as that enlarged in the present volume. Unfortunately the view was given by <abbr>Mr.</abbr> Matthew very briefly in scattered passages in an appendix to a work on a different subject, so that it remained unnoticed until <abbr>Mr.</abbr> Matthew himself drew attention to it in the <i epub:type="se:name.publication.magazine">Gardeners Chronicle</i>, on <time datetime="1860-04-07">April 7, 1860</time>. The differences of <abbr>Mr.</abbr> Matthews views from mine are not of much importance: he seems to consider that the world was nearly depopulated at successive periods, and then restocked; and he gives as an alternative, that new forms may be generated “without the presence of any mold or germ of former aggregates.” I am not sure that I understand some passages; but it seems that he attributes much influence to the direct action of the conditions of life. He clearly saw, however, the full force of the principle of natural selection.</p>
<p>The celebrated geologist and naturalist, Von Buch, in his excellent <i epub:type="se:name.publication.book">Description Physique des Isles Canaries</i> (<time datetime="1836">1836</time>, page 147), clearly expresses his belief that varieties slowly become changed into permanent species, which are no longer capable of intercrossing.</p>