Tagged initials in names and also many taxonomy items.

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 				<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>
 				<p>As far as I am able to judge, after long attending to the subject, the conditions of life appear to act in two ways⁠—directly on the whole organisation or on certain parts alone and indirectly by affecting the reproductive system. With respect to the direct action, we must bear in mind that in every case, as Professor Weismann has lately insisted, and as I have incidently shown in my work on <i epub:type="se:name.publication.book">Variation under Domestication</i>, there are two factors: namely, the nature of the organism and the nature of the conditions. The former seems to be much the more important; for nearly similar variations sometimes arise under, as far as we can judge, dissimilar conditions; and, on the other hand, dissimilar variations arise under conditions which appear to be nearly uniform. The effects on the offspring are either definite or indefinite. They may be considered as definite when all or nearly all the offspring of individuals exposed to certain conditions during several generations are modified in the same manner. It is extremely difficult to come to any conclusion in regard to the extent of the changes which have been thus definitely induced. There can, however, be little doubt about many slight changes, such as size from the amount of food, colour from the nature of the food, thickness of the skin and hair from climate, <abbr class="eoc">etc.</abbr> Each of the endless variations which we see in the plumage of our fowls must have had some efficient cause; and if the same cause were to act uniformly during a long series of generations on many individuals, all probably would be modified in the same manner. Such facts as the complex and extraordinary outgrowths which variably follow from the insertion of a minute drop of poison by a gall-producing insect, shows us what singular modifications might result in the case of plants from a chemical change in the nature of the sap.</p>
 				<p>Indefinite variability is a much more common result of changed conditions than definite variability, and has probably played a more important part in the formation of our domestic races. We see indefinite variability in the endless slight peculiarities which distinguish the individuals of the same species, and which cannot be accounted for by inheritance from either parent or from some more remote ancestor. Even strongly-marked differences occasionally appear in the young of the same litter, and in seedlings from the same seed-capsule. At long intervals of time, out of millions of individuals reared in the same country and fed on nearly the same food, deviations of structure so strongly pronounced as to deserve to be called monstrosities arise; but monstrosities cannot be separated by any distinct line from slighter variations. All such changes of structure, whether extremely slight or strongly marked, which appear among many individuals living together, may be considered as the indefinite effects of the conditions of life on each individual organism, in nearly the same manner as the chill effects different men in an indefinite manner, according to their state of body or constitution, causing coughs or colds, rheumatism, or inflammation of various organs.</p>
-				<p>With respect to what I have called the indirect action of changed conditions, namely, through the reproductive system of being affected, we may infer that variability is thus induced, partly from the fact of this system being extremely sensitive to any change in the conditions, and partly from the similarity, as Kolreuter and others have remarked, between the variability which follows from the crossing of distinct species, and that which may be observed with plants and animals when reared under new or unnatural conditions. Many facts clearly show how eminently susceptible the reproductive system is to very slight changes in the surrounding conditions. Nothing is more easy than to tame an animal, and few things more difficult than to get it to breed freely under confinement, even when the male and female unite. How many animals there are which will not breed, though kept in an almost free state in their native country! This is generally, but erroneously attributed to vitiated instincts. Many cultivated plants display the utmost vigour, and yet rarely or never seed! In some few cases it has been discovered that a very trifling change, such as a little more or less water at some particular period of growth, will determine whether or not a plant will produce seeds. I cannot here give the details which I have collected and elsewhere published on this curious subject; but to show how singular the laws are which determine the reproduction of animals under confinement, I may mention that carnivorous animals, even from the tropics, breed in this country pretty freely under confinement, with the exception of the plantigrades or bear family, which seldom produce young; whereas, carnivorous birds, with the rarest exception, hardly ever lay fertile eggs. Many exotic plants have pollen utterly worthless, in the same condition as in the most sterile hybrids. When, on the one hand, we see domesticated animals and plants, though often weak and sickly, breeding freely under confinement; and when, on the other hand, we see individuals, though taken young from a state of nature perfectly tamed, long-lived, and healthy (of which I could give numerous instances), yet having their reproductive system so seriously affected by unperceived causes as to fail to act, we need not be surprised at this system, when it does act under confinement, acting irregularly, and producing offspring somewhat unlike their parents. I may add that as some organisms breed freely under the most unnatural conditions⁠—for instance, rabbits and ferrets kept in hutches⁠—showing that their reproductive organs are not easily affected; so will some animals and plants withstand domestication or cultivation, and vary very slightly⁠—perhaps hardly more than in a state of nature.</p>
+				<p>With respect to what I have called the indirect action of changed conditions, namely, through the reproductive system being affected, we may infer that variability is thus induced, partly from the fact of this system being extremely sensitive to any change in the conditions, and partly from the similarity, as Kolreuter and others have remarked, between the variability which follows from the crossing of distinct species, and that which may be observed with plants and animals when reared under new or unnatural conditions. Many facts clearly show how eminently susceptible the reproductive system is to very slight changes in the surrounding conditions. Nothing is more easy than to tame an animal, and few things more difficult than to get it to breed freely under confinement, even when the male and female unite. How many animals there are which will not breed, though kept in an almost free state in their native country! This is generally, but erroneously attributed to vitiated instincts. Many cultivated plants display the utmost vigour, and yet rarely or never seed! In some few cases it has been discovered that a very trifling change, such as a little more or less water at some particular period of growth, will determine whether or not a plant will produce seeds. I cannot here give the details which I have collected and elsewhere published on this curious subject; but to show how singular the laws are which determine the reproduction of animals under confinement, I may mention that carnivorous animals, even from the tropics, breed in this country pretty freely under confinement, with the exception of the plantigrades or bear family, which seldom produce young; whereas, carnivorous birds, with the rarest exception, hardly ever lay fertile eggs. Many exotic plants have pollen utterly worthless, in the same condition as in the most sterile hybrids. When, on the one hand, we see domesticated animals and plants, though often weak and sickly, breeding freely under confinement; and when, on the other hand, we see individuals, though taken young from a state of nature perfectly tamed, long-lived, and healthy (of which I could give numerous instances), yet having their reproductive system so seriously affected by unperceived causes as to fail to act, we need not be surprised at this system, when it does act under confinement, acting irregularly, and producing offspring somewhat unlike their parents. I may add that as some organisms breed freely under the most unnatural conditions⁠—for instance, rabbits and ferrets kept in hutches⁠—showing that their reproductive organs are not easily affected; so will some animals and plants withstand domestication or cultivation, and vary very slightly⁠—perhaps hardly more than in a state of nature.</p>
 				<p>Some naturalists have maintained that all variations are connected with the act of sexual reproduction; but this is certainly an error; for I have given in another work a long list of “sporting plants;” as they are called by gardeners; that is, of plants which have suddenly produced a single bud with a new and sometimes widely different character from that of the other buds on the same plant. These bud variations, as they may be named, can be propagated by grafts, offsets, <abbr>etc.</abbr>, and sometimes by seed. They occur rarely under nature, but are far from rare under culture. As a single bud out of many thousands produced year after year on the same tree under uniform conditions, has been known suddenly to assume a new character; and as buds on distinct trees, growing under different conditions, have sometimes yielded nearly the same variety⁠—for instance, buds on peach-trees producing nectarines, and buds on common roses producing moss-roses⁠—we clearly see that the nature of the conditions is of subordinate importance in comparison with the nature of the organism in determining each particular form of variation; perhaps of not more importance than the nature of the spark, by which a mass of combustible matter is ignited, has in determining the nature of the flames.</p>
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 				<p>It has often been assumed that man has chosen for domestication animals and plants having an extraordinary inherent tendency to vary, and likewise to withstand diverse climates. I do not dispute that these capacities have added largely to the value of most of our domesticated productions; but how could a savage possibly know, when he first tamed an animal, whether it would vary in succeeding generations, and whether it would endure other climates? Has the little variability of the ass and goose, or the small power of endurance of warmth by the reindeer, or of cold by the common camel, prevented their domestication? I cannot doubt that if other animals and plants, equal in number to our domesticated productions, and belonging to equally diverse classes and countries, were taken from a state of nature, and could be made to breed for an equal number of generations under domestication, they would on an average vary as largely as the parent species of our existing domesticated productions have varied.</p>
 				<p>In the case of most of our anciently domesticated animals and plants, it is not possible to come to any definite conclusion, whether they are descended from one or several wild species. The argument mainly relied on by those who believe in the multiple origin of our domestic animals is, that we find in the most ancient times, on the monuments of Egypt, and in the lake-habitations of Switzerland, much diversity in the breeds; and that some of these ancient breeds closely resemble, or are even identical with, those still existing. But this only throws far backward the history of civilisation, and shows that animals were domesticated at a much earlier period than has hitherto been supposed. The lake-inhabitants of Switzerland cultivated several kinds of wheat and barley, the pea, the poppy for oil and flax; and they possessed several domesticated animals. They also carried on commerce with other nations. All this clearly shows, as Heer has remarked, that they had at this early age progressed considerably in civilisation; and this again implies a long continued previous period of less advanced civilisation, during which the domesticated animals, kept by different tribes in different districts, might have varied and given rise to distinct races. Since the discovery of flint tools in the superficial formations of many parts of the world, all geologists believe that barbarian men existed at an enormously remote period; and we know that at the present day there is hardly a tribe so barbarous as not to have domesticated at least the dog.</p>
 				<p>The origin of most of our domestic animals will probably forever remain vague. But I may here state that, looking to the domestic dogs of the whole world, I have, after a laborious collection of all known facts, come to the conclusion that several wild species of Canidae have been tamed, and that their blood, in some cases mingled together, flows in the veins of our domestic breeds. In regard to sheep and goats I can form no decided opinion. From facts communicated to me by <abbr>Mr.</abbr> Blyth, on the habits, voice, constitution and structure of the humped Indian cattle, it is almost certain that they are descended from a different aboriginal stock from our European cattle; and some competent judges believe that these latter have had two or three wild progenitors, whether or not these deserve to be called species. This conclusion, as well as that of the specific distinction between the humped and common cattle, may, indeed, be looked upon as established by the admirable researches of Professor Rutimeyer. With respect to horses, from reasons which I cannot here give, I am doubtfully inclined to believe, in opposition to several authors, that all the races belong to the same species. Having kept nearly all the English breeds of the fowl alive, having bred and crossed them, and examined their skeletons, it appears to me almost certain that all are the descendants of the wild Indian fowl, Gallus bankiva; and this is the conclusion of <abbr>Mr.</abbr> Blyth, and of others who have studied this bird in India. In regard to ducks and rabbits, some breeds of which differ much from each other, the evidence is clear that they are all descended from the common duck and wild rabbit.</p>
-				<p>The doctrine of the origin of our several domestic races from several aboriginal stocks, has been carried to an absurd extreme by some authors. They believe that every race which breeds true, let the distinctive characters be ever so slight, has had its wild prototype. At this rate there must have existed at least a score of species of wild cattle, as many sheep, and several goats, in Europe alone, and several even within Great Britain. One author believes that there formerly existed eleven wild species of sheep peculiar to Great Britain! When we bear in mind that Britain has now not one peculiar mammal, and France but few distinct from those of Germany, and so with Hungary, Spain, <abbr>etc.</abbr>, but that each of these kingdoms possesses several peculiar breeds of cattle, sheep, <abbr>etc.</abbr>, we must admit that many domestic breeds must have originated in Europe; for whence otherwise could they have been derived? So it is in India. Even in the case of the breeds of the domestic dog throughout the world, which I admit are descended from several wild species, it cannot be doubted that there has been an immense amount of inherited variation; for who will believe that animals closely resembling the Italian greyhound, the bloodhound, the bull-dog, pug-dog, or Blenheim spaniel, <abbr>etc.</abbr>⁠—so unlike all wild Canidae⁠—ever existed in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species; but by crossing we can only get forms in some degree intermediate between their parents; and if we account for our several domestic races by this process, we must admit the former existence of the most extreme forms, as the Italian greyhound, bloodhound, bull-dog, <abbr>etc.</abbr>, in the wild state. Moreover, the possibility of making distinct races by crossing has been greatly exaggerated. Many cases are on record showing that a race may be modified by occasional crosses if aided by the careful selection of the individuals which present the desired character; but to obtain a race intermediate between two quite distinct races would be very difficult. Sir J. Sebright expressly experimented with this object and failed. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) quite uniform in character, and everything seems simple enough; but when these mongrels are crossed one with another for several generations, hardly two of them are alike, and then the difficulty of the task becomes manifest.</p>
+				<p>The doctrine of the origin of our several domestic races from several aboriginal stocks, has been carried to an absurd extreme by some authors. They believe that every race which breeds true, let the distinctive characters be ever so slight, has had its wild prototype. At this rate there must have existed at least a score of species of wild cattle, as many sheep, and several goats, in Europe alone, and several even within Great Britain. One author believes that there formerly existed eleven wild species of sheep peculiar to Great Britain! When we bear in mind that Britain has now not one peculiar mammal, and France but few distinct from those of Germany, and so with Hungary, Spain, <abbr>etc.</abbr>, but that each of these kingdoms possesses several peculiar breeds of cattle, sheep, <abbr>etc.</abbr>, we must admit that many domestic breeds must have originated in Europe; for whence otherwise could they have been derived? So it is in India. Even in the case of the breeds of the domestic dog throughout the world, which I admit are descended from several wild species, it cannot be doubted that there has been an immense amount of inherited variation; for who will believe that animals closely resembling the Italian greyhound, the bloodhound, the bull-dog, pug-dog, or Blenheim spaniel, <abbr>etc.</abbr>⁠—so unlike all wild Canidae⁠—ever existed in a state of nature? It has often been loosely said that all our races of dogs have been produced by the crossing of a few aboriginal species; but by crossing we can only get forms in some degree intermediate between their parents; and if we account for our several domestic races by this process, we must admit the former existence of the most extreme forms, as the Italian greyhound, bloodhound, bull-dog, <abbr>etc.</abbr>, in the wild state. Moreover, the possibility of making distinct races by crossing has been greatly exaggerated. Many cases are on record showing that a race may be modified by occasional crosses if aided by the careful selection of the individuals which present the desired character; but to obtain a race intermediate between two quite distinct races would be very difficult. Sir <abbr class="name">J.</abbr> Sebright expressly experimented with this object and failed. The offspring from the first cross between two pure breeds is tolerably and sometimes (as I have found with pigeons) quite uniform in character, and everything seems simple enough; but when these mongrels are crossed one with another for several generations, hardly two of them are alike, and then the difficulty of the task becomes manifest.</p>
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 				<h3 epub:type="title">Breeds of the Domestic Pigeon, Their Differences and Origin</h3>
-				<p>Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the <abbr>Hon.</abbr> W. Elliot from India, and by the <abbr>Hon.</abbr> C. Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs. The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head, and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels. The runt is a bird of great size, with long, massive beak and large feet; some of the sub-breeds of runts have very long necks, others very long wings and tails, others singularly short tails. The barb is allied to the carrier, but, instead of a long beak, has a very short and broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter. The turbit has a short and conical beak, with a line of reversed feathers down the breast; and it has the habit of continually expanding, slightly, the upper part of the oesophagus. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood, and it has, proportionally to its size, elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds. The fantail has thirty or even forty tail-feathers, instead of twelve or fourteen, the normal number in all the members of the great pigeon family: these feathers are kept expanded and are carried so erect that in good birds the head and tail touch: the oil-gland is quite aborted. Several other less distinct breeds might be specified.</p>
+				<p>Believing that it is always best to study some special group, I have, after deliberation, taken up domestic pigeons. I have kept every breed which I could purchase or obtain, and have been most kindly favoured with skins from several quarters of the world, more especially by the <abbr>Hon.</abbr> <abbr class="name">W.</abbr> Elliot from India, and by the <abbr>Hon.</abbr> <abbr class="name">C.</abbr> Murray from Persia. Many treatises in different languages have been published on pigeons, and some of them are very important, as being of considerable antiquity. I have associated with several eminent fanciers, and have been permitted to join two of the London Pigeon Clubs. The diversity of the breeds is something astonishing. Compare the English carrier and the short-faced tumbler, and see the wonderful difference in their beaks, entailing corresponding differences in their skulls. The carrier, more especially the male bird, is also remarkable from the wonderful development of the carunculated skin about the head, and this is accompanied by greatly elongated eyelids, very large external orifices to the nostrils, and a wide gape of mouth. The short-faced tumbler has a beak in outline almost like that of a finch; and the common tumbler has the singular inherited habit of flying at a great height in a compact flock, and tumbling in the air head over heels. The runt is a bird of great size, with long, massive beak and large feet; some of the sub-breeds of runts have very long necks, others very long wings and tails, others singularly short tails. The barb is allied to the carrier, but, instead of a long beak, has a very short and broad one. The pouter has a much elongated body, wings, and legs; and its enormously developed crop, which it glories in inflating, may well excite astonishment and even laughter. The turbit has a short and conical beak, with a line of reversed feathers down the breast; and it has the habit of continually expanding, slightly, the upper part of the oesophagus. The Jacobin has the feathers so much reversed along the back of the neck that they form a hood, and it has, proportionally to its size, elongated wing and tail feathers. The trumpeter and laugher, as their names express, utter a very different coo from the other breeds. The fantail has thirty or even forty tail-feathers, instead of twelve or fourteen, the normal number in all the members of the great pigeon family: these feathers are kept expanded and are carried so erect that in good birds the head and tail touch: the oil-gland is quite aborted. Several other less distinct breeds might be specified.</p>
 				<p>In the skeletons of the several breeds, the development of the bones of the face, in length and breadth and curvature, differs enormously. The shape, as well as the breadth and length of the ramus of the lower jaw, varies in a highly remarkable manner. The caudal and sacral vertebrae vary in number; as does the number of the ribs, together with their relative breadth and the presence of processes. The size and shape of the apertures in the sternum are highly variable; so is the degree of divergence and relative size of the two arms of the furcula. The proportional width of the gape of mouth, the proportional length of the eyelids, of the orifice of the nostrils, of the tongue (not always in strict correlation with the length of beak), the size of the crop and of the upper part of the oesophagus; the development and abortion of the oil-gland; the number of the primary wing and caudal feathers; the relative length of the wing and tail to each other and to the body; the relative length of the leg and foot; the number of scutellae on the toes, the development of skin between the toes, are all points of structure which are variable. The period at which the perfect plumage is acquired varies, as does the state of the down with which the nestling birds are clothed when hatched. The shape and size of the eggs vary. The manner of flight, and in some breeds the voice and disposition, differ remarkably. Lastly, in certain breeds, the males and females have come to differ in a slight degree from each other.</p>
 				<p>Altogether at least a score of pigeons might be chosen, which, if shown to an ornithologist, and he were told that they were wild birds, would certainly be ranked by him as well-defined species. Moreover, I do not believe that any ornithologist would in this case place the English carrier, the short-faced tumbler, the runt, the barb, pouter, and fantail in the same genus; more especially as in each of these breeds several truly-inherited sub-breeds, or species, as he would call them, could be shown him.</p>
-				<p>Great as are the differences between the breeds of the pigeon, I am fully convinced that the common opinion of naturalists is correct, namely, that all are descended from the rock-pigeon (Columba livia), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects. As several of the reasons which have led me to this belief are in some degree applicable in other cases, I will here briefly give them. If the several breeds are not varieties, and have not proceeded from the rock-pigeon, they must have descended from at least seven or eight aboriginal stocks; for it is impossible to make the present domestic breeds by the crossing of any lesser number: how, for instance, could a pouter be produced by crossing two breeds unless one of the parent-stocks possessed the characteristic enormous crop? The supposed aboriginal stocks must all have been rock-pigeons, that is, they did not breed or willingly perch on trees. But besides C. livia, with its geographical sub-species, only two or three other species of rock-pigeons are known; and these have not any of the characters of the domestic breeds. Hence the supposed aboriginal stocks must either still exist in the countries where they were originally domesticated, and yet be unknown to ornithologists; and this, considering their size, habits and remarkable characters, seems improbable; or they must have become extinct in the wild state. But birds breeding on precipices, and good flyers, are unlikely to be exterminated; and the common rock-pigeon, which has the same habits with the domestic breeds, has not been exterminated even on several of the smaller British islets, or on the shores of the Mediterranean. Hence the supposed extermination of so many species having similar habits with the rock-pigeon seems a very rash assumption. Moreover, the several above-named domesticated breeds have been transported to all parts of the world, and, therefore, some of them must have been carried back again into their native country; but not one has become wild or feral, though the dovecot-pigeon, which is the rock-pigeon in a very slightly altered state, has become feral in several places. Again, all recent experience shows that it is difficult to get wild animals to breed freely under domestication; yet on the hypothesis of the multiple origin of our pigeons, it must be assumed that at least seven or eight species were so thoroughly domesticated in ancient times by half-civilized man, as to be quite prolific under confinement.</p>
+				<p>Great as are the differences between the breeds of the pigeon, I am fully convinced that the common opinion of naturalists is correct, namely, that all are descended from the rock-pigeon (<i epub:type="z3998:taxonomy">Columba livia</i>), including under this term several geographical races or sub-species, which differ from each other in the most trifling respects. As several of the reasons which have led me to this belief are in some degree applicable in other cases, I will here briefly give them. If the several breeds are not varieties, and have not proceeded from the rock-pigeon, they must have descended from at least seven or eight aboriginal stocks; for it is impossible to make the present domestic breeds by the crossing of any lesser number: how, for instance, could a pouter be produced by crossing two breeds unless one of the parent-stocks possessed the characteristic enormous crop? The supposed aboriginal stocks must all have been rock-pigeons, that is, they did not breed or willingly perch on trees. But besides <i epub:type="z3998:taxonomy">C. livia</i>, with its geographical sub-species, only two or three other species of rock-pigeons are known; and these have not any of the characters of the domestic breeds. Hence the supposed aboriginal stocks must either still exist in the countries where they were originally domesticated, and yet be unknown to ornithologists; and this, considering their size, habits and remarkable characters, seems improbable; or they must have become extinct in the wild state. But birds breeding on precipices, and good flyers, are unlikely to be exterminated; and the common rock-pigeon, which has the same habits with the domestic breeds, has not been exterminated even on several of the smaller British islets, or on the shores of the Mediterranean. Hence the supposed extermination of so many species having similar habits with the rock-pigeon seems a very rash assumption. Moreover, the several above-named domesticated breeds have been transported to all parts of the world, and, therefore, some of them must have been carried back again into their native country; but not one has become wild or feral, though the dovecot-pigeon, which is the rock-pigeon in a very slightly altered state, has become feral in several places. Again, all recent experience shows that it is difficult to get wild animals to breed freely under domestication; yet on the hypothesis of the multiple origin of our pigeons, it must be assumed that at least seven or eight species were so thoroughly domesticated in ancient times by half-civilized man, as to be quite prolific under confinement.</p>
 				<p>An argument of great weight, and applicable in several other cases, is, that the above-specified breeds, though agreeing generally with the wild rock-pigeon in constitution, habits, voice, colouring, and in most parts of their structure, yet are certainly highly abnormal in other parts; we may look in vain through the whole great family of Columbidae for a beak like that of the English carrier, or that of the short-faced tumbler, or barb; for reversed feathers like those of the Jacobin; for a crop like that of the pouter; for tail-feathers like those of the fantail. Hence it must be assumed, not only that half-civilized man succeeded in thoroughly domesticating several species, but that he intentionally or by chance picked out extraordinarily abnormal species; and further, that these very species have since all become extinct or unknown. So many strange contingencies are improbable in the highest degree.</p>
-				<p>Some facts in regard to the colouring of pigeons well deserve consideration. The rock-pigeon is of a slaty-blue, with white loins; but the Indian sub-species, C. intermedia of Strickland, has this part bluish. The tail has a terminal dark bar, with the outer feathers externally edged at the base with white. The wings have two black bars. Some semi-domestic breeds, and some truly wild breeds, have, besides the two black bars, the wings chequered with black. These several marks do not occur together in any other species of the whole family. Now, in every one of the domestic breeds, taking thoroughly well-bred birds, all the above marks, even to the white edging of the outer tail-feathers, sometimes concur perfectly developed. Moreover, when birds belonging to two or more distinct breeds are crossed, none of which are blue or have any of the above-specified marks, the mongrel offspring are very apt suddenly to acquire these characters. To give one instance out of several which I have observed: I crossed some white fantails, which breed very true, with some black barbs⁠—and it so happens that blue varieties of barbs are so rare that I never heard of an instance in England; and the mongrels were black, brown and mottled. I also crossed a barb with a spot, which is a white bird with a red tail and red spot on the forehead, and which notoriously breeds very true; the mongrels were dusky and mottled. I then crossed one of the mongrel barb-fantails with a mongrel barb-spot, and they produced a bird of as beautiful a blue colour, with the white loins, double black wing-bar, and barred and white-edged tail-feathers, as any wild rock-pigeon! We can understand these facts, on the well-known principle of reversion to ancestral characters, if all the domestic breeds are descended from the rock-pigeon. But if we deny this, we must make one of the two following highly improbable suppositions. Either, first, that all the several imagined aboriginal stocks were coloured and marked like the rock-pigeon, although no other existing species is thus coloured and marked, so that in each separate breed there might be a tendency to revert to the very same colours and markings. Or, secondly, that each breed, even the purest, has within a dozen, or at most within a score, of generations, been crossed by the rock-pigeon: I say within a dozen or twenty generations, for no instance is known of crossed descendants reverting to an ancestor of foreign blood, removed by a greater number of generations. In a breed which has been crossed only once the tendency to revert to any character derived from such a cross will naturally become less and less, as in each succeeding generation there will be less of the foreign blood; but when there has been no cross, and there is a tendency in the breed to revert to a character which was lost during some former generation, this tendency, for all that we can see to the contrary, may be transmitted undiminished for an indefinite number of generations. These two distinct cases of reversion are often confounded together by those who have written on inheritance.</p>
+				<p>Some facts in regard to the colouring of pigeons well deserve consideration. The rock-pigeon is of a slaty-blue, with white loins; but the Indian sub-species, <i epub:type="z3998:taxonomy">C. intermedia</i> of Strickland, has this part bluish. The tail has a terminal dark bar, with the outer feathers externally edged at the base with white. The wings have two black bars. Some semi-domestic breeds, and some truly wild breeds, have, besides the two black bars, the wings chequered with black. These several marks do not occur together in any other species of the whole family. Now, in every one of the domestic breeds, taking thoroughly well-bred birds, all the above marks, even to the white edging of the outer tail-feathers, sometimes concur perfectly developed. Moreover, when birds belonging to two or more distinct breeds are crossed, none of which are blue or have any of the above-specified marks, the mongrel offspring are very apt suddenly to acquire these characters. To give one instance out of several which I have observed: I crossed some white fantails, which breed very true, with some black barbs⁠—and it so happens that blue varieties of barbs are so rare that I never heard of an instance in England; and the mongrels were black, brown and mottled. I also crossed a barb with a spot, which is a white bird with a red tail and red spot on the forehead, and which notoriously breeds very true; the mongrels were dusky and mottled. I then crossed one of the mongrel barb-fantails with a mongrel barb-spot, and they produced a bird of as beautiful a blue colour, with the white loins, double black wing-bar, and barred and white-edged tail-feathers, as any wild rock-pigeon! We can understand these facts, on the well-known principle of reversion to ancestral characters, if all the domestic breeds are descended from the rock-pigeon. But if we deny this, we must make one of the two following highly improbable suppositions. Either, first, that all the several imagined aboriginal stocks were coloured and marked like the rock-pigeon, although no other existing species is thus coloured and marked, so that in each separate breed there might be a tendency to revert to the very same colours and markings. Or, secondly, that each breed, even the purest, has within a dozen, or at most within a score, of generations, been crossed by the rock-pigeon: I say within a dozen or twenty generations, for no instance is known of crossed descendants reverting to an ancestor of foreign blood, removed by a greater number of generations. In a breed which has been crossed only once the tendency to revert to any character derived from such a cross will naturally become less and less, as in each succeeding generation there will be less of the foreign blood; but when there has been no cross, and there is a tendency in the breed to revert to a character which was lost during some former generation, this tendency, for all that we can see to the contrary, may be transmitted undiminished for an indefinite number of generations. These two distinct cases of reversion are often confounded together by those who have written on inheritance.</p>
 				<p>Lastly, the hybrids or mongrels from between all the breeds of the pigeon are perfectly fertile, as I can state from my own observations, purposely made, on the most distinct breeds. Now, hardly any cases have been ascertained with certainty of hybrids from two quite distinct species of animals being perfectly fertile. Some authors believe that long-continued domestication eliminates this strong tendency to sterility in species. From the history of the dog, and of some other domestic animals, this conclusion is probably quite correct, if applied to species closely related to each other. But to extend it so far as to suppose that species, aboriginally as distinct as carriers, tumblers, pouters, and fantails now are, should yield offspring perfectly fertile, inter se, seems to me rash in the extreme.</p>
 				<p>From these several reasons, namely, the improbability of man having formerly made seven or eight supposed species of pigeons to breed freely under domestication⁠—these supposed species being quite unknown in a wild state, and their not having become anywhere feral⁠—these species presenting certain very abnormal characters, as compared with all other Columbidae, though so like the rock-pigeon in most other respects⁠—the occasional reappearance of the blue colour and various black marks in all the breeds, both when kept pure and when crossed⁠—and lastly, the mongrel offspring being perfectly fertile⁠—from these several reasons, taken together, we may safely conclude that all our domestic breeds are descended from the rock-pigeon or Columba livia with its geographical sub-species.</p>
-				<p>In favour of this view, I may add, firstly, that the wild C. livia has been found capable of domestication in Europe and in India; and that it agrees in habits and in a great number of points of structure with all the domestic breeds. Secondly, that although an English carrier or a short-faced tumbler differs immensely in certain characters from the rock-pigeon, yet that by comparing the several sub-breeds of these two races, more especially those brought from distant countries, we can make, between them and the rock-pigeon, an almost perfect series; so we can in some other cases, but not with all the breeds. Thirdly, those characters which are mainly distinctive of each breed are in each eminently variable, for instance, the wattle and length of beak of the carrier, the shortness of that of the tumbler, and the number of tail-feathers in the fantail; and the explanation of this fact will be obvious when we treat of selection. Fourthly, pigeons have been watched and tended with the utmost care, and loved by many people. They have been domesticated for thousands of years in several quarters of the world; the earliest known record of pigeons is in the fifth Aegyptian dynasty, about 3000 <abbr class="era">BC</abbr>., as was pointed out to me by Professor Lepsius; but <abbr>Mr.</abbr> Birch informs me that pigeons are given in a bill of fare in the previous dynasty. In the time of the Romans, as we hear from Pliny, immense prices were given for pigeons; “nay, they are come to this pass, that they can reckon up their pedigree and race.” Pigeons were much valued by Akber Khan in India, about the year 1600; never less than 20,000 pigeons were taken with the court. “The monarchs of Iran and Turan sent him some very rare birds;” and, continues the courtly historian, “His Majesty, by crossing the breeds, which method was never practised before, has improved them astonishingly.” About this same period the Dutch were as eager about pigeons as were the old Romans. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone, will likewise be obvious when we treat of selection. We shall then, also, see how it is that the several breeds so often have a somewhat monstrous character. It is also a most favourable circumstance for the production of distinct breeds, that male and female pigeons can be easily mated for life; and thus different breeds can be kept together in the same aviary.</p>
+				<p>In favour of this view, I may add, firstly, that the wild <i epub:type="z3998:taxonomy">C. livia</i> has been found capable of domestication in Europe and in India; and that it agrees in habits and in a great number of points of structure with all the domestic breeds. Secondly, that although an English carrier or a short-faced tumbler differs immensely in certain characters from the rock-pigeon, yet that by comparing the several sub-breeds of these two races, more especially those brought from distant countries, we can make, between them and the rock-pigeon, an almost perfect series; so we can in some other cases, but not with all the breeds. Thirdly, those characters which are mainly distinctive of each breed are in each eminently variable, for instance, the wattle and length of beak of the carrier, the shortness of that of the tumbler, and the number of tail-feathers in the fantail; and the explanation of this fact will be obvious when we treat of selection. Fourthly, pigeons have been watched and tended with the utmost care, and loved by many people. They have been domesticated for thousands of years in several quarters of the world; the earliest known record of pigeons is in the fifth Aegyptian dynasty, about 3000 <abbr class="era">BC</abbr>., as was pointed out to me by Professor Lepsius; but <abbr>Mr.</abbr> Birch informs me that pigeons are given in a bill of fare in the previous dynasty. In the time of the Romans, as we hear from Pliny, immense prices were given for pigeons; “nay, they are come to this pass, that they can reckon up their pedigree and race.” Pigeons were much valued by Akber Khan in India, about the year 1600; never less than 20,000 pigeons were taken with the court. “The monarchs of Iran and Turan sent him some very rare birds;” and, continues the courtly historian, “His Majesty, by crossing the breeds, which method was never practised before, has improved them astonishingly.” About this same period the Dutch were as eager about pigeons as were the old Romans. The paramount importance of these considerations in explaining the immense amount of variation which pigeons have undergone, will likewise be obvious when we treat of selection. We shall then, also, see how it is that the several breeds so often have a somewhat monstrous character. It is also a most favourable circumstance for the production of distinct breeds, that male and female pigeons can be easily mated for life; and thus different breeds can be kept together in the same aviary.</p>
 				<p>I have discussed the probable origin of domestic pigeons at some, yet quite insufficient, length; because when I first kept pigeons and watched the several kinds, well knowing how truly they breed, I felt fully as much difficulty in believing that since they had been domesticated they had all proceeded from a common parent, as any naturalist could in coming to a similar conclusion in regard to the many species of finches, or other groups of birds, in nature. One circumstance has struck me much; namely, that nearly all the breeders of the various domestic animals and the cultivators of plants, with whom I have conversed, or whose treatises I have read, are firmly convinced that the several breeds to which each has attended, are descended from so many aboriginally distinct species. Ask, as I have asked, a celebrated raiser of Hereford cattle, whether his cattle might not have descended from Long-horns, or both from a common parent-stock, and he will laugh you to scorn. I have never met a pigeon, or poultry, or duck, or rabbit fancier, who was not fully convinced that each main breed was descended from a distinct species. Van Mons, in his treatise on pears and apples, shows how utterly he disbelieves that the several sorts, for instance a Ribston-pippin or Codlin-apple, could ever have proceeded from the seeds of the same tree. Innumerable other examples could be given. The explanation, I think, is simple: from long-continued study they are strongly impressed with the differences between the several races; and though they well know that each race varies slightly, for they win their prizes by selecting such slight differences, yet they ignore all general arguments, and refuse to sum up in their minds slight differences accumulated during many successive generations. May not those naturalists who, knowing far less of the laws of inheritance than does the breeder, and knowing no more than he does of the intermediate links in the long lines of descent, yet admit that many of our domestic races are descended from the same parents⁠—may they not learn a lesson of caution, when they deride the idea of species in a state of nature being lineal descendants of other species?</p>
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 				<p epub:type="bridgehead">On the Absence of Intermediate Varieties at the Present Day⁠—On the Nature of Extinct Intermediate Varieties; On Their Number⁠—On the Lapse of Time, as Inferred from the Rate of Denudation and of Deposition Number⁠—On the Lapse of Time as Estimated by Years⁠—On the Poorness of Our Palaeontological Collections⁠—On the Intermittence of Geological Formations⁠—On the Denudation of Granitic Areas⁠—On the Absence of Intermediate Varieties in Any One Formation⁠—On the Sudden Appearance of Groups of Species⁠—On Their Sudden Appearance in the Lowest Known Fossiliferous Strata⁠—Antiquity of the Habitable Earth.</p>
 			</header>
 			<p>In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. Most of them have now been discussed. One, namely, the distinctness of specific forms and their not being blended together by innumerable transitional links, is a very obvious difficulty. I assigned reasons why such links do not commonly occur at the present day under the circumstances apparently most favourable for their presence, namely, on an extensive and continuous area with graduated physical conditions. I endeavoured to show, that the life of each species depends in a more important manner on the presence of other already defined organic forms, than on climate, and, therefore, that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. I endeavoured, also, to show that intermediate varieties, from existing in lesser numbers than the forms which they connect, will generally be beaten out and exterminated during the course of further modification and improvement. The main cause, however, of innumerable intermediate links not now occurring everywhere throughout nature depends, on the very process of natural selection, through which new varieties continually take the places of and supplant their parent-forms. But just in proportion as this process of extermination has acted on an enormous scale, so must the number of intermediate varieties, which have formerly existed, be truly enormous. Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this, perhaps, is the most obvious and serious objection which can be urged against my theory. The explanation lies, as I believe, in the extreme imperfection of the geological record.</p>
-			<p>In the first place, it should always be borne in mind what sort of intermediate forms must, on the theory, have formerly existed. I have found it difficult, when looking at any two species, to avoid picturing to myself forms <em>directly</em> intermediate between them. But this is a wholly false view; we should always look for forms intermediate between each species and a common but unknown progenitor; and the progenitor will generally have differed in some respects from all its modified descendants. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon; if we possessed all the intermediate varieties which have ever existed, we should have an extremely close series between both and the rock-pigeon; but we should have no varieties directly intermediate between the fantail and pouter; none, for instance, combining a tail somewhat expanded with a crop somewhat enlarged, the characteristic features of these two breeds. These two breeds, moreover, have become so much modified, that, if we had no historical or indirect evidence regarding their origin, it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon, C. livia, whether they had descended from this species or from some other allied species, such as C. oenas.</p>
+			<p>In the first place, it should always be borne in mind what sort of intermediate forms must, on the theory, have formerly existed. I have found it difficult, when looking at any two species, to avoid picturing to myself forms <em>directly</em> intermediate between them. But this is a wholly false view; we should always look for forms intermediate between each species and a common but unknown progenitor; and the progenitor will generally have differed in some respects from all its modified descendants. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon; if we possessed all the intermediate varieties which have ever existed, we should have an extremely close series between both and the rock-pigeon; but we should have no varieties directly intermediate between the fantail and pouter; none, for instance, combining a tail somewhat expanded with a crop somewhat enlarged, the characteristic features of these two breeds. These two breeds, moreover, have become so much modified, that, if we had no historical or indirect evidence regarding their origin, it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon, <i epub:type="z3998:taxonomy">C. livia</i>, whether they had descended from this species or from some other allied species, such as <i epub:type="z3998:taxonomy">C. oenas</i>.</p>
 			<p>So with natural species, if we look to forms very distinct, for instance to the horse and tapir, we have no reason to suppose that links directly intermediate between them ever existed, but between each and an unknown common parent. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse; but in some points of structure may have differed considerably from both, even perhaps more than they differ from each other. Hence, in all such cases, we should be unable to recognise the parent-form of any two or more species, even if we closely compared the structure of the parent with that of its modified descendants, unless at the same time we had a nearly perfect chain of the intermediate links.</p>
 			<p>It is just possible, by the theory, that one of two living forms might have descended from the other; for instance, a horse from a tapir; and in this case <em>direct</em> intermediate links will have existed between them. But such a case would imply that one form had remained for a very long period unaltered, whilst its descendants had undergone a vast amount of change; and the principle of competition between organism and organism, between child and parent, will render this a very rare event; for in all cases the new and improved forms of life tend to supplant the old and unimproved forms.</p>
 			<p>By the theory of natural selection all living species have been connected with the parent-species of each genus, by differences not greater than we see between the natural and domestic varieties of the same species at the present day; and these parent-species, now generally extinct, have in their turn been similarly connected with more ancient forms; and so on backwards, always converging to the common ancestor of each great class. So that the number of intermediate and transitional links, between all living and extinct species, must have been inconceivably great. But assuredly, if this theory be true, such have lived upon the earth.</p>
@@ -44,14 +44,14 @@
 		<section id="chapter-10-2" epub:type="z3998:subchapter">
 			<h3 epub:type="title">On the Poorness of Palaeontological Collections</h3>
 			<p>Now let us turn to our richest museums, and what a paltry display we behold! That our collections are imperfect is admitted by everyone. The remark of that admirable palaeontologist, Edward Forbes, should never be forgotten, namely, that very many fossil species are known and named from single and often broken specimens, or from a few specimens collected on some one spot. Only a small portion of the surface of the earth has been geologically explored, and no part with sufficient care, as the important discoveries made every year in Europe prove. No organism wholly soft can be preserved. Shells and bones decay and disappear when left on the bottom of the sea, where sediment is not accumulating. We probably take a quite erroneous view, when we assume that sediment is being deposited over nearly the whole bed of the sea, at a rate sufficiently quick to embed and preserve fossil remains. Throughout an enormously large proportion of the ocean, the bright blue tint of the water bespeaks its purity. The many cases on record of a formation conformably covered, after an immense interval of time, by another and later formation, without the underlying bed having suffered in the interval any wear and tear, seem explicable only on the view of the bottom of the sea not rarely lying for ages in an unaltered condition. The remains which do become embedded, if in sand or gravel, will, when the beds are upraised, generally be dissolved by the percolation of rain water charged with carbonic acid. Some of the many kinds of animals which live on the beach between high and low water mark seem to be rarely preserved. For instance, the several species of the Chthamalinae (a subfamily of sessile cirripedes) coat the rocks all over the world in infinite numbers: they are all strictly littoral, with the exception of a single Mediterranean species, which inhabits deep water and this has been found fossil in Sicily, whereas not one other species has hitherto been found in any tertiary formation: yet it is known that the genus Chthamalus existed during the Chalk period. Lastly, many great deposits, requiring a vast length of time for their accumulation, are entirely destitute of organic remains, without our being able to assign any reason: one of the most striking instances is that of the Flysch formation, which consists of shale and sandstone, several thousand, occasionally even six thousand feet in thickness, and extending for at least 300 miles from Vienna to Switzerland; and although this great mass has been most carefully searched, no fossils, except a few vegetable remains, have been found.</p>
-			<p>With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods, it is superfluous to state that our evidence is fragmentary in an extreme degree. For instance, until recently not a land-shell was known belonging to either of these vast periods, with the exception of one species discovered by Sir C. Lyell and <abbr>Dr.</abbr> Dawson in the carboniferous strata of North America; but now land-shells have been found in the lias. In regard to mammiferous remains, a glance at the historical table published in Lyell’s Manual, will bring home the truth, how accidental and rare is their preservation, far better than pages of detail. Nor is their rarity surprising, when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits; and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations.</p>
-			<p>But the imperfection in the geological record largely results from another and more important cause than any of the foregoing; namely, from the several formations being separated from each other by wide intervals of time. This doctrine has been emphatically admitted by many geologists and palaeontologists, who, like E. Forbes, entirely disbelieve in the change of species. When we see the formations tabulated in written works, or when we follow them in nature, it is difficult to avoid believing that they are closely consecutive. But we know, for instance, from Sir R. Murchison’s great work on Russia, what wide gaps there are in that country between the superimposed formations; so it is in North America, and in many other parts of the world. The most skilful geologist, if his attention had been confined exclusively to these large territories, would never have suspected that during the periods which were blank and barren in his own country, great piles of sediment, charged with new and peculiar forms of life, had elsewhere been accumulated. And if, in every separate territory, hardly any idea can be formed of the length of time which has elapsed between the consecutive formations, we may infer that this could nowhere be ascertained. The frequent and great changes in the mineralogical composition of consecutive formations, generally implying great changes in the geography of the surrounding lands, whence the sediment was derived, accord with the belief of vast intervals of time having elapsed between each formation.</p>
+			<p>With respect to the terrestrial productions which lived during the Secondary and Palaeozoic periods, it is superfluous to state that our evidence is fragmentary in an extreme degree. For instance, until recently not a land-shell was known belonging to either of these vast periods, with the exception of one species discovered by Sir <abbr class="name">C.</abbr> Lyell and <abbr>Dr.</abbr> Dawson in the carboniferous strata of North America; but now land-shells have been found in the lias. In regard to mammiferous remains, a glance at the historical table published in Lyell’s Manual, will bring home the truth, how accidental and rare is their preservation, far better than pages of detail. Nor is their rarity surprising, when we remember how large a proportion of the bones of tertiary mammals have been discovered either in caves or in lacustrine deposits; and that not a cave or true lacustrine bed is known belonging to the age of our secondary or palaeozoic formations.</p>
+			<p>But the imperfection in the geological record largely results from another and more important cause than any of the foregoing; namely, from the several formations being separated from each other by wide intervals of time. This doctrine has been emphatically admitted by many geologists and palaeontologists, who, like <abbr class="name">E.</abbr> Forbes, entirely disbelieve in the change of species. When we see the formations tabulated in written works, or when we follow them in nature, it is difficult to avoid believing that they are closely consecutive. But we know, for instance, from Sir <abbr class="name">R.</abbr> Murchison’s great work on Russia, what wide gaps there are in that country between the superimposed formations; so it is in North America, and in many other parts of the world. The most skilful geologist, if his attention had been confined exclusively to these large territories, would never have suspected that during the periods which were blank and barren in his own country, great piles of sediment, charged with new and peculiar forms of life, had elsewhere been accumulated. And if, in every separate territory, hardly any idea can be formed of the length of time which has elapsed between the consecutive formations, we may infer that this could nowhere be ascertained. The frequent and great changes in the mineralogical composition of consecutive formations, generally implying great changes in the geography of the surrounding lands, whence the sediment was derived, accord with the belief of vast intervals of time having elapsed between each formation.</p>
 			<p>We can, I think, see why the geological formations of each region are almost invariably intermittent; that is, have not followed each other in close sequence. Scarcely any fact struck me more when examining many hundred miles of the South American coasts, which have been upraised several hundred feet within the recent period, than the absence of any recent deposits sufficiently extensive to last for even a short geological period. Along the whole west coast, which is inhabited by a peculiar marine fauna, tertiary beds are so poorly developed that no record of several successive and peculiar marine faunas will probably be preserved to a distant age. A little reflection will explain why, along the rising coast of the western side of South America, no extensive formations with recent or tertiary remains can anywhere be found, though the supply of sediment must for ages have been great, from the enormous degradation of the coast rocks and from the muddy streams entering the sea. The explanation, no doubt, is that the littoral and sublittoral deposits are continually worn away, as soon as they are brought up by the slow and gradual rising of the land within the grinding action of the coast-waves.</p>
 			<p>We may, I think, conclude that sediment must be accumulated in extremely thick, solid, or extensive masses, in order to withstand the incessant action of the waves, when first upraised and during subsequent oscillations of level, as well as the subsequent subaerial degradation. Such thick and extensive accumulations of sediment may be formed in two ways; either in profound depths of the sea, in which case the bottom will not be inhabited by so many and such varied forms of life as the more shallow seas; and the mass when upraised will give an imperfect record of the organisms which existed in the neighbourhood during the period of its accumulation. Or sediment may be deposited to any thickness and extent over a shallow bottom, if it continue slowly to subside. In this latter case, as long as the rate of subsidence and supply of sediment nearly balance each other, the sea will remain shallow and favourable for many and varied forms, and thus a rich fossiliferous formation, thick enough, when upraised, to resist a large amount of denudation, may be formed.</p>
 			<p>I am convinced that nearly all our ancient formations, which are throughout the greater part of their thickness <em>rich in fossils</em>, have thus been formed during subsidence. Since publishing my views on this subject in 1845, I have watched the progress of geology, and have been surprised to note how author after author, in treating of this or that great formation, has come to the conclusion that it was accumulated during subsidence. I may add, that the only ancient tertiary formation on the west coast of South America, which has been bulky enough to resist such degradation as it has as yet suffered, but which will hardly last to a distant geological age, was deposited during a downward oscillation of level, and thus gained considerable thickness.</p>
 			<p>All geological facts tell us plainly that each area has undergone numerous slow oscillations of level, and apparently these oscillations have affected wide spaces. Consequently, formations rich in fossils and sufficiently thick and extensive to resist subsequent degradation, will have been formed over wide spaces during periods of subsidence, but only where the supply of sediment was sufficient to keep the sea shallow and to embed and preserve the remains before they had time to decay. On the other hand, as long as the bed of the sea remained stationary, <em>thick</em> deposits cannot have been accumulated in the shallow parts, which are the most favourable to life. Still less can this have happened during the alternate periods of elevation; or, to speak more accurately, the beds which were then accumulated will generally have been destroyed by being upraised and brought within the limits of the coast-action.</p>
 			<p>These remarks apply chiefly to littoral and sublittoral deposits. In the case of an extensive and shallow sea, such as that within a large part of the Malay Archipelago, where the depth varies from thirty or forty to sixty fathoms, a widely extended formation might be formed during a period of elevation, and yet not suffer excessively from denudation during its slow upheaval; but the thickness of the formation could not be great, for owing to the elevatory movement it would be less than the depth in which it was formed; nor would the deposit be much consolidated, nor be capped by overlying formations, so that it would run a good chance of being worn away by atmospheric degradation and by the action of the sea during subsequent oscillations of level. It has, however, been suggested by <abbr>Mr.</abbr> Hopkins, that if one part of the area, after rising and before being denuded, subsided, the deposit formed during the rising movement, though not thick, might afterwards become protected by fresh accumulations, and thus be preserved for a long period.</p>
-			<p><abbr>Mr.</abbr> Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. But all geologists, excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe, will admit that these latter rocks have been stripped of their covering to an enormous extent. For it is scarcely possible that such rocks could have been solidified and crystallised while uncovered; but if the metamorphic action occurred at profound depths of the ocean, the former protecting mantle of rock may not have been very thick. Admitting then that gneiss, mica-schist, granite, diorite, <abbr>etc.</abbr>, were once necessarily covered up, how can we account for the naked and extensive areas of such rocks in many parts of the world, except on the belief that they have subsequently been completely denuded of all overlying strata? That such extensive areas do exist cannot be doubted: the granitic region of Parime is described by Humboldt as being at least nineteen times as large as Switzerland. South of the Amazon, Boue colours an area composed of rocks of this nature as equal to that of Spain, France, Italy, part of Germany, and the British Islands, all conjoined. This region has not been carefully explored, but from the concurrent testimony of travellers, the granitic area is very large: thus Von Eschwege gives a detailed section of these rocks, stretching from Rio de Janeiro for 260 geographical miles inland in a straight line; and I travelled for 150 miles in another direction, and saw nothing but granitic rocks. Numerous specimens, collected along the whole coast, from near Rio de Janeiro to the mouth of the Plata, a distance of 1,100 geographical miles, were examined by me, and they all belonged to this class. Inland, along the whole northern bank of the Plata, I saw, besides modern tertiary beds, only one small patch of slightly metamorphosed rock, which alone could have formed a part of the original capping of the granitic series. Turning to a well-known region, namely, to the United States and Canada, as shown in Professor H.D. Rogers’ beautiful map, I have estimated the areas by cutting out and weighing the paper, and I find that the metamorphic (excluding the “semi-metamorphic”) and granite rocks exceed, in the proportion of 19 to 12.5, the whole of the newer Palaeozoic formations. In many regions the metamorphic and granite rocks would be found much more widely extended than they appear to be, if all the sedimentary beds were removed which rest unconformably on them, and which could not have formed part of the original mantle under which they were crystallised. Hence, it is probable that in some parts of the world whole formations have been completely denuded, with not a wreck left behind.</p>
+			<p><abbr>Mr.</abbr> Hopkins also expresses his belief that sedimentary beds of considerable horizontal extent have rarely been completely destroyed. But all geologists, excepting the few who believe that our present metamorphic schists and plutonic rocks once formed the primordial nucleus of the globe, will admit that these latter rocks have been stripped of their covering to an enormous extent. For it is scarcely possible that such rocks could have been solidified and crystallised while uncovered; but if the metamorphic action occurred at profound depths of the ocean, the former protecting mantle of rock may not have been very thick. Admitting then that gneiss, mica-schist, granite, diorite, <abbr>etc.</abbr>, were once necessarily covered up, how can we account for the naked and extensive areas of such rocks in many parts of the world, except on the belief that they have subsequently been completely denuded of all overlying strata? That such extensive areas do exist cannot be doubted: the granitic region of Parime is described by Humboldt as being at least nineteen times as large as Switzerland. South of the Amazon, Boue colours an area composed of rocks of this nature as equal to that of Spain, France, Italy, part of Germany, and the British Islands, all conjoined. This region has not been carefully explored, but from the concurrent testimony of travellers, the granitic area is very large: thus Von Eschwege gives a detailed section of these rocks, stretching from Rio de Janeiro for 260 geographical miles inland in a straight line; and I travelled for 150 miles in another direction, and saw nothing but granitic rocks. Numerous specimens, collected along the whole coast, from near Rio de Janeiro to the mouth of the Plata, a distance of 1,100 geographical miles, were examined by me, and they all belonged to this class. Inland, along the whole northern bank of the Plata, I saw, besides modern tertiary beds, only one small patch of slightly metamorphosed rock, which alone could have formed a part of the original capping of the granitic series. Turning to a well-known region, namely, to the United States and Canada, as shown in Professor <abbr class="name">H. D.</abbr> Rogers’ beautiful map, I have estimated the areas by cutting out and weighing the paper, and I find that the metamorphic (excluding the “semi-metamorphic”) and granite rocks exceed, in the proportion of 19 to 12.5, the whole of the newer Palaeozoic formations. In many regions the metamorphic and granite rocks would be found much more widely extended than they appear to be, if all the sedimentary beds were removed which rest unconformably on them, and which could not have formed part of the original mantle under which they were crystallised. Hence, it is probable that in some parts of the world whole formations have been completely denuded, with not a wreck left behind.</p>
 			<p>One remark is here worth a passing notice. During periods of elevation the area of the land and of the adjoining shoal parts of the sea will be increased and new stations will often be formed⁠—all circumstances favourable, as previously explained, for the formation of new varieties and species; but during such periods there will generally be a blank in the geological record. On the other hand, during subsidence, the inhabited area and number of inhabitants will decrease (excepting on the shores of a continent when first broken up into an archipelago), and consequently during subsidence, though there will be much extinction, few new varieties or species will be formed; and it is during these very periods of subsidence that the deposits which are richest in fossils have been accumulated.</p>
 		</section>
 		<section id="chapter-10-3" epub:type="z3998:subchapter">
@@ -60,13 +60,13 @@
 			<p>Although each formation may mark a very long lapse of years, each probably is short compared with the period requisite to change one species into another. I am aware that two palaeontologists, whose opinions are worthy of much deference, namely Bronn and Woodward, have concluded that the average duration of each formation is twice or thrice as long as the average duration of specific forms. But insuperable difficulties, as it seems to me, prevent us from coming to any just conclusion on this head. When we see a species first appearing in the middle of any formation, it would be rash in the extreme to infer that it had not elsewhere previously existed. So again, when we find a species disappearing before the last layers have been deposited, it would be equally rash to suppose that it then became extinct. We forget how small the area of Europe is compared with the rest of the world; nor have the several stages of the same formation throughout Europe been correlated with perfect accuracy.</p>
 			<p>We may safely infer that with marine animals of all kinds there has been a large amount of migration due to climatal and other changes; and when we see a species first appearing in any formation, the probability is that it only then first immigrated into that area. It is well known, for instance, that several species appear somewhat earlier in the palaeozoic beds of North America than in those of Europe; time having apparently been required for their migration from the American to the European seas. In examining the latest deposits, in various quarters of the world, it has everywhere been noted, that some few still existing species are common in the deposit, but have become extinct in the immediately surrounding sea; or, conversely, that some are now abundant in the neighbouring sea, but are rare or absent in this particular deposit. It is an excellent lesson to reflect on the ascertained amount of migration of the inhabitants of Europe during the glacial epoch, which forms only a part of one whole geological period; and likewise to reflect on the changes of level, on the extreme change of climate, and on the great lapse of time, all included within this same glacial period. Yet it may be doubted whether, in any quarter of the world, sedimentary deposits, <em>including fossil remains</em>, have gone on accumulating within the same area during the whole of this period. It is not, for instance, probable that sediment was deposited during the whole of the glacial period near the mouth of the Mississippi, within that limit of depth at which marine animals can best flourish: for we know that great geographical changes occurred in other parts of America during this space of time. When such beds as were deposited in shallow water near the mouth of the Mississippi during some part of the glacial period shall have been upraised, organic remains will probably first appear and disappear at different levels, owing to the migrations of species and to geographical changes. And in the distant future, a geologist, examining these beds, would be tempted to conclude that the average duration of life of the embedded fossils had been less than that of the glacial period, instead of having been really far greater, that is, extending from before the glacial epoch to the present day.</p>
 			<p>In order to get a perfect gradation between two forms in the upper and lower parts of the same formation, the deposit must have gone on continuously accumulating during a long period, sufficient for the slow process of modification; hence, the deposit must be a very thick one; and the species undergoing change must have lived in the same district throughout the whole time. But we have seen that a thick formation, fossiliferous throughout its entire thickness, can accumulate only during a period of subsidence; and to keep the depth approximately the same, which is necessary that the same marine species may live on the same space, the supply of sediment must nearly counterbalance the amount of subsidence. But this same movement of subsidence will tend to submerge the area whence the sediment is derived, and thus diminish the supply, whilst the downward movement continues. In fact, this nearly exact balancing between the supply of sediment and the amount of subsidence is probably a rare contingency; for it has been observed by more than one palaeontologist that very thick deposits are usually barren of organic remains, except near their upper or lower limits.</p>
-			<p>It would seem that each separate formation, like the whole pile of formations in any country, has generally been intermittent in its accumulation. When we see, as is so often the case, a formation composed of beds of widely different mineralogical composition, we may reasonably suspect that the process of deposition has been more or less interrupted. Nor will the closest inspection of a formation give us any idea of the length of time which its deposition may have consumed. Many instances could be given of beds, only a few feet in thickness, representing formations which are elsewhere thousands of feet in thickness, and which must have required an enormous period for their accumulation; yet no one ignorant of this fact would have even suspected the vast lapse of time represented by the thinner formation. Many cases could be given of the lower beds of a formation having been upraised, denuded, submerged, and then re-covered by the upper beds of the same formation⁠—facts, showing what wide, yet easily overlooked, intervals have occurred in its accumulation. In other cases we have the plainest evidence in great fossilised trees, still standing upright as they grew, of many long intervals of time and changes of level during the process of deposition, which would not have been suspected, had not the trees been preserved: thus Sir C. Lyell and <abbr>Dr.</abbr> Dawson found carboniferous beds 1,400 feet thick in Nova Scotia, with ancient root-bearing strata, one above the other, at no less than sixty-eight different levels. Hence, when the same species occurs at the bottom, middle, and top of a formation, the probability is that it has not lived on the same spot during the whole period of deposition, but has disappeared and reappeared, perhaps many times, during the same geological period. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation, a section would not include all the fine intermediate gradations which must on our theory have existed, but abrupt, though perhaps slight, changes of form.</p>
+			<p>It would seem that each separate formation, like the whole pile of formations in any country, has generally been intermittent in its accumulation. When we see, as is so often the case, a formation composed of beds of widely different mineralogical composition, we may reasonably suspect that the process of deposition has been more or less interrupted. Nor will the closest inspection of a formation give us any idea of the length of time which its deposition may have consumed. Many instances could be given of beds, only a few feet in thickness, representing formations which are elsewhere thousands of feet in thickness, and which must have required an enormous period for their accumulation; yet no one ignorant of this fact would have even suspected the vast lapse of time represented by the thinner formation. Many cases could be given of the lower beds of a formation having been upraised, denuded, submerged, and then re-covered by the upper beds of the same formation⁠—facts, showing what wide, yet easily overlooked, intervals have occurred in its accumulation. In other cases we have the plainest evidence in great fossilised trees, still standing upright as they grew, of many long intervals of time and changes of level during the process of deposition, which would not have been suspected, had not the trees been preserved: thus Sir <abbr class="name">C.</abbr> Lyell and <abbr>Dr.</abbr> Dawson found carboniferous beds 1,400 feet thick in Nova Scotia, with ancient root-bearing strata, one above the other, at no less than sixty-eight different levels. Hence, when the same species occurs at the bottom, middle, and top of a formation, the probability is that it has not lived on the same spot during the whole period of deposition, but has disappeared and reappeared, perhaps many times, during the same geological period. Consequently if it were to undergo a considerable amount of modification during the deposition of any one geological formation, a section would not include all the fine intermediate gradations which must on our theory have existed, but abrupt, though perhaps slight, changes of form.</p>
 			<p>It is all-important to remember that naturalists have no golden rule by which to distinguish species and varieties; they grant some little variability to each species, but when they meet with a somewhat greater amount of difference between any two forms, they rank both as species, unless they are enabled to connect them together by the closest intermediate gradations; and this, from the reasons just assigned, we can seldom hope to effect in any one geological section. Supposing B and C to be two species, and a third, A, to be found in an older and underlying bed; even if A were strictly intermediate between B and C, it would simply be ranked as a third and distinct species, unless at the same time it could be closely connected by intermediate varieties with either one or both forms. Nor should it be forgotten, as before explained, that A might be the actual progenitor of B and C, and yet would not necessarily be strictly intermediate between them in all respects. So that we might obtain the parent-species and its several modified descendants from the lower and upper beds of the same formation, and unless we obtained numerous transitional gradations, we should not recognise their blood-relationship, and should consequently rank them as distinct species.</p>
 			<p>It is notorious on what excessively slight differences many palaeontologists have founded their species; and they do this the more readily if the specimens come from different sub-stages of the same formation. Some experienced conchologists are now sinking many of the very fine species of D’Orbigny and others into the rank of varieties; and on this view we do find the kind of evidence of change which on the theory we ought to find. Look again at the later tertiary deposits, which include many shells believed by the majority of naturalists to be identical with existing species; but some excellent naturalists, as Agassiz and Pictet, maintain that all these tertiary species are specifically distinct, though the distinction is admitted to be very slight; so that here, unless we believe that these eminent naturalists have been misled by their imaginations, and that these late tertiary species really present no difference whatever from their living representatives, or unless we admit, in opposition to the judgment of most naturalists, that these tertiary species are all truly distinct from the recent, we have evidence of the frequent occurrence of slight modifications of the kind required. If we look to rather wider intervals of time, namely, to distinct but consecutive stages of the same great formation, we find that the embedded fossils, though universally ranked as specifically different, yet are far more closely related to each other than are the species found in more widely separated formations; so that here again we have undoubted evidence of change in the direction required by the theory; but to this latter subject I shall return in the following chapter.</p>
 			<p>With animals and plants that propagate rapidly and do not wander much, there is reason to suspect, as we have formerly seen, that their varieties are generally at first local; and that such local varieties do not spread widely and supplant their parent-form until they have been modified and perfected in some considerable degree. According to this view, the chance of discovering in a formation in any one country all the early stages of transition between any two forms, is small, for the successive changes are supposed to have been local or confined to some one spot. Most marine animals have a wide range; and we have seen that with plants it is those which have the widest range, that oftenest present varieties, so that, with shells and other marine animals, it is probable that those which had the widest range, far exceeding the limits of the known geological formations in Europe, have oftenest given rise, first to local varieties and ultimately to new species; and this again would greatly lessen the chance of our being able to trace the stages of transition in any one geological formation.</p>
 			<p>It is a more important consideration, leading to the same result, as lately insisted on by <abbr>Dr.</abbr> Falconer, namely, that the period during which each species underwent modification, though long as measured by years, was probably short in comparison with that during which it remained without undergoing any change.</p>
 			<p>It should not be forgotten, that at the present day, with perfect specimens for examination, two forms can seldom be connected by intermediate varieties, and thus proved to be the same species, until many specimens are collected from many places; and with fossil species this can rarely be done. We shall, perhaps, best perceive the improbability of our being enabled to connect species by numerous, fine, intermediate, fossil links, by asking ourselves whether, for instance, geologists at some future period will be able to prove that our different breeds of cattle, sheep, horses, and dogs are descended from a single stock or from several aboriginal stocks; or, again, whether certain seashells inhabiting the shores of North America, which are ranked by some conchologists as distinct species from their European representatives, and by other conchologists as only varieties, are really varieties, or are, as it is called, specifically distinct. This could be effected by the future geologist only by his discovering in a fossil state numerous intermediate gradations; and such success is improbable in the highest degree.</p>
-			<p>It has been asserted over and over again, by writers who believe in the immutability of species, that geology yields no linking forms. This assertion, as we shall see in the next chapter, is certainly erroneous. As Sir J. Lubbock has remarked, “Every species is a link between other allied forms.” If we take a genus having a score of species, recent and extinct, and destroy four-fifths of them, no one doubts that the remainder will stand much more distinct from each other. If the extreme forms in the genus happen to have been thus destroyed, the genus itself will stand more distinct from other allied genera. What geological research has not revealed, is the former existence of infinitely numerous gradations, as fine as existing varieties, connecting together nearly all existing and extinct species. But this ought not to be expected; yet this has been repeatedly advanced as a most serious objection against my views.</p>
+			<p>It has been asserted over and over again, by writers who believe in the immutability of species, that geology yields no linking forms. This assertion, as we shall see in the next chapter, is certainly erroneous. As Sir <abbr class="name">J.</abbr> Lubbock has remarked, “Every species is a link between other allied forms.” If we take a genus having a score of species, recent and extinct, and destroy four-fifths of them, no one doubts that the remainder will stand much more distinct from each other. If the extreme forms in the genus happen to have been thus destroyed, the genus itself will stand more distinct from other allied genera. What geological research has not revealed, is the former existence of infinitely numerous gradations, as fine as existing varieties, connecting together nearly all existing and extinct species. But this ought not to be expected; yet this has been repeatedly advanced as a most serious objection against my views.</p>
 			<p>It may be worth while to sum up the foregoing remarks on the causes of the imperfection of the geological record under an imaginary illustration. The Malay Archipelago is about the size of Europe from the North Cape to the Mediterranean, and from Britain to Russia; and therefore equals all the geological formations which have been examined with any accuracy, excepting those of the United States of America. I fully agree with <abbr>Mr.</abbr> Godwin-Austen, that the present condition of the Malay Archipelago, with its numerous large islands separated by wide and shallow seas, probably represents the former state of Europe, while most of our formations were accumulating. The Malay Archipelago is one of the richest regions in organic beings; yet if all the species were to be collected which have ever lived there, how imperfectly would they represent the natural history of the world!</p>
 			<p>But we have every reason to believe that the terrestrial productions of the archipelago would be preserved in an extremely imperfect manner in the formations which we suppose to be there accumulating. Not many of the strictly littoral animals, or of those which lived on naked submarine rocks, would be embedded; and those embedded in gravel or sand would not endure to a distant epoch. Wherever sediment did not accumulate on the bed of the sea, or where it did not accumulate at a sufficient rate to protect organic bodies from decay, no remains could be preserved.</p>
 			<p>Formations rich in fossils of many kinds, and of thickness sufficient to last to an age as distant in futurity as the secondary formations lie in the past, would generally be formed in the archipelago only during periods of subsidence. These periods of subsidence would be separated from each other by immense intervals of time, during which the area would be either stationary or rising; whilst rising, the fossiliferous formations on the steeper shores would be destroyed, almost as soon as accumulated, by the incessant coast-action, as we now see on the shores of South America. Even throughout the extensive and shallow seas within the archipelago, sedimentary beds could hardly be accumulated of great thickness during the periods of elevation, or become capped and protected by subsequent deposits, so as to have a good chance of enduring to a very distant future. During the periods of subsidence, there would probably be much extinction of life; during the periods of elevation, there would be much variation, but the geological record would then be less perfect.</p>
@@ -79,18 +79,18 @@
 			<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 Review 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 Palaeontology, published in 1844⁠–⁠46 and in 1853⁠–⁠57, 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 Fossil Sessile Cirripedes, 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, M. 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>I may give another instance, which, from having passed under my own eyes has much struck me. In a memoir on Fossil Sessile Cirripedes, 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 Palaeontology 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>
 		</section>
 		<section id="chapter-10-5" epub:type="z3998:subchapter">
 			<h3 epub:type="title">On the Sudden Appearance of Groups of Allied Species in the Lowest Known Fossiliferous Strata</h3>
 			<p>There is another and allied difficulty, which is much more serious. I allude to the manner in which species belonging to several of the main divisions of the animal kingdom suddenly appear in the lowest known fossiliferous rocks. Most of the arguments which have convinced me that all the existing species of the same group are descended from a single progenitor, apply with equal force to the earliest known species. For instance, it cannot be doubted that all the Cambrian and Silurian trilobites are descended from some one crustacean, which must have lived long before the Cambrian age, and which probably differed greatly from any known animal. Some of the most ancient animals, as the Nautilus, Lingula, <abbr>etc.</abbr>, do not differ much from living species; and it cannot on our theory be supposed, that these old species were the progenitors of all the species belonging to the same groups which have subsequently appeared, for they are not in any degree intermediate in character.</p>
-			<p>Consequently, if the theory be true, it is indisputable that before the lowest Cambrian stratum was deposited long periods elapsed, as long as, or probably far longer than, the whole interval from the Cambrian age to the present day; and that during these vast periods the world swarmed with living creatures. Here we encounter a formidable objection; for it seems doubtful whether the earth, in a fit state for the habitation of living creatures, has lasted long enough. Sir W. Thompson concludes that the consolidation of the crust can hardly have occurred less than twenty or more than four hundred million years ago, but probably not less than ninety-eight or more than two hundred million years. These very wide limits show how doubtful the data are; and other elements may have hereafter to be introduced into the problem. <abbr>Mr.</abbr> Croll estimates that about sixty million years have elapsed since the Cambrian period, but this, judging from the small amount of organic change since the commencement of the Glacial epoch, appears a very short time for the many and great mutations of life, which have certainly occurred since the Cambrian formation; and the previous one hundred and forty million years can hardly be considered as sufficient for the development of the varied forms of life which already existed during the Cambrian period. It is, however, probable, as Sir William Thompson insists, that the world at a very early period was subjected to more rapid and violent changes in its physical conditions than those now occurring; and such changes would have tended to induce changes at a corresponding rate in the organisms which then existed.</p>
-			<p>To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system, I can give no satisfactory answer. Several eminent geologists, with Sir R. Murchison at their head, were until recently convinced that we beheld in the organic remains of the lowest Silurian stratum the first dawn of life. Other highly competent judges, as Lyell and E. Forbes, have disputed this conclusion. We should not forget that only a small portion of the world is known with accuracy. Not very long ago M. Barrande added another and lower stage, abounding with new and peculiar species, beneath the then known Silurian system; and now, still lower down in the Lower Cambrian formation, <abbr>Mr.</abbr> Hicks has found South Wales beds rich in trilobites, and containing various molluscs and annelids. The presence of phosphatic nodules and bituminous matter, even in some of the lowest azotic rocks, probably indicates life at these periods; and the existence of the Eozoon in the Laurentian formation of Canada is generally admitted. There are three great series of strata beneath the Silurian system in Canada, in the lowest of which the Eozoon is found. Sir W. Logan states that their “united thickness may possibly far surpass that of all the succeeding rocks, from the base of the palaeozoic series to the present time. We are thus carried back to a period so remote, that the appearance of the so-called primordial fauna (of Barrande) may by some be considered as a comparatively modern event.” The Eozoon belongs to the most lowly organised of all classes of animals, but is highly organised for its class; it existed in countless numbers, and, as <abbr>Dr.</abbr> Dawson has remarked, certainly preyed on other minute organic beings, which must have lived in great numbers. Thus the words, which I wrote in 1859, about the existence of living beings long before the Cambrian period, and which are almost the same with those since used by Sir W. Logan, have proved true. Nevertheless, the difficulty of assigning any good reason for the absence of vast piles of strata rich in fossils beneath the Cambrian system is very great. It does not seem probable that the most ancient beds have been quite worn away by denudation, or that their fossils have been wholly obliterated by metamorphic action, for if this had been the case we should have found only small remnants of the formations next succeeding them in age, and these would always have existed in a partially metamorphosed condition. But the descriptions which we possess of the Silurian deposits over immense territories in Russia and in North America, do not support the view that the older a formation is the more invariably it has suffered extreme denudation and metamorphism.</p>
+			<p>Consequently, if the theory be true, it is indisputable that before the lowest Cambrian stratum was deposited long periods elapsed, as long as, or probably far longer than, the whole interval from the Cambrian age to the present day; and that during these vast periods the world swarmed with living creatures. Here we encounter a formidable objection; for it seems doubtful whether the earth, in a fit state for the habitation of living creatures, has lasted long enough. Sir <abbr class="name">W.</abbr> Thompson concludes that the consolidation of the crust can hardly have occurred less than twenty or more than four hundred million years ago, but probably not less than ninety-eight or more than two hundred million years. These very wide limits show how doubtful the data are; and other elements may have hereafter to be introduced into the problem. <abbr>Mr.</abbr> Croll estimates that about sixty million years have elapsed since the Cambrian period, but this, judging from the small amount of organic change since the commencement of the Glacial epoch, appears a very short time for the many and great mutations of life, which have certainly occurred since the Cambrian formation; and the previous one hundred and forty million years can hardly be considered as sufficient for the development of the varied forms of life which already existed during the Cambrian period. It is, however, probable, as Sir William Thompson insists, that the world at a very early period was subjected to more rapid and violent changes in its physical conditions than those now occurring; and such changes would have tended to induce changes at a corresponding rate in the organisms which then existed.</p>
+			<p>To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system, I can give no satisfactory answer. Several eminent geologists, with Sir <abbr class="name">R.</abbr> Murchison at their head, were until recently convinced that we beheld in the organic remains of the lowest Silurian stratum the first dawn of life. Other highly competent judges, as Lyell and <abbr class="name">E.</abbr> Forbes, have disputed this conclusion. We should not forget that only a small portion of the world is known with accuracy. Not very long ago <abbr class="name">M.</abbr> Barrande added another and lower stage, abounding with new and peculiar species, beneath the then known Silurian system; and now, still lower down in the Lower Cambrian formation, <abbr>Mr.</abbr> Hicks has found South Wales beds rich in trilobites, and containing various molluscs and annelids. The presence of phosphatic nodules and bituminous matter, even in some of the lowest azotic rocks, probably indicates life at these periods; and the existence of the Eozoon in the Laurentian formation of Canada is generally admitted. There are three great series of strata beneath the Silurian system in Canada, in the lowest of which the Eozoon is found. Sir <abbr class="name">W.</abbr> Logan states that their “united thickness may possibly far surpass that of all the succeeding rocks, from the base of the palaeozoic series to the present time. We are thus carried back to a period so remote, that the appearance of the so-called primordial fauna (of Barrande) may by some be considered as a comparatively modern event.” The Eozoon belongs to the most lowly organised of all classes of animals, but is highly organised for its class; it existed in countless numbers, and, as <abbr>Dr.</abbr> Dawson has remarked, certainly preyed on other minute organic beings, which must have lived in great numbers. Thus the words, which I wrote in 1859, about the existence of living beings long before the Cambrian period, and which are almost the same with those since used by Sir <abbr class="name">W.</abbr> Logan, have proved true. Nevertheless, the difficulty of assigning any good reason for the absence of vast piles of strata rich in fossils beneath the Cambrian system is very great. It does not seem probable that the most ancient beds have been quite worn away by denudation, or that their fossils have been wholly obliterated by metamorphic action, for if this had been the case we should have found only small remnants of the formations next succeeding them in age, and these would always have existed in a partially metamorphosed condition. But the descriptions which we possess of the Silurian deposits over immense territories in Russia and in North America, do not support the view that the older a formation is the more invariably it has suffered extreme denudation and metamorphism.</p>
 			<p>The case at present must remain inexplicable; and may be truly urged as a valid argument against the views here entertained. To show that it may hereafter receive some explanation, I will give the following hypothesis. From the nature of the organic remains which do not appear to have inhabited profound depths, in the several formations of Europe and of the United States; and from the amount of sediment, miles in thickness, of which the formations are composed, we may infer that from first to last large islands or tracts of land, whence the sediment was derived, occurred in the neighbourhood of the now existing continents of Europe and North America. This same view has since been maintained by Agassiz and others. But we do not know what was the state of things in the intervals between the several successive formations; whether Europe and the United States during these intervals existed as dry land, or as a submarine surface near land, on which sediment was not deposited, or as the bed of an open and unfathomable sea.</p>
 			<p>Looking to the existing oceans, which are thrice as extensive as the land, we see them studded with many islands; but hardly one truly oceanic island (with the exception of New Zealand, if this can be called a truly oceanic island) is as yet known to afford even a remnant of any palaeozoic or secondary formation. Hence, we may perhaps infer, that during the palaeozoic and secondary periods, neither continents nor continental islands existed where our oceans now extend; for had they existed, palaeozoic and secondary formations would in all probability have been accumulated from sediment derived from their wear and tear; and would have been at least partially upheaved by the oscillations of level, which must have intervened during these enormously long periods. If, then, we may infer anything from these facts, we may infer that, where our oceans now extend, oceans have extended from the remotest period of which we have any record; and on the other hand, that where continents now exist, large tracts of land have existed, subjected, no doubt, to great oscillations of level, since the Cambrian period. The coloured map appended to my volume on Coral Reefs, led me to conclude that the great oceans are still mainly areas of subsidence, the great archipelagoes still areas of oscillations of level, and the continents areas of elevation. But we have no reason to assume that things have thus remained from the beginning of the world. Our continents seem to have been formed by a preponderance, during many oscillations of level, of the force of elevation. But may not the areas of preponderant movement have changed in the lapse of ages? At a period long antecedent to the Cambrian epoch, continents may have existed where oceans are now spread out, and clear and open oceans may have existed where our continents now stand. Nor should we be justified in assuming that if, for instance, the bed of the Pacific Ocean were now converted into a continent we should there find sedimentary formations, in recognisable condition, older than the Cambrian strata, supposing such to have been formerly deposited; for it might well happen that strata which had subsided some miles nearer to the centre of the earth, and which had been pressed on by an enormous weight of superincumbent water, might have undergone far more metamorphic action than strata which have always remained nearer to the surface. The immense areas in some parts of the world, for instance in South America, of naked metamorphic rocks, which must have been heated under great pressure, have always seemed to me to require some special explanation; and we may perhaps believe that we see in these large areas the many formations long anterior to the Cambrian epoch in a completely metamorphosed and denuded condition.</p>
-			<p>The several difficulties here discussed, namely, that, though we find in our geological formations many links between the species which now exist and which formerly existed, we do not find infinitely numerous fine transitional forms closely joining them all together. The sudden manner in which several groups of species first appear in our European formations, the almost entire absence, as at present known, of formations rich in fossils beneath the Cambrian strata, are all undoubtedly of the most serious nature. We see this in the fact that the most eminent palaeontologists, namely, Cuvier, Agassiz, Barrande, Pictet, Falconer, E. Forbes, <abbr>etc.</abbr>, and all our greatest geologists, as Lyell, Murchison, Sedgwick, <abbr>etc.</abbr>, have unanimously, often vehemently, maintained the immutability of species. But Sir Charles Lyell now gives the support of his high authority to the opposite side, and most geologists and palaeontologists are much shaken in their former belief. Those who believe that the geological record is in any degree perfect, will undoubtedly at once reject my theory. For my part, following out Lyell’s metaphor, I look at the geological record as a history of the world imperfectly kept and written in a changing dialect. Of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved, and of each page, only here and there a few lines. Each word of the slowly-changing language, more or less different in the successive chapters, may represent the forms of life, which are entombed in our consecutive formations, and which falsely appear to have been abruptly introduced. On this view the difficulties above discussed are greatly diminished or even disappear.</p>
+			<p>The several difficulties here discussed, namely, that, though we find in our geological formations many links between the species which now exist and which formerly existed, we do not find infinitely numerous fine transitional forms closely joining them all together. The sudden manner in which several groups of species first appear in our European formations, the almost entire absence, as at present known, of formations rich in fossils beneath the Cambrian strata, are all undoubtedly of the most serious nature. We see this in the fact that the most eminent palaeontologists, namely, Cuvier, Agassiz, Barrande, Pictet, Falconer, <abbr class="name">E.</abbr> Forbes, <abbr>etc.</abbr>, and all our greatest geologists, as Lyell, Murchison, Sedgwick, <abbr>etc.</abbr>, have unanimously, often vehemently, maintained the immutability of species. But Sir Charles Lyell now gives the support of his high authority to the opposite side, and most geologists and palaeontologists are much shaken in their former belief. Those who believe that the geological record is in any degree perfect, will undoubtedly at once reject my theory. For my part, following out Lyell’s metaphor, I look at the geological record as a history of the world imperfectly kept and written in a changing dialect. Of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved, and of each page, only here and there a few lines. Each word of the slowly-changing language, more or less different in the successive chapters, may represent the forms of life, which are entombed in our consecutive formations, and which falsely appear to have been abruptly introduced. On this view the difficulties above discussed are greatly diminished or even disappear.</p>
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 			<p>Let us now see whether the several facts and laws relating to the geological succession of organic beings accord best with the common view of the immutability of species, or with that of their slow and gradual modification, through variation and natural selection.</p>
 			<p>New species have appeared very slowly, one after another, both on the land and in the waters. Lyell has shown that it is hardly possible to resist the evidence on this head in the case of the several tertiary stages; and every year tends to fill up the blanks between the stages, and to make the proportion between the lost and existing forms more gradual. In some of the most recent beds, though undoubtedly of high antiquity if measured by years, only one or two species are extinct, and only one or two are new, having appeared there for the first time, either locally, or, as far as we know, on the face of the earth. The secondary formations are more broken; but, as Bronn has remarked, neither the appearance nor disappearance of the many species embedded in each formation has been simultaneous.</p>
-			<p>Species belonging to different genera and classes have not changed at the same rate, or in the same degree. In the older tertiary beds a few living shells may still be found in the midst of a multitude of extinct forms. Falconer has given a striking instance of a similar fact, for an existing crocodile is associated with many lost mammals and reptiles in the sub-Himalayan deposits. The Silurian Lingula differs but little from the living species of this genus; whereas most of the other Silurian Molluscs and all the Crustaceans have changed greatly. The productions of the land seem to have changed at a quicker rate than those of the sea, of which a striking instance has been observed in Switzerland. There is some reason to believe that organisms high in the scale, change more quickly than those that are low: though there are exceptions to this rule. The amount of organic change, as Pictet has remarked, is not the same in each successive so-called formation. Yet if we compare any but the most closely related formations, all the species will be found to have undergone some change. When a species has once disappeared from the face of the earth, we have no reason to believe that the same identical form ever reappears. The strongest apparent exception to this latter rule is that of the so-called “colonies” of M. Barrande, which intrude for a period in the midst of an older formation, and then allow the preexisting fauna to reappear; but Lyell’s explanation, namely, that it is a case of temporary migration from a distinct geographical province, seems satisfactory.</p>
+			<p>Species belonging to different genera and classes have not changed at the same rate, or in the same degree. In the older tertiary beds a few living shells may still be found in the midst of a multitude of extinct forms. Falconer has given a striking instance of a similar fact, for an existing crocodile is associated with many lost mammals and reptiles in the sub-Himalayan deposits. The Silurian Lingula differs but little from the living species of this genus; whereas most of the other Silurian Molluscs and all the Crustaceans have changed greatly. The productions of the land seem to have changed at a quicker rate than those of the sea, of which a striking instance has been observed in Switzerland. There is some reason to believe that organisms high in the scale, change more quickly than those that are low: though there are exceptions to this rule. The amount of organic change, as Pictet has remarked, is not the same in each successive so-called formation. Yet if we compare any but the most closely related formations, all the species will be found to have undergone some change. When a species has once disappeared from the face of the earth, we have no reason to believe that the same identical form ever reappears. The strongest apparent exception to this latter rule is that of the so-called “colonies” of <abbr class="name">M.</abbr> Barrande, which intrude for a period in the midst of an older formation, and then allow the preexisting fauna to reappear; but Lyell’s explanation, namely, that it is a case of temporary migration from a distinct geographical province, seems satisfactory.</p>
 			<p>These several facts accord well with our theory, which includes no fixed law of development, causing all the inhabitants of an area to change abruptly, or simultaneously, or to an equal degree. The process of modification must be slow, and will generally affect only a few species at the same time; for the variability of each species is independent of that of all others. Whether such variations or individual differences as may arise will be accumulated through natural selection in a greater or less degree, thus causing a greater or less amount of permanent modification, will depend on many complex contingencies⁠—on the variations being of a beneficial nature, on the freedom of intercrossing, on the slowly changing physical conditions of the country, on the immigration of new colonists, and on the nature of the other inhabitants with which the varying species come into competition. Hence it is by no means surprising that one species should retain the same identical form much longer than others; or, if changing, should change in a less degree. We find similar relations between the existing inhabitants of distinct countries; for instance, the land-shells and coleopterous insects of Madeira have come to differ considerably from their nearest allies on the continent of Europe, whereas the marine shells and birds have remained unaltered. We can perhaps understand the apparently quicker rate of change in terrestrial and in more highly organised productions compared with marine and lower productions, by the more complex relations of the higher beings to their organic and inorganic conditions of life, as explained in a former chapter. When many of the inhabitants of any area have become modified and improved, we can understand, on the principle of competition, and from the all-important relations of organism to organism in the struggle for life, that any form which did not become in some degree modified and improved, would be liable to extermination. Hence, we see why all the species in the same region do at last, if we look to long enough intervals of time, become modified; for otherwise they would become extinct.</p>
 			<p>In members of the same class the average amount of change, during long and equal periods of time, may, perhaps, be nearly the same; but as the accumulation of enduring formations, rich in fossils, depends on great masses of sediment being deposited on subsiding areas, our formations have been almost necessarily accumulated at wide and irregularly intermittent intervals of time; consequently the amount of organic change exhibited by the fossils embedded in consecutive formations is not equal. Each formation, on this view, does not mark a new and complete act of creation, but only an occasional scene, taken almost at hazard, in an ever slowly changing drama.</p>
 			<p>We can clearly understand why a species when once lost should never reappear, even if the very same conditions of life, organic and inorganic, should recur. For though the offspring of one species might be adapted (and no doubt this has occurred in innumerable instances) to fill the place of another species in the economy of nature, and thus supplant it; yet the two forms⁠—the old and the new⁠—would not be identically the same; for both would almost certainly inherit different characters from their distinct progenitors; and organisms already differing would vary in a different manner. For instance, it is possible, if all our fantail-pigeons were destroyed, that fanciers might make a new breed hardly distinguishable from the present breed; but if the parent rock-pigeon were likewise destroyed, and under nature we have every reason to believe that parent forms are generally supplanted and exterminated by their improved offspring, it is incredible that a fantail, identical with the existing breed, could be raised from any other species of pigeon, or even from any other well established race of the domestic pigeon, for the successive variations would almost certainly be in some degree different, and the newly-formed variety would probably inherit from its progenitor some characteristic differences.</p>
-			<p>Groups of species, that is, genera and families, follow the same general rules in their appearance and disappearance as do single species, changing more or less quickly, and in a greater or lesser degree. A group, when it has once disappeared, never reappears; that is, its existence, as long as it lasts, is continuous. I am aware that there are some apparent exceptions to this rule, but the exceptions are surprisingly few, so few that E. Forbes, Pictet, and Woodward (though all strongly opposed to such views as I maintain) admit its truth; and the rule strictly accords with the theory. For all the species of the same group, however long it may have lasted, are the modified descendants one from the other, and all from a common progenitor. In the genus Lingula, for instance, the species which have successively appeared at all ages must have been connected by an unbroken series of generations, from the lowest Silurian stratum to the present day.</p>
+			<p>Groups of species, that is, genera and families, follow the same general rules in their appearance and disappearance as do single species, changing more or less quickly, and in a greater or lesser degree. A group, when it has once disappeared, never reappears; that is, its existence, as long as it lasts, is continuous. I am aware that there are some apparent exceptions to this rule, but the exceptions are surprisingly few, so few that <abbr class="name">E.</abbr> Forbes, Pictet, and Woodward (though all strongly opposed to such views as I maintain) admit its truth; and the rule strictly accords with the theory. For all the species of the same group, however long it may have lasted, are the modified descendants one from the other, and all from a common progenitor. In the genus Lingula, for instance, the species which have successively appeared at all ages must have been connected by an unbroken series of generations, from the lowest Silurian stratum to the present day.</p>
 			<p>We have seen in the last chapter that whole groups of species sometimes falsely appear to have been abruptly developed; and I have attempted to give an explanation of this fact, which if true would be fatal to my views. But such cases are certainly exceptional; the general rule being a gradual increase in number, until the group reaches its maximum, and then, sooner or later, a gradual decrease. If the number of the species included within a genus, or the number of the genera within a family, be represented by a vertical line of varying thickness, ascending through the successive geological formations, in which the species are found, the line will sometimes falsely appear to begin at its lower end, not in a sharp point, but abruptly; it then gradually thickens upwards, often keeping of equal thickness for a space, and ultimately thins out in the upper beds, marking the decrease and final extinction of the species. This gradual increase in number of the species of a group is strictly conformable with the theory; for the species of the same genus, and the genera of the same family, can increase only slowly and progressively; the process of modification and the production of a number of allied forms necessarily being a slow and gradual process, one species first giving rise to two or three varieties, these being slowly converted into species, which in their turn produce by equally slow steps other varieties and species, and so on, like the branching of a great tree from a single stem, till the group becomes large.</p>
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 			<p>Scarcely any palaeontological discovery is more striking than the fact that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant regions, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the organic remains in certain beds present an unmistakable resemblance to those of the Chalk. It is not that the same species are met with; for in some cases not one species is identically the same, but they belong to the same families, genera, and sections of genera, and sometimes are similarly characterised in such trifling points as mere superficial sculpture. Moreover, other forms, which are not found in the Chalk of Europe, but which occur in the formations either above or below, occur in the same order at these distant points of the world. In the several successive palaeozoic formations of Russia, Western Europe and North America, a similar parallelism in the forms of life has been observed by several authors; so it is, according to Lyell, with the European and North American tertiary deposits. Even if the few fossil species which are common to the Old and New Worlds were kept wholly out of view, the general parallelism in the successive forms of life, in the palaeozoic and tertiary stages, would still be manifest, and the several formations could be easily correlated.</p>
 			<p>These observations, however, relate to the marine inhabitants of the world: we have not sufficient data to judge whether the productions of the land and of fresh water at distant points change in the same parallel manner. We may doubt whether they have thus changed: if the Megatherium, Mylodon, Macrauchenia, and Toxodon had been brought to Europe from La Plata, without any information in regard to their geological position, no one would have suspected that they had coexisted with seashells all still living; but as these anomalous monsters coexisted with the Mastodon and Horse, it might at least have been inferred that they had lived during one of the later tertiary stages.</p>
 			<p>When the marine forms of life are spoken of as having changed simultaneously throughout the world, it must not be supposed that this expression relates to the same year, or even to the same century, or even that it has a very strict geological sense; for if all the marine animals now living in Europe, and all those that lived in Europe during the pleistocene period (a very remote period as measured by years, including the whole glacial epoch) were compared with those now existing in South America or in Australia, the most skilful naturalist would hardly be able to say whether the present or the pleistocene inhabitants of Europe resembled most closely those of the southern hemisphere. So, again, several highly competent observers maintain that the existing productions of the United States are more closely related to those which lived in Europe during certain late tertiary stages, than to the present inhabitants of Europe; and if this be so, it is evident that fossiliferous beds now deposited on the shores of North America would hereafter be liable to be classed with somewhat older European beds. Nevertheless, looking to a remotely future epoch, there can be little doubt that all the more modern <em>marine</em> formations, namely, the upper pliocene, the pleistocene and strictly modern beds of Europe, North and South America, and Australia, from containing fossil remains in some degree allied, and from not including those forms which are found only in the older underlying deposits, would be correctly ranked as simultaneous in a geological sense.</p>
-			<p>The fact of the forms of life changing simultaneously in the above large sense, at distant parts of the world, has greatly struck those admirable observers, <abbr>MM.</abbr> de Verneuil and d’Archiac. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe, they add, “If struck by this strange sequence, we turn our attention to North America, and there discover a series of analogous phenomena, it will appear certain that all these modifications of species, their extinction, and the introduction of new ones, cannot be owing to mere changes in marine currents or other causes more or less local and temporary, but depend on general laws which govern the whole animal kingdom.” M. Barrande has made forcible remarks to precisely the same effect. It is, indeed, quite futile to look to changes of currents, climate, or other physical conditions, as the cause of these great mutations in the forms of life throughout the world, under the most different climates. We must, as Barrande has remarked, look to some special law. We shall see this more clearly when we treat of the present distribution of organic beings, and find how slight is the relation between the physical conditions of various countries and the nature of their inhabitants.</p>
+			<p>The fact of the forms of life changing simultaneously in the above large sense, at distant parts of the world, has greatly struck those admirable observers, <abbr>MM.</abbr> de Verneuil and d’Archiac. After referring to the parallelism of the palaeozoic forms of life in various parts of Europe, they add, “If struck by this strange sequence, we turn our attention to North America, and there discover a series of analogous phenomena, it will appear certain that all these modifications of species, their extinction, and the introduction of new ones, cannot be owing to mere changes in marine currents or other causes more or less local and temporary, but depend on general laws which govern the whole animal kingdom.” <abbr class="name">M.</abbr> Barrande has made forcible remarks to precisely the same effect. It is, indeed, quite futile to look to changes of currents, climate, or other physical conditions, as the cause of these great mutations in the forms of life throughout the world, under the most different climates. We must, as Barrande has remarked, look to some special law. We shall see this more clearly when we treat of the present distribution of organic beings, and find how slight is the relation between the physical conditions of various countries and the nature of their inhabitants.</p>
 			<p>This great fact of the parallel succession of the forms of life throughout the world, is explicable on the theory of natural selection. New species are formed by having some advantage over older forms; and the forms, which are already dominant, or have some advantage over the other forms in their own country, give birth to the greatest number of new varieties or incipient species. We have distinct evidence on this head, in the plants which are dominant, that is, which are commonest and most widely diffused, producing the greatest number of new varieties. It is also natural that the dominant, varying and far-spreading species, which have already invaded, to a certain extent, the territories of other species, should be those which would have the best chance of spreading still further, and of giving rise in new countries to other new varieties and species. The process of diffusion would often be very slow, depending on climatal and geographical changes, on strange accidents, and on the gradual acclimatization of new species to the various climates through which they might have to pass, but in the course of time the dominant forms would generally succeed in spreading and would ultimately prevail. The diffusion would, it is probable, be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. We might therefore expect to find, as we do find, a less strict degree of parallelism in the succession of the productions of the land than with those of the sea.</p>
 			<p>Thus, as it seems to me, the parallel, and, taken in a large sense, simultaneous, succession of the same forms of life throughout the world, accords well with the principle of new species having been formed by dominant species spreading widely and varying; the new species thus produced being themselves dominant, owing to their having had some advantage over their already dominant parents, as well as over other species; and again spreading, varying, and producing new forms. The old forms which are beaten and which yield their places to the new and victorious forms, will generally be allied in groups, from inheriting some inferiority in common; and, therefore, as new and improved groups spread throughout the world, old groups disappear from the world; and the succession of forms everywhere tends to correspond both in their first appearance and final disappearance.</p>
 			<p>There is one other remark connected with this subject worth making. I have given my reasons for believing that most of our great formations, rich in fossils, were deposited during periods of subsidence; and that blank intervals of vast duration, as far as fossils are concerned, occurred during the periods when the bed of the sea was either stationary or rising, and likewise when sediment was not thrown down quickly enough to embed and preserve organic remains. During these long and blank intervals I suppose that the inhabitants of each region underwent a considerable amount of modification and extinction, and that there was much migration from other parts of the world. As we have reason to believe that large areas are affected by the same movement, it is probable that strictly contemporaneous formations have often been accumulated over very wide spaces in the same quarter of the world; but we are very far from having any right to conclude that this has invariably been the case, and that large areas have invariably been affected by the same movements. When two formations have been deposited in two regions during nearly, but not exactly, the same period, we should find in both, from the causes explained in the foregoing paragraphs, the same general succession in the forms of life; but the species would not exactly correspond; for there will have been a little more time in the one region than in the other for modification, extinction, and immigration.</p>
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 			<h3 epub:type="title">On the Affinities of Extinct Species to Each Other, and to Living Forms</h3>
-			<p>Let us now look to the mutual affinities of extinct and living species. All fall into a few grand classes; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, extinct species can all be classed either in still existing groups, or between them. That the extinct forms of life help to fill up the intervals between existing genera, families, and orders, is certainly true; but as this statement has often been ignored or even denied, it may be well to make some remarks on this subject, and to give some instances. If we confine our attention either to the living or to the extinct species of the same class, the series is far less perfect than if we combine both into one general system. In the writings of Professor Owen we continually meet with the expression of generalised forms, as applied to extinct animals; and in the writings of Agassiz, of prophetic or synthetic types; and these terms imply that such forms are, in fact, intermediate or connecting links. Another distinguished palaeontologist, M. Gaudry, has shown in the most striking manner that many of the fossil mammals discovered by him in Attica serve to break down the intervals between existing genera. Cuvier ranked the Ruminants and Pachyderms as two of the most distinct orders of mammals; but so many fossil links have been disentombed that Owen has had to alter the whole classification, and has placed certain Pachyderms in the same suborder with ruminants; for example, he dissolves by gradations the apparently wide interval between the pig and the camel. The Ungulata or hoofed quadrupeds are now divided into the even-toed or odd-toed divisions; but the Macrauchenia of South America connects to a certain extent these two grand divisions. No one will deny that the Hipparion is intermediate between the existing horse and certain other ungulate forms. What a wonderful connecting link in the chain of mammals is the Typotherium from South America, as the name given to it by Professor Gervais expresses, and which cannot be placed in any existing order. The Sirenia form a very distinct group of the mammals, and one of the most remarkable peculiarities in existing dugong and lamentin is the entire absence of hind limbs, without even a rudiment being left; but the extinct Halitherium had, according to Professor Flower, an ossified thighbone “articulated to a well-defined acetabulum in the pelvis,” and it thus makes some approach to ordinary hoofed quadrupeds, to which the Sirenia are in other respects allied. The cetaceans or whales are widely different from all other mammals, but the tertiary Zeuglodon and Squalodon, which have been placed by some naturalists in an order by themselves, are considered by Professor Huxley to be undoubtedly cetaceans, “and to constitute connecting links with the aquatic carnivora.”</p>
+			<p>Let us now look to the mutual affinities of extinct and living species. All fall into a few grand classes; and this fact is at once explained on the principle of descent. The more ancient any form is, the more, as a general rule, it differs from living forms. But, as Buckland long ago remarked, extinct species can all be classed either in still existing groups, or between them. That the extinct forms of life help to fill up the intervals between existing genera, families, and orders, is certainly true; but as this statement has often been ignored or even denied, it may be well to make some remarks on this subject, and to give some instances. If we confine our attention either to the living or to the extinct species of the same class, the series is far less perfect than if we combine both into one general system. In the writings of Professor Owen we continually meet with the expression of generalised forms, as applied to extinct animals; and in the writings of Agassiz, of prophetic or synthetic types; and these terms imply that such forms are, in fact, intermediate or connecting links. Another distinguished palaeontologist, <abbr class="name">M.</abbr> Gaudry, has shown in the most striking manner that many of the fossil mammals discovered by him in Attica serve to break down the intervals between existing genera. Cuvier ranked the Ruminants and Pachyderms as two of the most distinct orders of mammals; but so many fossil links have been disentombed that Owen has had to alter the whole classification, and has placed certain Pachyderms in the same suborder with ruminants; for example, he dissolves by gradations the apparently wide interval between the pig and the camel. The Ungulata or hoofed quadrupeds are now divided into the even-toed or odd-toed divisions; but the Macrauchenia of South America connects to a certain extent these two grand divisions. No one will deny that the Hipparion is intermediate between the existing horse and certain other ungulate forms. What a wonderful connecting link in the chain of mammals is the Typotherium from South America, as the name given to it by Professor Gervais expresses, and which cannot be placed in any existing order. The Sirenia form a very distinct group of the mammals, and one of the most remarkable peculiarities in existing dugong and lamentin is the entire absence of hind limbs, without even a rudiment being left; but the extinct Halitherium had, according to Professor Flower, an ossified thighbone “articulated to a well-defined acetabulum in the pelvis,” and it thus makes some approach to ordinary hoofed quadrupeds, to which the Sirenia are in other respects allied. The cetaceans or whales are widely different from all other mammals, but the tertiary Zeuglodon and Squalodon, which have been placed by some naturalists in an order by themselves, are considered by Professor Huxley to be undoubtedly cetaceans, “and to constitute connecting links with the aquatic carnivora.”</p>
 			<p>Even the wide interval between birds and reptiles has been shown by the naturalist just quoted to be partially bridged over in the most unexpected manner, on the one hand, by the ostrich and extinct Archeopteryx, and on the other hand by the Compsognathus, one of the Dinosaurians⁠—that group which includes the most gigantic of all terrestrial reptiles. Turning to the Invertebrata, Barrande asserts, a higher authority could not be named, that he is every day taught that, although palaeozoic animals can certainly be classed under existing groups, yet that at this ancient period the groups were not so distinctly separated from each other as they now are.</p>
 			<p>Some writers have objected to any extinct species, or group of species, being considered as intermediate between any two living species, or groups of species. If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms or groups, the objection is probably valid. But in a natural classification many fossil species certainly stand between living species, and some extinct genera between living genera, even between genera belonging to distinct families. The most common case, especially with respect to very distinct groups, such as fish and reptiles, seems to be that, supposing them to be distinguished at the present day by a score of characters, the ancient members are separated by a somewhat lesser number of characters, so that the two groups formerly made a somewhat nearer approach to each other than they now do.</p>
 			<p>It is a common belief that the more ancient a form is, by so much the more it tends to connect by some of its characters groups now widely separated from each other. This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages; and it would be difficult to prove the truth of the proposition, for every now and then even a living animal, as the Lepidosiren, is discovered having affinities directed towards very distinct groups. Yet if we compare the older Reptiles and Batrachians, the older Fish, the older Cephalopods, and the eocene Mammals, with the recent members of the same classes, we must admit that there is truth in the remark.</p>
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 			<h3 epub:type="title">Means of Dispersal</h3>
-			<p>Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration. A region now impassable to certain organisms from the nature of its climate, might have been a high road for migration, when the climate was different. I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas; submerge it, or let it formerly have been submerged, and the two faunas will now blend together, or may formerly have blended. Where the sea now extends, land may at a former period have connected islands or possibly even continents together, and thus have allowed terrestrial productions to pass from one to the other. No geologist disputes that great mutations of level have occurred within the period of existing organisms. Edward Forbes insisted that all the islands in the Atlantic must have been recently connected with Europe or Africa, and Europe likewise with America. Other authors have thus hypothetically bridged over every ocean, and united almost every island with some mainland. If, indeed, the arguments used by Forbes are to be trusted, it must be admitted that scarcely a single island exists which has not recently been united to some continent. This view cuts the Gordian knot of the dispersal of the same species to the most distant points, and removes many a difficulty; but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea; but not of such vast changes in the position and extension of our continents, as to have united them within the recent period to each other and to the several intervening oceanic islands. I freely admit the former existence of many islands, now buried beneath the sea, which may have served as halting places for plants and for many animals during their migration. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. Whenever it is fully admitted, as it will some day be, that each species has proceeded from a single birthplace, and when in the course of time we know something definite about the means of distribution, we shall be enabled to speculate with security on the former extension of the land. But I do not believe that it will ever be proved that within the recent period most of our continents which now stand quite separate, have been continuously, or almost continuously united with each other, and with the many existing oceanic islands. Several facts in distribution⁠—such as the great difference in the marine faunas on the opposite sides of almost every continent⁠—the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants⁠—the degree of affinity between the mammals inhabiting islands with those of the nearest continent, being in part determined (as we shall hereafter see) by the depth of the intervening ocean⁠—these and other such facts are opposed to the admission of such prodigious geographical revolutions within the recent period, as are necessary on the view advanced by Forbes and admitted by his followers. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity of continents. Nor does the almost universally volcanic composition of such islands favour the admission that they are the wrecks of sunken continents; if they had originally existed as continental mountain ranges, some at least of the islands would have been formed, like other mountain summits, of granite, metamorphic schists, old fossiliferous and other rocks, instead of consisting of mere piles of volcanic matter.</p>
+			<p>Sir <abbr class="name">C.</abbr> Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration. A region now impassable to certain organisms from the nature of its climate, might have been a high road for migration, when the climate was different. I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas; submerge it, or let it formerly have been submerged, and the two faunas will now blend together, or may formerly have blended. Where the sea now extends, land may at a former period have connected islands or possibly even continents together, and thus have allowed terrestrial productions to pass from one to the other. No geologist disputes that great mutations of level have occurred within the period of existing organisms. Edward Forbes insisted that all the islands in the Atlantic must have been recently connected with Europe or Africa, and Europe likewise with America. Other authors have thus hypothetically bridged over every ocean, and united almost every island with some mainland. If, indeed, the arguments used by Forbes are to be trusted, it must be admitted that scarcely a single island exists which has not recently been united to some continent. This view cuts the Gordian knot of the dispersal of the same species to the most distant points, and removes many a difficulty; but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea; but not of such vast changes in the position and extension of our continents, as to have united them within the recent period to each other and to the several intervening oceanic islands. I freely admit the former existence of many islands, now buried beneath the sea, which may have served as halting places for plants and for many animals during their migration. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. Whenever it is fully admitted, as it will some day be, that each species has proceeded from a single birthplace, and when in the course of time we know something definite about the means of distribution, we shall be enabled to speculate with security on the former extension of the land. But I do not believe that it will ever be proved that within the recent period most of our continents which now stand quite separate, have been continuously, or almost continuously united with each other, and with the many existing oceanic islands. Several facts in distribution⁠—such as the great difference in the marine faunas on the opposite sides of almost every continent⁠—the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants⁠—the degree of affinity between the mammals inhabiting islands with those of the nearest continent, being in part determined (as we shall hereafter see) by the depth of the intervening ocean⁠—these and other such facts are opposed to the admission of such prodigious geographical revolutions within the recent period, as are necessary on the view advanced by Forbes and admitted by his followers. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity of continents. Nor does the almost universally volcanic composition of such islands favour the admission that they are the wrecks of sunken continents; if they had originally existed as continental mountain ranges, some at least of the islands would have been formed, like other mountain summits, of granite, metamorphic schists, old fossiliferous and other rocks, instead of consisting of mere piles of volcanic matter.</p>
 			<p>I must now say a few words on what are called accidental means, but which more properly should be called occasional means of distribution. I shall here confine myself to plants. In botanical works, this or that plant is often stated to be ill adapted for wide dissemination; but the greater or less facilities for transport across the sea may be said to be almost wholly unknown. Until I tried, with <abbr>Mr.</abbr> Berkeley’s aid, a few experiments, it was not even known how far seeds could resist the injurious action of seawater. To my surprise I found that out of eighty-seven kinds, sixty-four germinated after an immersion of twenty-eight days, and a few survived an immersion of 137 days. It deserves notice that certain orders were far more injured than others: nine Leguminosae were tried, and, with one exception, they resisted the saltwater badly; seven species of the allied orders, Hydrophyllaceae and Polemoniaceae, were all killed by a month’s immersion. For convenience sake I chiefly tried small seeds without the capsules or fruit; and as all of these sank in a few days, they could not have been floated across wide spaces of the sea, whether or not they were injured by salt water. Afterwards I tried some larger fruits, capsules, <abbr>etc.</abbr>, and some of these floated for a long time. It is well known what a difference there is in the buoyancy of green and seasoned timber; and it occurred to me that floods would often wash into the sea dried plants or branches with seed-capsules or fruit attached to them. Hence I was led to dry the stems and branches of ninety-four plants with ripe fruit, and to place them on seawater. The majority sank quickly, but some which, whilst green, floated for a very short time, when dried floated much longer; for instance, ripe hazelnuts sank immediately, but when dried they floated for ninety days, and afterwards when planted germinated; an asparagus plant with ripe berries floated for twenty-three days, when dried it floated for eighty-five days, and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days, when dried they floated for above ninety days, and afterwards germinated. Altogether, out of the ninety-four dried plants, eighteen floated for above twenty-eight days; and some of the eighteen floated for a very much longer period. So that as 64/87 kinds of seeds germinated after an immersion of twenty-eight days; and as 18/94 distinct species with ripe fruit (but not all the same species as in the foregoing experiment) floated, after being dried, for above twenty-eight days, we may conclude, as far as anything can be inferred from these scanty facts, that the seeds of 14/100 kinds of plants of any country might be floated by sea-currents during twenty-eight days, and would retain their power of germination. In Johnston’s Physical Atlas, the average rate of the several Atlantic currents is thirty-three miles per diem (some currents running at the rate of sixty miles per diem); on this average, the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country; and when stranded, if blown by an inland gale to a favourable spot, would germinate.</p>
-			<p>Subsequently to my experiments, M. Martens tried similar ones, but in a much better manner, for he placed the seeds in a box in the actual sea, so that they were alternately wet and exposed to the air like really floating plants. He tried ninety-eight seeds, mostly different from mine, but he chose many large fruits, and likewise seeds, from plants which live near the sea; and this would have favoured both the average length of their flotation and their resistance to the injurious action of the saltwater. On the other hand, he did not previously dry the plants or branches with the fruit; and this, as we have seen, would have caused some of them to have floated much longer. The result was that 18/98 of his seeds of different kinds floated for forty-two days, and were then capable of germination. But I do not doubt that plants exposed to the waves would float for a less time than those protected from violent movement as in our experiments. Therefore, it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora, after having been dried, could be floated across a space of sea 900 miles in width, and would then germinate. The fact of the larger fruits often floating longer than the small, is interesting; as plants with large seeds or fruit which, as Alph. de Candolle has shown, generally have restricted ranges, could hardly be transported by any other means.</p>
+			<p>Subsequently to my experiments, <abbr class="name">M.</abbr> Martens tried similar ones, but in a much better manner, for he placed the seeds in a box in the actual sea, so that they were alternately wet and exposed to the air like really floating plants. He tried ninety-eight seeds, mostly different from mine, but he chose many large fruits, and likewise seeds, from plants which live near the sea; and this would have favoured both the average length of their flotation and their resistance to the injurious action of the saltwater. On the other hand, he did not previously dry the plants or branches with the fruit; and this, as we have seen, would have caused some of them to have floated much longer. The result was that 18/98 of his seeds of different kinds floated for forty-two days, and were then capable of germination. But I do not doubt that plants exposed to the waves would float for a less time than those protected from violent movement as in our experiments. Therefore, it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora, after having been dried, could be floated across a space of sea 900 miles in width, and would then germinate. The fact of the larger fruits often floating longer than the small, is interesting; as plants with large seeds or fruit which, as Alph. de Candolle has shown, generally have restricted ranges, could hardly be transported by any other means.</p>
 			<p>Seeds may be occasionally transported in another manner. Drift timber is thrown up on most islands, even on those in the midst of the widest oceans; and the natives of the coral islands in the Pacific procure stones for their tools, solely from the roots of drifted trees, these stones being a valuable royal tax. I find that when irregularly shaped stones are embedded in the roots of trees, small parcels of earth are very frequently enclosed in their interstices and behind them, so perfectly that not a particle could be washed away during the longest transport: out of one small portion of earth thus <em>completely</em> enclosed by the roots of an oak about fifty years old, three dicotyledonous plants germinated: I am certain of the accuracy of this observation. Again, I can show that the carcasses of birds, when floating on the sea, sometimes escape being immediately devoured; and many kinds of seeds in the crops of floating birds long retain their vitality: peas and vetches, for instance, are killed by even a few days’ immersion in seawater; but some taken out of the crop of a pigeon, which had floated on artificial seawater for thirty days, to my surprise nearly all germinated.</p>
 			<p>Living birds can hardly fail to be highly effective agents in the transportation of seeds. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. We may safely assume that under such circumstances their rate of flight would often be thirty-five miles an hour; and some authors have given a far higher estimate. I have never seen an instance of nutritious seeds passing through the intestines of a bird; but hard seeds of fruit pass uninjured through even the digestive organs of a turkey. In the course of two months, I picked up in my garden twelve kinds of seeds, out of the excrement of small birds, and these seemed perfect, and some of them, which were tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not, as I know by trial, injure in the least the germination of seeds; now, after a bird has found and devoured a large supply of food, it is positively asserted that all the grains do not pass into the gizzard for twelve or even eighteen hours. A bird in this interval might easily be blown to the distance of five hundred miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. Some hawks and owls bolt their prey whole, and after an interval of from twelve to twenty hours, disgorge pellets, which, as I know from experiments made in the Zoological Gardens, include seeds capable of germination. Some seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey; and two seeds of beet grew after having been thus retained for two days and fourteen hours. Freshwater fish, I find, eat seeds of many land and water plants; fish are frequently devoured by birds, and thus the seeds might be transported from place to place. I forced many kinds of seeds into the stomachs of dead fish, and then gave their bodies to fishing-eagles, storks, and pelicans; these birds, after an interval of many hours, either rejected the seeds in pellets or passed them in their excrement; and several of these seeds retained the power of germination. Certain seeds, however, were always killed by this process.</p>
-			<p>Locusts are sometimes blown to great distances from the land. I myself caught one 370 miles from the coast of Africa, and have heard of others caught at greater distances. The <abbr>Rev.</abbr> <abbr class="name">R. T.</abbr> Lowe informed Sir C. Lyell that in November, 1844, swarms of locusts visited the island of Madeira. They were in countless numbers, as thick as the flakes of snow in the heaviest snowstorm, and extended upward as far as could be seen with a telescope. During two or three days they slowly careered round and round in an immense ellipse, at least five or six miles in diameter, and at night alighted on the taller trees, which were completely coated with them. They then disappeared over the sea, as suddenly as they had appeared, and have not since visited the island. Now, in parts of Natal it is believed by some farmers, though on insufficient evidence, that injurious seeds are introduced into their grassland in the dung left by the great flights of locusts which often visit that country. In consequence of this belief <abbr>Mr.</abbr> Weale sent me in a letter a small packet of the dried pellets, out of which I extracted under the microscope several seeds, and raised from them seven grass plants, belonging to two species, of two genera. Hence a swarm of locusts, such as that which visited Madeira, might readily be the means of introducing several kinds of plants into an island lying far from the mainland.</p>
+			<p>Locusts are sometimes blown to great distances from the land. I myself caught one 370 miles from the coast of Africa, and have heard of others caught at greater distances. The <abbr>Rev.</abbr> <abbr class="name">R. T.</abbr> Lowe informed Sir <abbr class="name">C.</abbr> Lyell that in November, 1844, swarms of locusts visited the island of Madeira. They were in countless numbers, as thick as the flakes of snow in the heaviest snowstorm, and extended upward as far as could be seen with a telescope. During two or three days they slowly careered round and round in an immense ellipse, at least five or six miles in diameter, and at night alighted on the taller trees, which were completely coated with them. They then disappeared over the sea, as suddenly as they had appeared, and have not since visited the island. Now, in parts of Natal it is believed by some farmers, though on insufficient evidence, that injurious seeds are introduced into their grassland in the dung left by the great flights of locusts which often visit that country. In consequence of this belief <abbr>Mr.</abbr> Weale sent me in a letter a small packet of the dried pellets, out of which I extracted under the microscope several seeds, and raised from them seven grass plants, belonging to two species, of two genera. Hence a swarm of locusts, such as that which visited Madeira, might readily be the means of introducing several kinds of plants into an island lying far from the mainland.</p>
 			<p>Although the beaks and feet of birds are generally clean, earth sometimes adheres to them: in one case I removed sixty-one grains, and in another case twenty-two grains of dry argillaceous earth from the foot of a partridge, and in the earth there was a pebble as large as the seed of a vetch. Here is a better case: the leg of a woodcock was sent to me by a friend, with a little cake of dry earth attached to the shank, weighing only nine grains; and this contained a seed of the toad-rush (Juncus bufonius) which germinated and flowered. <abbr>Mr.</abbr> Swaysland, of Brighton, who during the last forty years has paid close attention to our migratory birds, informs me that he has often shot wagtails (Motacillae), wheatears, and whinchats (Saxicolae), on their first arrival on our shores, before they had alighted; and he has several times noticed little cakes of earth attached to their feet. Many facts could be given showing how generally soil is charged with seeds. For instance, Professor Newton sent me the leg of a red-legged partridge (Caccabis rufa) which had been wounded and could not fly, with a ball of hard earth adhering to it, and weighing six and a half ounces. The earth had been kept for three years, but when broken, watered and placed under a bell glass, no less than eighty-two plants sprung from it: these consisted of twelve monocotyledons, including the common oat, and at least one kind of grass, and of seventy dicotyledons, which consisted, judging from the young leaves, of at least three distinct species. With such facts before us, can we doubt that the many birds which are annually blown by gales across great spaces of ocean, and which annually migrate⁠—for instance, the millions of quails across the Mediterranean⁠—must occasionally transport a few seeds embedded in dirt adhering to their feet or beaks? But I shall have to recur to this subject.</p>
-			<p>As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, it can hardly be doubted that they must occasionally, as suggested by Lyell, have transported seeds from one part to another of the arctic and antarctic regions; and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of plants common to Europe, in comparison with the species on the other islands of the Atlantic, which stand nearer to the mainland, and (as remarked by <abbr>Mr.</abbr> <abbr class="name">H. C.</abbr> Watson) from their somewhat northern character, in comparison with the latitude, I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch. At my request Sir C. Lyell wrote to M. Hartung to inquire whether he had observed erratic boulders on these islands, and he answered that he had found large fragments of granite and other rocks, which do not occur in the archipelago. Hence we may safely infer that icebergs formerly landed their rocky burdens on the shores of these mid-ocean islands, and it is at least possible that they may have brought thither the seeds of northern plants.</p>
+			<p>As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, it can hardly be doubted that they must occasionally, as suggested by Lyell, have transported seeds from one part to another of the arctic and antarctic regions; and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of plants common to Europe, in comparison with the species on the other islands of the Atlantic, which stand nearer to the mainland, and (as remarked by <abbr>Mr.</abbr> <abbr class="name">H. C.</abbr> Watson) from their somewhat northern character, in comparison with the latitude, I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch. At my request Sir <abbr class="name">C.</abbr> Lyell wrote to <abbr class="name">M.</abbr> Hartung to inquire whether he had observed erratic boulders on these islands, and he answered that he had found large fragments of granite and other rocks, which do not occur in the archipelago. Hence we may safely infer that icebergs formerly landed their rocky burdens on the shores of these mid-ocean islands, and it is at least possible that they may have brought thither the seeds of northern plants.</p>
 			<p>Considering that these several means of transport, and that other means, which without doubt remain to be discovered, have been in action year after year for tens of thousands of years, it would, I think, be a marvellous fact if many plants had not thus become widely transported. These means of transport are sometimes called accidental, but this is not strictly correct: the currents of the sea are not accidental, nor is the direction of prevalent gales of wind. It should be observed that scarcely any means of transport would carry seeds for very great distances; for seeds do not retain their vitality when exposed for a great length of time to the action of sea water; nor could they be long carried in the crops or intestines of birds. These means, however, would suffice for occasional transport across tracts of sea some hundred miles in breadth, or from island to island, or from a continent to a neighbouring island, but not from one distant continent to another. The floras of distant continents would not by such means become mingled; but would remain as distinct as they now are. The currents, from their course, would never bring seeds from North America to Britain, though they might and do bring seeds from the West Indies to our western shores, where, if not killed by their very long immersion in salt water, they could not endure our climate. Almost every year, one or two land-birds are blown across the whole Atlantic Ocean, from North America to the western shores of Ireland and England; but seeds could be transported by these rare wanderers only by one means, namely, by dirt adhering to their feet or beaks, which is in itself a rare accident. Even in this case, how small would be the chance of a seed falling on favourable soil, and coming to maturity! But it would be a great error to argue that because a well-stocked island, like Great Britain, has not, as far as is known (and it would be very difficult to prove this), received within the last few centuries, through occasional means of transport, immigrants from Europe or any other continent, that a poorly-stocked island, though standing more remote from the mainland, would not receive colonists by similar means. Out of a hundred kinds of seeds or animals transported to an island, even if far less well-stocked than Britain, perhaps not more than one would be so well fitted to its new home, as to become naturalised. But this is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, whilst the island was being upheaved, and before it had become fully stocked with inhabitants. On almost bare land, with few or no destructive insects or birds living there, nearly every seed which chanced to arrive, if fitted for the climate, would germinate and survive.</p>
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 			<h3 epub:type="title">Alternate Glacial Periods in the North and South</h3>
 			<p>But we must return to our more immediate subject. I am convinced that Forbes’s view may be largely extended. In Europe we meet with the plainest evidence of the Glacial period, from the western shores of Britain to the Ural range, and southward to the Pyrenees. We may infer from the frozen mammals and nature of the mountain vegetation, that Siberia was similarly affected. In the Lebanon, according to <abbr>Dr.</abbr> Hooker, perpetual snow formerly covered the central axis, and fed glaciers which rolled 4,000 feet down the valleys. The same observer has recently found great moraines at a low level on the Atlas range in North Africa. Along the Himalaya, at points 900 miles apart, glaciers have left the marks of their former low descent; and in Sikkim, <abbr>Dr.</abbr> Hooker saw maize growing on ancient and gigantic moraines. Southward of the Asiatic continent, on the opposite side of the equator, we know, from the excellent researches of <abbr>Dr.</abbr> <abbr class="name">J.</abbr> Haast and <abbr>Dr.</abbr> Hector, that in New Zealand immense glaciers formerly descended to a low level; and the same plants, found by <abbr>Dr.</abbr> Hooker on widely separated mountains in this island tell the same story of a former cold period. From facts communicated to me by the <abbr>Rev.</abbr> <abbr class="name">W. B.</abbr> Clarke, it appears also that there are traces of former glacial action on the mountains of the southeastern corner of Australia.</p>
 			<p>Looking to America: in the northern half, ice-borne fragments of rock have been observed on the eastern side of the continent, as far south as latitude 36 and 37 degrees, and on the shores of the Pacific, where the climate is now so different, as far south as latitude 46 degrees. Erratic boulders have, also, been noticed on the Rocky Mountains. In the Cordillera of South America, nearly under the equator, glaciers once extended far below their present level. In central Chile I examined a vast mound of detritus with great boulders, crossing the Portillo valley, which, there can hardly be a doubt, once formed a huge moraine; and <abbr>Mr.</abbr> <abbr class="name">D.</abbr> Forbes informs me that he found in various parts of the Cordillera, from latitude 13 to 30 degrees south, at about the height of 12,000 feet, deeply-furrowed rocks, resembling those with which he was familiar in Norway, and likewise great masses of detritus, including grooved pebbles. Along this whole space of the Cordillera true glaciers do not now exist even at much more considerable heights. Further south, on both sides of the continent, from latitude 41 degrees to the southernmost extremity, we have the clearest evidence of former glacial action, in numerous immense boulders transported far from their parent source.</p>
-			<p>From these several facts, namely, from the glacial action having extended all round the northern and southern hemispheres⁠—from the period having been in a geological sense recent in both hemispheres⁠—from its having lasted in both during a great length of time, as may be inferred from the amount of work effected⁠—and lastly, from glaciers having recently descended to a low level along the whole line of the Cordillera, it at one time appeared to me that we could not avoid the conclusion that the temperature of the whole world had been simultaneously lowered during the Glacial period. But now, <abbr>Mr.</abbr> Croll, in a series of admirable memoirs, has attempted to show that a glacial condition of climate is the result of various physical causes, brought into operation by an increase in the eccentricity of the earth’s orbit. All these causes tend towards the same end; but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. According to <abbr>Mr.</abbr> Croll, cold periods regularly recur every ten or fifteen thousand years; and these at long intervals are extremely severe, owing to certain contingencies, of which the most important, as Sir C. Lyell has shown, is the relative position of the land and water. <abbr>Mr.</abbr> Croll believes that the last great glacial period occurred about 240,000 years ago, and endured, with slight alterations of climate, for about 160,000 years. With respect to more ancient glacial periods, several geologists are convinced, from direct evidence, that such occurred during the miocene and eocene formations, not to mention still more ancient formations. But the most important result for us, arrived at by <abbr>Mr.</abbr> Croll, is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised, with the winters rendered much milder, chiefly through changes in the direction of the ocean currents. So conversely it will be with the northern hemisphere, while the southern passes through a glacial period. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it; but I will first give the facts which demand an explanation.</p>
+			<p>From these several facts, namely, from the glacial action having extended all round the northern and southern hemispheres⁠—from the period having been in a geological sense recent in both hemispheres⁠—from its having lasted in both during a great length of time, as may be inferred from the amount of work effected⁠—and lastly, from glaciers having recently descended to a low level along the whole line of the Cordillera, it at one time appeared to me that we could not avoid the conclusion that the temperature of the whole world had been simultaneously lowered during the Glacial period. But now, <abbr>Mr.</abbr> Croll, in a series of admirable memoirs, has attempted to show that a glacial condition of climate is the result of various physical causes, brought into operation by an increase in the eccentricity of the earth’s orbit. All these causes tend towards the same end; but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. According to <abbr>Mr.</abbr> Croll, cold periods regularly recur every ten or fifteen thousand years; and these at long intervals are extremely severe, owing to certain contingencies, of which the most important, as Sir <abbr class="name">C.</abbr> Lyell has shown, is the relative position of the land and water. <abbr>Mr.</abbr> Croll believes that the last great glacial period occurred about 240,000 years ago, and endured, with slight alterations of climate, for about 160,000 years. With respect to more ancient glacial periods, several geologists are convinced, from direct evidence, that such occurred during the miocene and eocene formations, not to mention still more ancient formations. But the most important result for us, arrived at by <abbr>Mr.</abbr> Croll, is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised, with the winters rendered much milder, chiefly through changes in the direction of the ocean currents. So conversely it will be with the northern hemisphere, while the southern passes through a glacial period. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it; but I will first give the facts which demand an explanation.</p>
 			<p>In South America, <abbr>Dr.</abbr> Hooker has shown that besides many closely allied species, between forty and fifty of the flowering plants of Tierra del Fuego, forming no inconsiderable part of its scanty flora, are common to North America and Europe, enormously remote as these areas in opposite hemispheres are from each other. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. On the Organ Mountains of Brazil some few temperate European, some Antarctic and some Andean genera were found by Gardner which do not exist in the low intervening hot countries. On the Silla of Caraccas the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera.</p>
 			<p>In Africa, several forms characteristic of Europe, and some few representatives of the flora of the Cape of Good Hope, occur on the mountains of Abyssinia. At the Cape of Good Hope a very few European species, believed not to have been introduced by man, and on the mountains several representative European forms are found which have not been discovered in the intertropical parts of Africa. <abbr>Dr.</abbr> Hooker has also lately shown that several of the plants living on the upper parts of the lofty island of Fernando Po, and on the neighbouring Cameroon Mountains, in the Gulf of Guinea, are closely related to those on the mountains of Abyssinia, and likewise to those of temperate Europe. It now also appears, as I hear from <abbr>Dr.</abbr> Hooker, that some of these same temperate plants have been discovered by the <abbr>Rev.</abbr> <abbr class="name">R. T.</abbr> Lowe on the mountains of the Cape Verde Islands. This extension of the same temperate forms, almost under the equator, across the whole continent of Africa and to the mountains of the Cape Verde archipelago, is one of the most astonishing facts ever recorded in the distribution of plants.</p>
 			<p>On the Himalaya, and on the isolated mountain ranges of the peninsula of India, on the heights of Ceylon, and on the volcanic cones of Java, many plants occur either identically the same or representing each other, and at the same time representing plants of Europe not found in the intervening hot lowlands. A list of the genera of plants collected on the loftier peaks of Java, raises a picture of a collection made on a hillock in Europe. Still more striking is the fact that peculiar Australian forms are represented by certain plants growing on the summits of the mountains of Borneo. Some of these Australian forms, as I hear from <abbr>Dr.</abbr> Hooker, extend along the heights of the peninsula of Malacca, and are thinly scattered on the one hand over India, and on the other hand as far north as Japan.</p>
 			<p>On the southern mountains of Australia, <abbr>Dr.</abbr> <abbr class="name">F.</abbr> Muller has discovered several European species; other species, not introduced by man, occur on the lowlands; and a long list can be given, as I am informed by <abbr>Dr.</abbr> Hooker, of European genera, found in Australia, but not in the intermediate torrid regions. In the admirable “Introduction to the Flora of New Zealand,” by <abbr>Dr.</abbr> Hooker, analogous and striking facts are given in regard to the plants of that large island. Hence, we see that certain plants growing on the more lofty mountains of the tropics in all parts of the world, and on the temperate plains of the north and south, are either the same species or varieties of the same species. It should, however, be observed that these plants are not strictly arctic forms; for, as <abbr>Mr.</abbr> <abbr class="name">H. C.</abbr> Watson has remarked, “in receding from polar toward equatorial latitudes, the Alpine or mountain flora really become less and less Arctic.” Besides these identical and closely allied forms, many species inhabiting the same widely sundered areas, belong to genera not now found in the intermediate tropical lowlands.</p>
-			<p>These brief remarks apply to plants alone; but some few analogous facts could be given in regard to terrestrial animals. In marine productions, similar cases likewise occur; as an example, I may quote a statement by the highest authority, <abbr>Prof.</abbr> Dana, that “it is certainly a wonderful fact that New Zealand should have a closer resemblance in its crustacea to Great Britain, its antipode, than to any other part of the world.” Sir J. Richardson, also, speaks of the reappearance on the shores of New Zealand, Tasmania, <abbr>etc.</abbr>, of northern forms of fish. <abbr>Dr.</abbr> Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe, but have not been found in the intermediate tropical seas.</p>
+			<p>These brief remarks apply to plants alone; but some few analogous facts could be given in regard to terrestrial animals. In marine productions, similar cases likewise occur; as an example, I may quote a statement by the highest authority, <abbr>Prof.</abbr> Dana, that “it is certainly a wonderful fact that New Zealand should have a closer resemblance in its crustacea to Great Britain, its antipode, than to any other part of the world.” Sir <abbr class="name">J.</abbr> Richardson, also, speaks of the reappearance on the shores of New Zealand, Tasmania, <abbr>etc.</abbr>, of northern forms of fish. <abbr>Dr.</abbr> Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe, but have not been found in the intermediate tropical seas.</p>
 			<p>From the foregoing facts, namely, the presence of temperate forms on the highlands across the whole of equatorial Africa, and along the Peninsula of India, to Ceylon and the Malay Archipelago, and in a less well-marked manner across the wide expanse of tropical South America, it appears almost certain that at some former period, no doubt during the most severe part of a Glacial period, the lowlands of these great continents were everywhere tenanted under the equator by a considerable number of temperate forms. At this period the equatorial climate at the level of the sea was probably about the same with that now experienced at the height of from five to six thousand feet under the same latitude, or perhaps even rather cooler. During this, the coldest period, the lowlands under the equator must have been clothed with a mingled tropical and temperate vegetation, like that described by Hooker as growing luxuriantly at the height of from four to five thousand feet on the lower slopes of the Himalaya, but with perhaps a still greater preponderance of temperate forms. So again in the mountainous island of Fernando Po, in the Gulf of Guinea, <abbr>Mr.</abbr> Mann found temperate European forms beginning to appear at the height of about five thousand feet. On the mountains of Panama, at the height of only two thousand feet, <abbr>Dr.</abbr> Seemann found the vegetation like that of Mexico, “with forms of the torrid zone harmoniously blended with those of the temperate.”</p>
 			<p>Now let us see whether <abbr>Mr.</abbr> Croll’s conclusion that when the northern hemisphere suffered from the extreme cold of the great Glacial period, the southern hemisphere was actually warmer, throws any clear light on the present apparently inexplicable distribution of various organisms in the temperate parts of both hemispheres, and on the mountains of the tropics. The Glacial period, as measured by years, must have been very long; and when we remember over what vast spaces some naturalised plants and animals have spread within a few centuries, this period will have been ample for any amount of migration. As the cold became more and more intense, we know that Arctic forms invaded the temperate regions; and from the facts just given, there can hardly be a doubt that some of the more vigorous, dominant and widest-spreading temperate forms invaded the equatorial lowlands. The inhabitants of these hot lowlands would at the same time have migrated to the tropical and subtropical regions of the south, for the southern hemisphere was at this period warmer. On the decline of the Glacial period, as both hemispheres gradually recovered their former temperature, the northern temperate forms living on the lowlands under the equator, would have been driven to their former homes or have been destroyed, being replaced by the equatorial forms returning from the south. Some, however, of the northern temperate forms would almost certainly have ascended any adjoining high land, where, if sufficiently lofty, they would have long survived like the Arctic forms on the mountains of Europe. They might have survived, even if the climate was not perfectly fitted for them, for the change of temperature must have been very slow, and plants undoubtedly possess a certain capacity for acclimatisation, as shown by their transmitting to their offspring different constitutional powers of resisting heat and cold.</p>
 			<p>In the regular course of events the southern hemisphere would in its turn be subjected to a severe Glacial period, with the northern hemisphere rendered warmer; and then the southern temperate forms would invade the equatorial lowlands. The northern forms which had before been left on the mountains would now descend and mingle with the southern forms. These latter, when the warmth returned, would return to their former homes, leaving some few species on the mountains, and carrying southward with them some of the northern temperate forms which had descended from their mountain fastnesses. Thus, we should have some few species identically the same in the northern and southern temperate zones and on the mountains of the intermediate tropical regions. But the species left during a long time on these mountains, or in opposite hemispheres, would have to compete with many new forms and would be exposed to somewhat different physical conditions; hence, they would be eminently liable to modification, and would generally now exist as varieties or as representative species; and this is the case. We must, also, bear in mind the occurrence in both hemispheres of former Glacial periods; for these will account, in accordance with the same principles, for the many quite distinct species inhabiting the same widely separated areas, and belonging to genera not now found in the intermediate torrid zones.</p>
@@ -73,8 +73,7 @@
 			<p>The same principles apply to the distribution of terrestrial animals and of marine productions, in the northern and southern temperate zones, and on the intertropical mountains. When, during the height of the Glacial period, the ocean-currents were widely different to what they now are, some of the inhabitants of the temperate seas might have reached the equator; of these a few would perhaps at once be able to migrate southwards, by keeping to the cooler currents, while others might remain and survive in the colder depths until the southern hemisphere was in its turn subjected to a glacial climate and permitted their further progress; in nearly the same manner as, according to Forbes, isolated spaces inhabited by Arctic productions exist to the present day in the deeper parts of the northern temperate seas.</p>
 			<p>I am far from supposing that all the difficulties in regard to the distribution and affinities of the identical and allied species, which now live so widely separated in the north and south, and sometimes on the intermediate mountain ranges, are removed on the views above given. The exact lines of migration cannot be indicated. We cannot say why certain species and not others have migrated; why certain species have been modified and have given rise to new forms, while others have remained unaltered. We cannot hope to explain such facts, until we can say why one species and not another becomes naturalised by man’s agency in a foreign land; why one species ranges twice or thrice as far, and is twice or thrice as common, as another species within their own homes.</p>
 			<p>Various special difficulties also remain to be solved; for instance, the occurrence, as shown by <abbr>Dr.</abbr> Hooker, of the same plants at points so enormously remote as Kerguelen Land, New Zealand, and Fuegia; but icebergs, as suggested by Lyell, may have been concerned in their dispersal. The existence at these and other distant points of the southern hemisphere, of species, which, though distinct, belong to genera exclusively confined to the south, is a more remarkable case. Some of these species are so distinct, that we cannot suppose that there has been time since the commencement of the last Glacial period for their migration and subsequent modification to the necessary degree. The facts seem to indicate that distinct species belonging to the same genera have migrated in radiating lines from a common centre; and I am inclined to look in the southern, as in the northern hemisphere, to a former and warmer period, before the commencement of the last Glacial period, when the Antarctic lands, now covered with ice, supported a highly peculiar and isolated flora. It may be suspected that before this flora was exterminated during the last Glacial epoch, a few forms had been already widely dispersed to various points of the southern hemisphere by occasional means of transport, and by the aid, as halting-places, of now sunken islands. Thus the southern shores of America, Australia, and New Zealand may have become slightly tinted by the same peculiar forms of life.</p>
-			<p>Sir C. Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alternations of climate throughout the world on geographical distribution. And we have now seen that <abbr>Mr.</abbr> Croll’s conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere, together with the admission of the slow modification of species, explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. The living waters have flowed during one period from the north and during another from the south, and in both cases have reached the equator; but the stream of life has flowed with greater force from the north than in the opposite direction, and has consequently more freely inundated the south. As the tide leaves its drift in horizontal lines, rising higher on the shores where the tide rises highest, so have the living waters left their living drift on our mountain summits, in a line gently rising from the Arctic lowlands to a great latitude under the equator. The various beings thus left stranded may be compared with savage races of man, driven up and surviving in the mountain fastnesses of almost every land, which serves as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.</p>
+			<p>Sir <abbr class="name">C.</abbr> Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alternations of climate throughout the world on geographical distribution. And we have now seen that <abbr>Mr.</abbr> Croll’s conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere, together with the admission of the slow modification of species, explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. The living waters have flowed during one period from the north and during another from the south, and in both cases have reached the equator; but the stream of life has flowed with greater force from the north than in the opposite direction, and has consequently more freely inundated the south. As the tide leaves its drift in horizontal lines, rising higher on the shores where the tide rises highest, so have the living waters left their living drift on our mountain summits, in a line gently rising from the Arctic lowlands to a great latitude under the equator. The various beings thus left stranded may be compared with savage races of man, driven up and surviving in the mountain fastnesses of almost every land, which serves as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.</p>
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 			<p>The most striking and important fact for us is the affinity of the species which inhabit islands to those of the nearest mainland, without being actually the same. Numerous instances could be given. The Galapagos Archipelago, situated under the equator, lies at a distance of between 500 and 600 miles from the shores of South America. Here almost every product of the land and of the water bears the unmistakable stamp of the American continent. There are twenty-six land birds. Of these twenty-one, or perhaps twenty-three, are ranked as distinct species, and would commonly be assumed to have been here created; yet the close affinity of most of these birds to American species is manifest in every character in their habits, gestures, and tones of voice. So it is with the other animals, and with a large proportion of the plants, as shown by <abbr>Dr.</abbr> Hooker in his admirable Flora of this archipelago. The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, feels that he is standing on American land. Why should this be so? Why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plainly the stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which closely resembles the conditions of the South American coast. In fact, there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in the climate, height, and size of the islands, between the Galapagos and Cape Verde Archipelagos: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape Verde Islands are related to those of Africa, like those of the Galapagos to America. Facts, such as these, admit of no sort of explanation on the ordinary view of independent creation; whereas, on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists from America, whether by occasional means of transport or (though I do not believe in this doctrine) by formerly continuous land, and the Cape Verde Islands from Africa; such colonists would be liable to modification⁠—the principle of inheritance still betraying their original birthplace.</p>
 			<p>Many analogous facts could be given: indeed it is an almost universal rule that the endemic productions of islands are related to those of the nearest continent, or of the nearest large island. The exceptions are few, and most of them can be explained. Thus, although Kerguelen Land stands nearer to Africa than to America, the plants are related, and that very closely, as we know from <abbr>Dr.</abbr> Hooker’s account, to those of America: but on the view that this island has been mainly stocked by seeds brought with earth and stones on icebergs, drifted by the prevailing currents, this anomaly disappears. New Zealand in its endemic plants is much more closely related to Australia, the nearest mainland, than to any other region: and this is what might have been expected; but it is also plainly related to South America, which, although the next nearest continent, is so enormously remote, that the fact becomes an anomaly. But this difficulty partially disappears on the view that New Zealand, South America, and the other southern lands, have been stocked in part from a nearly intermediate though distant point, namely, from the antarctic islands, when they were clothed with vegetation, during a warmer tertiary period, before the commencement of the last Glacial period. The affinity, which, though feeble, I am assured by <abbr>Dr.</abbr> Hooker is real, between the flora of the southwestern corner of Australia and of the Cape of Good Hope, is a far more remarkable case; but this affinity is confined to the plants, and will, no doubt, some day be explained.</p>
 			<p>The same law which has determined the relationship between the inhabitants of islands and the nearest mainland, is sometimes displayed on a small scale, but in a most interesting manner, within the limits of the same archipelago. Thus each separate island of the Galapagos Archipelago is tenanted, and the fact is a marvellous one, by many distinct species; but these species are related to each other in a very much closer manner than to the inhabitants of the American continent, or of any other quarter of the world. This is what might have been expected, for islands situated so near to each other would almost necessarily receive immigrants from the same original source, and from each other. But how is it that many of the immigrants have been differently modified, though only in a small degree, in islands situated within sight of each other, having the same geological nature, the same height, climate, etc? This long appeared to me a great difficulty: but it arises in chief part from the deeply-seated error of considering the physical conditions of a country as the most important; whereas it cannot be disputed that the nature of the other species with which each has to compete, is at least as important, and generally a far more important element of success. Now if we look to the species which inhabit the Galapagos Archipelago, and are likewise found in other parts of the world, we find that they differ considerably in the several islands. This difference might indeed have been expected if the islands have been stocked by occasional means of transport⁠—a seed, for instance, of one plant having been brought to one island, and that of another plant to another island, though all proceeding from the same general source. Hence, when in former times an immigrant first settled on one of the islands, or when it subsequently spread from one to another, it would undoubtedly be exposed to different conditions in the different islands, for it would have to compete with a different set of organisms; a plant, for instance, would find the ground best-fitted for it occupied by somewhat different species in the different islands, and would be exposed to the attacks of somewhat different enemies. If, then, it varied, natural selection would probably favour different varieties in the different islands. Some species, however, might spread and yet retain the same character throughout the group, just as we see some species spreading widely throughout a continent and remaining the same.</p>
-			<p>The really surprising fact in this case of the Galapagos Archipelago, and in a lesser degree in some analogous cases, is that each new species after being formed in any one island, did not spread quickly to the other islands. But the islands, though in sight of each other, are separated by deep arms of the sea, in most cases wider than the British Channel, and there is no reason to suppose that they have at any former period been continuously united. The currents of the sea are rapid and deep between the islands, and gales of wind are extraordinarily rare; so that the islands are far more effectually separated from each other than they appear on a map. Nevertheless, some of the species, both of those found in other parts of the world and of those confined to the archipelago, are common to the several islands; and we may infer from the present manner of distribution that they have spread from one island to the others. But we often take, I think, an erroneous view of the probability of closely allied species invading each other’s territory, when put into free intercommunication. Undoubtedly, if one species has any advantage over another, it will in a very brief time wholly or in part supplant it; but if both are equally well fitted for their own places, both will probably hold their separate places for almost any length of time. Being familiar with the fact that many species, naturalised through man’s agency, have spread with astonishing rapidity over wide areas, we are apt to infer that most species would thus spread; but we should remember that the species which become naturalised in new countries are not generally closely allied to the aboriginal inhabitants, but are very distinct forms, belonging in a large proportion of cases, as shown by Alph. de Candolle, to distinct genera. In the Galapagos Archipelago, many even of the birds, though so well adapted for flying from island to island, differ on the different islands; thus there are three closely allied species of mocking-thrush, each confined to its own island. Now let us suppose the mocking-thrush of Chatham Island to be blown to Charles Island, which has its own mocking-thrush; why should it succeed in establishing itself there? We may safely infer that Charles Island is well stocked with its own species, for annually more eggs are laid and young birds hatched than can possibly be reared; and we may infer that the mocking-thrush peculiar to Charles Island is at least as well fitted for its home as is the species peculiar to Chatham Island. Sir C. Lyell and <abbr>Mr.</abbr> Wollaston have communicated to me a remarkable fact bearing on this subject; namely, that Madeira and the adjoining islet of Porto Santo possess many distinct but representative species of land-shells, some of which live in crevices of stone; and although large quantities of stone are annually transported from Porto Santo to Madeira, yet this latter island has not become colonised by the Porto Santo species: nevertheless, both islands have been colonised by some European land-shells, which no doubt had some advantage over the indigenous species. From these considerations I think we need not greatly marvel at the endemic species which inhabit the several islands of the Galapagos Archipelago not having all spread from island to island. On the same continent, also, preoccupation has probably played an important part in checking the commingling of the species which inhabit different districts with nearly the same physical conditions. Thus, the southeast and southwest corners of Australia have nearly the same physical conditions, and are united by continuous land, yet they are inhabited by a vast number of distinct mammals, birds, and plants; so it is, according to <abbr>Mr.</abbr> Bates, with the butterflies and other animals inhabiting the great, open, and continuous valley of the Amazons.</p>
+			<p>The really surprising fact in this case of the Galapagos Archipelago, and in a lesser degree in some analogous cases, is that each new species after being formed in any one island, did not spread quickly to the other islands. But the islands, though in sight of each other, are separated by deep arms of the sea, in most cases wider than the British Channel, and there is no reason to suppose that they have at any former period been continuously united. The currents of the sea are rapid and deep between the islands, and gales of wind are extraordinarily rare; so that the islands are far more effectually separated from each other than they appear on a map. Nevertheless, some of the species, both of those found in other parts of the world and of those confined to the archipelago, are common to the several islands; and we may infer from the present manner of distribution that they have spread from one island to the others. But we often take, I think, an erroneous view of the probability of closely allied species invading each other’s territory, when put into free intercommunication. Undoubtedly, if one species has any advantage over another, it will in a very brief time wholly or in part supplant it; but if both are equally well fitted for their own places, both will probably hold their separate places for almost any length of time. Being familiar with the fact that many species, naturalised through man’s agency, have spread with astonishing rapidity over wide areas, we are apt to infer that most species would thus spread; but we should remember that the species which become naturalised in new countries are not generally closely allied to the aboriginal inhabitants, but are very distinct forms, belonging in a large proportion of cases, as shown by Alph. de Candolle, to distinct genera. In the Galapagos Archipelago, many even of the birds, though so well adapted for flying from island to island, differ on the different islands; thus there are three closely allied species of mocking-thrush, each confined to its own island. Now let us suppose the mocking-thrush of Chatham Island to be blown to Charles Island, which has its own mocking-thrush; why should it succeed in establishing itself there? We may safely infer that Charles Island is well stocked with its own species, for annually more eggs are laid and young birds hatched than can possibly be reared; and we may infer that the mocking-thrush peculiar to Charles Island is at least as well fitted for its home as is the species peculiar to Chatham Island. Sir <abbr class="name">C.</abbr> Lyell and <abbr>Mr.</abbr> Wollaston have communicated to me a remarkable fact bearing on this subject; namely, that Madeira and the adjoining islet of Porto Santo possess many distinct but representative species of land-shells, some of which live in crevices of stone; and although large quantities of stone are annually transported from Porto Santo to Madeira, yet this latter island has not become colonised by the Porto Santo species: nevertheless, both islands have been colonised by some European land-shells, which no doubt had some advantage over the indigenous species. From these considerations I think we need not greatly marvel at the endemic species which inhabit the several islands of the Galapagos Archipelago not having all spread from island to island. On the same continent, also, preoccupation has probably played an important part in checking the commingling of the species which inhabit different districts with nearly the same physical conditions. Thus, the southeast and southwest corners of Australia have nearly the same physical conditions, and are united by continuous land, yet they are inhabited by a vast number of distinct mammals, birds, and plants; so it is, according to <abbr>Mr.</abbr> Bates, with the butterflies and other animals inhabiting the great, open, and continuous valley of the Amazons.</p>
 			<p>The same principle which governs the general character of the inhabitants of oceanic islands, namely, the relation to the source whence colonists could have been most easily derived, together with their subsequent modification, is of the widest application throughout nature. We see this on every mountain-summit, in every lake and marsh. For Alpine species, excepting in as far as the same species have become widely spread during the Glacial epoch, are related to those of the surrounding lowlands; thus we have in South America, Alpine hummingbirds, Alpine rodents, Alpine plants, <abbr>etc.</abbr>, all strictly belonging to American forms; and it is obvious that a mountain, as it became slowly upheaved, would be colonised from the surrounding lowlands. So it is with the inhabitants of lakes and marshes, excepting in so far as great facility of transport has allowed the same forms to prevail throughout large portions of the world. We see the same principle in the character of most of the blind animals inhabiting the caves of America and of Europe. Other analogous facts could be given. It will, I believe, be found universally true, that wherever in two regions, let them be ever so distant, many closely allied or representative species occur, there will likewise be found some identical species; and wherever many closely-allied species occur, there will be found many forms which some naturalists rank as distinct species, and others as mere varieties; these doubtful forms showing us the steps in the process of modification.</p>
 			<p>The relation between the power and extent of migration in certain species, either at the present or at some former period, and the existence at remote points of the world of closely allied species, is shown in another and more general way. <abbr>Mr.</abbr> Gould remarked to me long ago, that in those genera of birds which range over the world, many of the species have very wide ranges. I can hardly doubt that this rule is generally true, though difficult of proof. Among mammals, we see it strikingly displayed in Bats, and in a lesser degree in the Felidae and Canidae. We see the same rule in the distribution of butterflies and beetles. So it is with most of the inhabitants of fresh water, for many of the genera in the most distinct classes range over the world, and many of the species have enormous ranges. It is not meant that all, but that some of the species have very wide ranges in the genera which range very widely. Nor is it meant that the species in such genera have, on an average, a very wide range; for this will largely depend on how far the process of modification has gone; for instance, two varieties of the same species inhabit America and Europe, and thus the species has an immense range; but, if variation were to be carried a little further, the two varieties would be ranked as distinct species, and their range would be greatly reduced. Still less is it meant, that species which have the capacity of crossing barriers and ranging widely, as in the case of certain powerfully-winged birds, will necessarily range widely; for we should never forget that to range widely implies not only the power of crossing barriers, but the more important power of being victorious in distant lands in the struggle for life with foreign associates. But according to the view that all the species of a genus, though distributed to the most remote points of the world, are descended from a single progenitor, we ought to find, and I believe as a general rule we do find, that some at least of the species range very widely.</p>
 			<p>We should bear in mind that many genera in all classes are of ancient origin, and the species in this case will have had ample time for dispersal and subsequent modification. There is also reason to believe, from geological evidence, that within each great class the lower organisms change at a slower rate than the higher; consequently they will have had a better chance of ranging widely and of still retaining the same specific character. This fact, together with that of the seeds and eggs of most lowly organised forms being very minute and better fitted for distant transportal, probably accounts for a law which has long been observed, and which has lately been discussed by Alph. de Candolle in regard to plants, namely, that the lower any group of organisms stands the more widely it ranges.</p>
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 			<p>That the mere physiological importance of an organ does not determine its classificatory value, is almost proved by the fact, that in allied groups, in which the same organ, as we have every reason to suppose, has nearly the same physiological value, its classificatory value is widely different. No naturalist can have worked at any group without being struck with this fact; and it has been fully acknowledged in the writings of almost every author. It will suffice to quote the highest authority, Robert Brown, who, in speaking of certain organs in the Proteaceae, says their generic importance, “like that of all their parts, not only in this, but, as I apprehend in every natural family, is very unequal, and in some cases seems to be entirely lost.” Again, in another work he says, the genera of the Connaraceae “differ in having one or more ovaria, in the existence or absence of albumen, in the imbricate or valvular aestivation. Any one of these characters singly is frequently of more than generic importance, though here even, when all taken together, they appear insufficient to separate Cnestis from Connarus.” To give an example among insects: in one great division of the Hymenoptera, the antennae, as Westwood has remarked, are most constant in structure; in another division they differ much, and the differences are of quite subordinate value in classification; yet no one will say that the antennae in these two divisions of the same order are of unequal physiological importance. Any number of instances could be given of the varying importance for classification of the same important organ within the same group of beings.</p>
 			<p>Again, no one will say that rudimentary or atrophied organs are of high physiological or vital importance; yet, undoubtedly, organs in this condition are often of much value in classification. No one will dispute that the rudimentary teeth in the upper jaws of young ruminants, and certain rudimentary bones of the leg, are highly serviceable in exhibiting the close affinity between Ruminants and Pachyderms. Robert Brown has strongly insisted on the fact that the position of the rudimentary florets is of the highest importance in the classification of the Grasses.</p>
 			<p>Numerous instances could be given of characters derived from parts which must be considered of very trifling physiological importance, but which are universally admitted as highly serviceable in the definition of whole groups. For instance, whether or not there is an open passage from the nostrils to the mouth, the only character, according to Owen, which absolutely distinguishes fishes and reptiles⁠—the inflection of the angle of the lower jaw in Marsupials⁠—the manner in which the wings of insects are folded⁠—mere colour in certain Algae⁠—mere pubescence on parts of the flower in grasses⁠—the nature of the dermal covering, as hair or feathers, in the Vertebrata. If the Ornithorhynchus had been covered with feathers instead of hair, this external and trifling character would have been considered by naturalists as an important aid in determining the degree of affinity of this strange creature to birds.</p>
-			<p>The importance, for classification, of trifling characters, mainly depends on their being correlated with many other characters of more or less importance. The value indeed of an aggregate of characters is very evident in natural history. Hence, as has often been remarked, a species may depart from its allies in several characters, both of high physiological importance, and of almost universal prevalence, and yet leave us in no doubt where it should be ranked. Hence, also, it has been found that a classification founded on any single character, however important that may be, has always failed; for no part of the organisation is invariably constant. The importance of an aggregate of characters, even when none are important, alone explains the aphorism enunciated by Linnaeus, namely, that the characters do not give the genus, but the genus gives the character; for this seems founded on the appreciation of many trifling points of resemblance, too slight to be defined. Certain plants, belonging to the Malpighiaceae, bear perfect and degraded flowers; in the latter, as A. de Jussieu has remarked, “The greater number of the characters proper to the species, to the genus, to the family, to the class, disappear, and thus laugh at our classification.” When Aspicarpa produced in France, during several years, only these degraded flowers, departing so wonderfully in a number of the most important points of structure from the proper type of the order, yet M. Richard sagaciously saw, as Jussieu observes, that this genus should still be retained among the Malpighiaceae. This case well illustrates the spirit of our classifications.</p>
+			<p>The importance, for classification, of trifling characters, mainly depends on their being correlated with many other characters of more or less importance. The value indeed of an aggregate of characters is very evident in natural history. Hence, as has often been remarked, a species may depart from its allies in several characters, both of high physiological importance, and of almost universal prevalence, and yet leave us in no doubt where it should be ranked. Hence, also, it has been found that a classification founded on any single character, however important that may be, has always failed; for no part of the organisation is invariably constant. The importance of an aggregate of characters, even when none are important, alone explains the aphorism enunciated by Linnaeus, namely, that the characters do not give the genus, but the genus gives the character; for this seems founded on the appreciation of many trifling points of resemblance, too slight to be defined. Certain plants, belonging to the Malpighiaceae, bear perfect and degraded flowers; in the latter, as <abbr class="name">A.</abbr> de Jussieu has remarked, “The greater number of the characters proper to the species, to the genus, to the family, to the class, disappear, and thus laugh at our classification.” When Aspicarpa produced in France, during several years, only these degraded flowers, departing so wonderfully in a number of the most important points of structure from the proper type of the order, yet <abbr class="name">M.</abbr> Richard sagaciously saw, as Jussieu observes, that this genus should still be retained among the Malpighiaceae. This case well illustrates the spirit of our classifications.</p>
 			<p>Practically, when naturalists are at work, they do not trouble themselves about the physiological value of the characters which they use in defining a group or in allocating any particular species. If they find a character nearly uniform, and common to a great number of forms, and not common to others, they use it as one of high value; if common to some lesser number, they use it as of subordinate value. This principle has been broadly confessed by some naturalists to be the true one; and by none more clearly than by that excellent botanist, <abbr class="name">Aug. St.</abbr> Hilaire. If several trifling characters are always found in combination, though no apparent bond of connection can be discovered between them, especial value is set on them. As in most groups of animals, important organs, such as those for propelling the blood, or for aerating it, or those for propagating the race, are found nearly uniform, they are considered as highly serviceable in classification; but in some groups all these, the most important vital organs, are found to offer characters of quite subordinate value. Thus, as Fritz Muller has lately remarked, in the same group of crustaceans, Cypridina is furnished with a heart, while in two closely allied genera, namely Cypris and Cytherea, there is no such organ; one species of Cypridina has well-developed branchiae, while another species is destitute of them.</p>
 			<p>We can see why characters derived from the embryo should be of equal importance with those derived from the adult, for a natural classification of course includes all ages. But it is by no means obvious, on the ordinary view, why the structure of the embryo should be more important for this purpose than that of the adult, which alone plays its full part in the economy of nature. Yet it has been strongly urged by those great naturalists, Milne Edwards and Agassiz, that embryological characters are the most important of all; and this doctrine has very generally been admitted as true. Nevertheless, their importance has sometimes been exaggerated, owing to the adaptive characters of larvae not having been excluded; in order to show this, Fritz Muller arranged, by the aid of such characters alone, the great class of crustaceans, and the arrangement did not prove a natural one. But there can be no doubt that embryonic, excluding larval characters, are of the highest value for classification, not only with animals but with plants. Thus the main divisions of flowering plants are founded on differences in the embryo⁠—on the number and position of the cotyledons, and on the mode of development of the plumule and radicle. We shall immediately see why these characters possess so high a value in classification, namely, from the natural system being genealogical in its arrangement.</p>
 			<p>Our classifications are often plainly influenced by chains of affinities. Nothing can be easier than to define a number of characters common to all birds; but with crustaceans, any such definition has hitherto been found impossible. There are crustaceans at the opposite ends of the series, which have hardly a character in common; yet the species at both ends, from being plainly allied to others, and these to others, and so onwards, can be recognised as unequivocally belonging to this, and to no other class of the Articulata.</p>
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 			<h3 epub:type="title">Development and Embryology</h3>
-			<p>This is one of the most important subjects in the whole round of natural history. The metamorphoses of insects, with which everyone is familiar, are generally effected abruptly by a few stages; but the transformations are in reality numerous and gradual, though concealed. A certain ephemerous insect (Chloeon) during its development, moults, as shown by Sir J. Lubbock, above twenty times, and each time undergoes a certain amount of change; and in this case we see the act of metamorphosis performed in a primary and gradual manner. Many insects, and especially certain crustaceans, show us what wonderful changes of structure can be effected during development. Such changes, however, reach their acme in the so-called alternate generations of some of the lower animals. It is, for instance, an astonishing fact that a delicate branching coralline, studded with polypi, and attached to a submarine rock, should produce, first by budding and then by transverse division, a host of huge floating jellyfishes; and that these should produce eggs, from which are hatched swimming animalcules, which attach themselves to rocks and become developed into branching corallines; and so on in an endless cycle. The belief in the essential identity of the process of alternate generation and of ordinary metamorphosis has been greatly strengthened by Wagner’s discovery of the larva or maggot of a fly, namely the Cecidomyia, producing asexually other larvae, and these others, which finally are developed into mature males and females, propagating their kind in the ordinary manner by eggs.</p>
+			<p>This is one of the most important subjects in the whole round of natural history. The metamorphoses of insects, with which everyone is familiar, are generally effected abruptly by a few stages; but the transformations are in reality numerous and gradual, though concealed. A certain ephemerous insect (Chloeon) during its development, moults, as shown by Sir <abbr class="name">J.</abbr> Lubbock, above twenty times, and each time undergoes a certain amount of change; and in this case we see the act of metamorphosis performed in a primary and gradual manner. Many insects, and especially certain crustaceans, show us what wonderful changes of structure can be effected during development. Such changes, however, reach their acme in the so-called alternate generations of some of the lower animals. It is, for instance, an astonishing fact that a delicate branching coralline, studded with polypi, and attached to a submarine rock, should produce, first by budding and then by transverse division, a host of huge floating jellyfishes; and that these should produce eggs, from which are hatched swimming animalcules, which attach themselves to rocks and become developed into branching corallines; and so on in an endless cycle. The belief in the essential identity of the process of alternate generation and of ordinary metamorphosis has been greatly strengthened by Wagner’s discovery of the larva or maggot of a fly, namely the Cecidomyia, producing asexually other larvae, and these others, which finally are developed into mature males and females, propagating their kind in the ordinary manner by eggs.</p>
 			<p>It may be worth notice that when Wagner’s remarkable discovery was first announced, I was asked how was it possible to account for the larvae of this fly having acquired the power of a sexual reproduction. As long as the case remained unique no answer could be given. But already Grimm has shown that another fly, a Chironomus, reproduces itself in nearly the same manner, and he believes that this occurs frequently in the order. It is the pupa, and not the larva, of the Chironomus which has this power; and Grimm further shows that this case, to a certain extent, “unites that of the Cecidomyia with the parthenogenesis of the Coccidae;” the term parthenogenesis implying that the mature females of the Coccidae are capable of producing fertile eggs without the concourse of the male. Certain animals belonging to several classes are now known to have the power of ordinary reproduction at an unusually early age; and we have only to accelerate parthenogenetic reproduction by gradual steps to an earlier and earlier age⁠—Chironomus showing us an almost exactly intermediate stage, <abbr>viz.</abbr>, that of the pupa⁠—and we can perhaps account for the marvellous case of the Cecidomyia.</p>
 			<p>It has already been stated that various parts in the same individual, which are exactly alike during an early embryonic period, become widely different and serve for widely different purposes in the adult state. So again it has been shown that generally the embryos of the most distinct species belonging to the same class are closely similar, but become, when fully developed, widely dissimilar. A better proof of this latter fact cannot be given than the statement by Von Baer that “the embryos of mammalia, of birds, lizards and snakes, probably also of chelonia, are in the earliest states exceedingly like one another, both as a whole and in the mode of development of their parts; so much so, in fact, that we can often distinguish the embryos only by their size. In my possession are two little embryos in spirit, whose names I have omitted to attach, and at present I am quite unable to say to what class they belong. They may be lizards or small birds, or very young mammalia, so complete is the similarity in the mode of formation of the head and trunk in these animals. The extremities, however, are still absent in these embryos. But even if they had existed in the earliest stage of their development we should learn nothing, for the feet of lizards and mammals, the wings and feet of birds, no less than the hands and feet of man, all arise from the same fundamental form.” The larvae of most crustaceans, at corresponding stages of development, closely resemble each other, however different the adults may become; and so it is with very many other animals. A trace of the law of embryonic resemblance occasionally lasts till a rather late age: thus birds of the same genus, and of allied genera, often resemble each other in their immature plumage; as we see in the spotted feathers in the young of the thrush group. In the cat tribe, most of the species when adult are striped or spotted in lines; and stripes or spots can be plainly distinguished in the whelp of the lion and the puma. We occasionally, though rarely, see something of the same kind in plants; thus the first leaves of the ulex or furze, and the first leaves of the phyllodineous acacias, are pinnate or divided like the ordinary leaves of the leguminosae.</p>
 			<p>The points of structure, in which the embryos of widely different animals within the same class resemble each other, often have no direct relation to their conditions of existence. We cannot, for instance, suppose that in the embryos of the vertebrata the peculiar loop-like courses of the arteries near the branchial slits are related to similar conditions⁠—in the young mammal which is nourished in the womb of its mother, in the egg of the bird which is hatched in a nest, and in the spawn of a frog under water. We have no more reason to believe in such a relation than we have to believe that the similar bones in the hand of a man, wing of a bat, and fin of a porpoise, are related to similar conditions of life. No one supposes that the stripes on the whelp of a lion, or the spots on the young blackbird, are of any use to these animals.</p>
-			<p>The case, however, is different when an animal, during any part of its embryonic career, is active, and has to provide for itself. The period of activity may come on earlier or later in life; but whenever it comes on, the adaptation of the larva to its conditions of life is just as perfect and as beautiful as in the adult animal. In how important a manner this has acted, has recently been well shown by Sir J. Lubbock in his remarks on the close similarity of the larvae of some insects belonging to very different orders, and on the dissimilarity of the larvae of other insects within the same order, according to their habits of life. Owing to such adaptations the similarity of the larvae of allied animals is sometimes greatly obscured; especially when there is a division of labour during the different stages of development, as when the same larva has during one stage to search for food, and during another stage has to search for a place of attachment. Cases can even be given of the larvae of allied species, or groups of species, differing more from each other than do the adults. In most cases, however, the larvae, though active, still obey, more or less closely, the law of common embryonic resemblance. Cirripedes afford a good instance of this: even the illustrious Cuvier did not perceive that a barnacle was a crustacean: but a glance at the larva shows this in an unmistakable manner. So again the two main divisions of cirripedes, the pedunculated and sessile, though differing widely in external appearance, have larvae in all their stages barely distinguishable.</p>
+			<p>The case, however, is different when an animal, during any part of its embryonic career, is active, and has to provide for itself. The period of activity may come on earlier or later in life; but whenever it comes on, the adaptation of the larva to its conditions of life is just as perfect and as beautiful as in the adult animal. In how important a manner this has acted, has recently been well shown by Sir <abbr class="name">J.</abbr> Lubbock in his remarks on the close similarity of the larvae of some insects belonging to very different orders, and on the dissimilarity of the larvae of other insects within the same order, according to their habits of life. Owing to such adaptations the similarity of the larvae of allied animals is sometimes greatly obscured; especially when there is a division of labour during the different stages of development, as when the same larva has during one stage to search for food, and during another stage has to search for a place of attachment. Cases can even be given of the larvae of allied species, or groups of species, differing more from each other than do the adults. In most cases, however, the larvae, though active, still obey, more or less closely, the law of common embryonic resemblance. Cirripedes afford a good instance of this: even the illustrious Cuvier did not perceive that a barnacle was a crustacean: but a glance at the larva shows this in an unmistakable manner. So again the two main divisions of cirripedes, the pedunculated and sessile, though differing widely in external appearance, have larvae in all their stages barely distinguishable.</p>
 			<p>The embryo in the course of development generally rises in organisation. I use this expression, though I am aware that it is hardly possible to define clearly what is meant by organisation being higher or lower. But no one probably will dispute that the butterfly is higher than the caterpillar. In some cases, however, the mature animal must be considered as lower in the scale than the larva, as with certain parasitic crustaceans. To refer once again to cirripedes: the larvae in the first stage have three pairs of locomotive organs, a simple single eye, and a probosciformed mouth, with which they feed largely, for they increase much in size. In the second stage, answering to the chrysalis stage of butterflies, they have six pairs of beautifully constructed natatory legs, a pair of magnificent compound eyes, and extremely complex antennae; but they have a closed and imperfect mouth, and cannot feed: their function at this stage is, to search out by their well-developed organs of sense, and to reach by their active powers of swimming, a proper place on which to become attached and to undergo their final metamorphosis. When this is completed they are fixed for life: their legs are now converted into prehensile organs; they again obtain a well-constructed mouth; but they have no antennae, and their two eyes are now reconverted into a minute, single, simple eyespot. In this last and complete state, cirripedes may be considered as either more highly or more lowly organised than they were in the larval condition. But in some genera the larvae become developed into hermaphrodites having the ordinary structure, or into what I have called complemental males; and in the latter the development has assuredly been retrograde; for the male is a mere sack, which lives for a short time and is destitute of mouth, stomach, and every other organ of importance, excepting those for reproduction.</p>
 			<p>We are so much accustomed to see a difference in structure between the embryo and the adult, that we are tempted to look at this difference as in some necessary manner contingent on growth. But there is no reason why, for instance, the wing of a bat, or the fin of a porpoise, should not have been sketched out with all their parts in proper proportion, as soon as any part became visible. In some whole groups of animals and in certain members of other groups this is the case, and the embryo does not at any period differ widely from the adult: thus Owen has remarked in regard to cuttlefish, “there is no metamorphosis; the cephalopodic character is manifested long before the parts of the embryo are completed.” Land-shells and freshwater crustaceans are born having their proper forms, while the marine members of the same two great classes pass through considerable and often great changes during their development. Spiders, again, barely undergo any metamorphosis. The larvae of most insects pass through a worm-like stage, whether they are active and adapted to diversified habits, or are inactive from being placed in the midst of proper nutriment, or from being fed by their parents; but in some few cases, as in that of Aphis, if we look to the admirable drawings of the development of this insect, by Professor Huxley, we see hardly any trace of the vermiform stage.</p>
 			<p>Sometimes it is only the earlier developmental stages which fail. Thus, Fritz Muller has made the remarkable discovery that certain shrimp-like crustaceans (allied to Penoeus) first appear under the simple nauplius-form, and after passing through two or more zoea-stages, and then through the mysis-stage, finally acquire their mature structure: now in the whole great malacostracan order, to which these crustaceans belong, no other member is as yet known to be first developed under the nauplius-form, though many appear as zoeas; nevertheless Muller assigns reasons for his belief, that if there had been no suppression of development, all these crustaceans would have appeared as nauplii.</p>
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 			<p>Now, let us apply these two principles to species in a state of nature. Let us take a group of birds, descended from some ancient form and modified through natural selection for different habits. Then, from the many slight successive variations having supervened in the several species at a not early age, and having been inherited at a corresponding age, the young will have been but little modified, and they will still resemble each other much more closely than do the adults, just as we have seen with the breeds of the pigeon. We may extend this view to widely distinct structures and to whole classes. The forelimbs, for instance, which once served as legs to a remote progenitor, may have become, through a long course of modification, adapted in one descendant to act as hands, in another as paddles, in another as wings; but on the above two principles the forelimbs will not have been much modified in the embryos of these several forms; although in each form the forelimb will differ greatly in the adult state. Whatever influence long continued use or disuse may have had in modifying the limbs or other parts of any species, this will chiefly or solely have affected it when nearly mature, when it was compelled to use its full powers to gain its own living; and the effects thus produced will have been transmitted to the offspring at a corresponding nearly mature age. Thus the young will not be modified, or will be modified only in a slight degree, through the effects of the increased use or disuse of parts.</p>
 			<p>With some animals the successive variations may have supervened at a very early period of life, or the steps may have been inherited at an earlier age than that at which they first occurred. In either of these cases the young or embryo will closely resemble the mature parent-form, as we have seen with the short-faced tumbler. And this is the rule of development in certain whole groups, or in certain subgroups alone, as with cuttlefish, land-shells, freshwater crustaceans, spiders, and some members of the great class of insects. With respect to the final cause of the young in such groups not passing through any metamorphosis, we can see that this would follow from the following contingencies: namely, from the young having to provide at a very early age for their own wants, and from their following the same habits of life with their parents; for in this case it would be indispensable for their existence that they should be modified in the same manner as their parents. Again, with respect to the singular fact that many terrestrial and freshwater animals do not undergo any metamorphosis, while marine members of the same groups pass through various transformations, Fritz Muller has suggested that the process of slowly modifying and adapting an animal to live on the land or in fresh water, instead of in the sea, would be greatly simplified by its not passing through any larval stage; for it is not probable that places well adapted for both the larval and mature stages, under such new and greatly changed habits of life, would commonly be found unoccupied or ill-occupied by other organisms. In this case the gradual acquirement at an earlier and earlier age of the adult structure would be favoured by natural selection; and all traces of former metamorphoses would finally be lost.</p>
 			<p>If, on the other hand, it profited the young of an animal to follow habits of life slightly different from those of the parent-form, and consequently to be constructed on a slightly different plan, or if it profited a larva already different from its parent to change still further, then, on the principle of inheritance at corresponding ages, the young or the larvae might be rendered by natural selection more and more different from their parents to any conceivable extent. Differences in the larva might, also, become correlated with successive stages of its development; so that the larva, in the first stage, might come to differ greatly from the larva in the second stage, as is the case with many animals. The adult might also become fitted for sites or habits, in which organs of locomotion or of the senses, <abbr>etc.</abbr>, would be useless; and in this case the metamorphosis would be retrograde.</p>
-			<p>From the remarks just made we can see how by changes of structure in the young, in conformity with changed habits of life, together with inheritance at corresponding ages, animals might come to pass through stages of development, perfectly distinct from the primordial condition of their adult progenitors. Most of our best authorities are now convinced that the various larval and pupal stages of insects have thus been acquired through adaptation, and not through inheritance from some ancient form. The curious case of Sitaris⁠—a beetle which passes through certain unusual stages of development⁠—will illustrate how this might occur. The first larval form is described by M. Fabre, as an active, minute insect, furnished with six legs, two long antennae, and four eyes. These larvae are hatched in the nests of bees; and when the male bees emerge from their burrows, in the spring, which they do before the females, the larvae spring on them, and afterwards crawl on to the females while paired with the males. As soon as the female bee deposits her eggs on the surface of the honey stored in the cells, the larvae of the Sitaris leap on the eggs and devour them. Afterwards they undergo a complete change; their eyes disappear; their legs and antennae become rudimentary, and they feed on honey; so that they now more closely resemble the ordinary larvae of insects; ultimately they undergo a further transformation, and finally emerge as the perfect beetle. Now, if an insect, undergoing transformations like those of the Sitaris, were to become the progenitor of a whole new class of insects, the course of development of the new class would be widely different from that of our existing insects; and the first larval stage certainly would not represent the former condition of any adult and ancient form.</p>
+			<p>From the remarks just made we can see how by changes of structure in the young, in conformity with changed habits of life, together with inheritance at corresponding ages, animals might come to pass through stages of development, perfectly distinct from the primordial condition of their adult progenitors. Most of our best authorities are now convinced that the various larval and pupal stages of insects have thus been acquired through adaptation, and not through inheritance from some ancient form. The curious case of Sitaris⁠—a beetle which passes through certain unusual stages of development⁠—will illustrate how this might occur. The first larval form is described by <abbr class="name">M.</abbr> Fabre, as an active, minute insect, furnished with six legs, two long antennae, and four eyes. These larvae are hatched in the nests of bees; and when the male bees emerge from their burrows, in the spring, which they do before the females, the larvae spring on them, and afterwards crawl on to the females while paired with the males. As soon as the female bee deposits her eggs on the surface of the honey stored in the cells, the larvae of the Sitaris leap on the eggs and devour them. Afterwards they undergo a complete change; their eyes disappear; their legs and antennae become rudimentary, and they feed on honey; so that they now more closely resemble the ordinary larvae of insects; ultimately they undergo a further transformation, and finally emerge as the perfect beetle. Now, if an insect, undergoing transformations like those of the Sitaris, were to become the progenitor of a whole new class of insects, the course of development of the new class would be widely different from that of our existing insects; and the first larval stage certainly would not represent the former condition of any adult and ancient form.</p>
 			<p>On the other hand it is highly probable that with many animals the embryonic or larval stages show us, more or less completely, the condition of the progenitor of the whole group in its adult state. In the great class of the Crustacea, forms wonderfully distinct from each other, namely, suctorial parasites, cirripedes, entomostraca, and even the malacostraca, appear at first as larvae under the nauplius-form; and as these larvae live and feed in the open sea, and are not adapted for any peculiar habits of life, and from other reasons assigned by Fritz Muller, it is probable that at some very remote period an independent adult animal, resembling the Nauplius, existed, and subsequently produced, along several divergent lines of descent, the above-named great Crustacean groups. So again, it is probable, from what we know of the embryos of mammals, birds, fishes and reptiles, that these animals are the modified descendants of some ancient progenitor, which was furnished in its adult state with branchiae, a swim-bladder, four fin-like limbs, and a long tail, all fitted for an aquatic life.</p>
 			<p>As all the organic beings, extinct and recent, which have ever lived, can be arranged within a few great classes; and as all within each class have, according to our theory, been connected together by fine gradations, the best, and, if our collections were nearly perfect, the only possible arrangement, would be genealogical; descent being the hidden bond of connection which naturalists have been seeking under the term of the Natural System. On this view we can understand how it is that, in the eyes of most naturalists, the structure of the embryo is even more important for classification than that of the adult. In two or more groups of animals, however much they may differ from each other in structure and habits in their adult condition, if they pass through closely similar embryonic stages, we may feel assured that they are all descended from one parent-form, and are therefore closely related. Thus, community in embryonic structure reveals community of descent; but dissimilarity in embryonic development does not prove discommunity of descent, for in one of two groups the developmental stages may have been suppressed, or may have been so greatly modified through adaptation to new habits of life as to be no longer recognisable. Even in groups, in which the adults have been modified to an extreme degree, community of origin is often revealed by the structure of the larvae; we have seen, for instance, that cirripedes, though externally so like shellfish, are at once known by their larvae to belong to the great class of crustaceans. As the embryo often shows us more or less plainly the structure of the less modified and ancient progenitor of the group, we can see why ancient and extinct forms so often resemble in their adult state the embryos of existing species of the same class. Agassiz believes this to be a universal law of nature; and we may hope hereafter to see the law proved true. It can, however, be proved true only in those cases in which the ancient state of the progenitor of the group has not been wholly obliterated, either by successive variations having supervened at a very early period of growth, or by such variations having been inherited at an earlier age than that at which they first appeared. It should also be borne in mind, that the law may be true, but yet, owing to the geological record not extending far enough back in time, may remain for a long period, or forever, incapable of demonstration. The law will not strictly hold good in those cases in which an ancient form became adapted in its larval state to some special line of life, and transmitted the same larval state to a whole group of descendants; for such larval state will not resemble any still more ancient form in its adult state.</p>
 			<p>Thus, as it seems to me, the leading facts in embryology, which are second to none in importance, are explained on the principle of variations in the many descendants from some one ancient progenitor, having appeared at a not very early period of life, and having been inherited at a corresponding period. Embryology rises greatly in interest, when we look at the embryo as a picture, more or less obscured, of the progenitor, either in its adult or larval state, of all the members of the same great class.</p>

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 			<h3 epub:type="title">Individual Differences</h3>
-			<p>The many slight differences which appear in the offspring from the same parents, or which it may be presumed have thus arisen, from being observed in the individuals of the same species inhabiting the same confined locality, may be called individual differences. No one supposes that all the individuals of the same species are cast in the same actual mould. These individual differences are of the highest importance for us, for they are often inherited, as must be familiar to everyone; and they thus afford materials for natural selection to act on and accumulate, in the same manner as man accumulates in any given direction individual differences in his domesticated productions. These individual differences generally affect what naturalists consider unimportant parts; but I could show, by a long catalogue of facts, that parts which must be called important, whether viewed under a physiological or classificatory point of view, sometimes vary in the individuals of the same species. I am convinced that the most experienced naturalist would be surprised at the number of the cases of variability, even in important parts of structure, which he could collect on good authority, as I have collected, during a course of years. It should be remembered that systematists are far from being pleased at finding variability in important characters, and that there are not many men who will laboriously examine internal and important organs, and compare them in many specimens of the same species. It would never have been expected that the branching of the main nerves close to the great central ganglion of an insect would have been variable in the same species; it might have been thought that changes of this nature could have been effected only by slow degrees; yet Sir J. Lubbock has shown a degree of variability in these main nerves in Coccus, which may almost be compared to the irregular branching of the stem of a tree. This philosophical naturalist, I may add, has also shown that the muscles in the larvae of certain insects are far from uniform. Authors sometimes argue in a circle when they state that important organs never vary; for these same authors practically rank those parts as important (as some few naturalists have honestly confessed) which do not vary; and, under this point of view, no instance will ever be found of an important part varying; but under any other point of view many instances assuredly can be given.</p>
+			<p>The many slight differences which appear in the offspring from the same parents, or which it may be presumed have thus arisen, from being observed in the individuals of the same species inhabiting the same confined locality, may be called individual differences. No one supposes that all the individuals of the same species are cast in the same actual mould. These individual differences are of the highest importance for us, for they are often inherited, as must be familiar to everyone; and they thus afford materials for natural selection to act on and accumulate, in the same manner as man accumulates in any given direction individual differences in his domesticated productions. These individual differences generally affect what naturalists consider unimportant parts; but I could show, by a long catalogue of facts, that parts which must be called important, whether viewed under a physiological or classificatory point of view, sometimes vary in the individuals of the same species. I am convinced that the most experienced naturalist would be surprised at the number of the cases of variability, even in important parts of structure, which he could collect on good authority, as I have collected, during a course of years. It should be remembered that systematists are far from being pleased at finding variability in important characters, and that there are not many men who will laboriously examine internal and important organs, and compare them in many specimens of the same species. It would never have been expected that the branching of the main nerves close to the great central ganglion of an insect would have been variable in the same species; it might have been thought that changes of this nature could have been effected only by slow degrees; yet Sir <abbr class="name">J.</abbr> Lubbock has shown a degree of variability in these main nerves in Coccus, which may almost be compared to the irregular branching of the stem of a tree. This philosophical naturalist, I may add, has also shown that the muscles in the larvae of certain insects are far from uniform. Authors sometimes argue in a circle when they state that important organs never vary; for these same authors practically rank those parts as important (as some few naturalists have honestly confessed) which do not vary; and, under this point of view, no instance will ever be found of an important part varying; but under any other point of view many instances assuredly can be given.</p>
 			<p>There is one point connected with individual differences which is extremely perplexing: I refer to those genera which have been called “protean” or “polymorphic,” in which species present an inordinate amount of variation. With respect to many of these forms, hardly two naturalists agree whether to rank them as species or as varieties. We may instance Rubus, Rosa, and Hieracium among plants, several genera of insects, and of Brachiopod shells. In most polymorphic genera some of the species have fixed and definite characters. Genera which are polymorphic in one country seem to be, with a few exceptions, polymorphic in other countries, and likewise, judging from Brachiopod shells, at former periods of time. These facts are very perplexing, for they seem to show that this kind of variability is independent of the conditions of life. I am inclined to suspect that we see, at least in some of these polymorphic genera, variations which are of no service or disservice to the species, and which consequently have not been seized on and rendered definite by natural selection, as hereafter to be explained.</p>
 			<p>Individuals of the same species often present, as is known to everyone, great differences of structure, independently of variation, as in the two sexes of various animals, in the two or three castes of sterile females or workers among insects, and in the immature and larval states of many of the lower animals. There are, also, cases of dimorphism and trimorphism, both with animals and plants. Thus, <abbr>Mr.</abbr> Wallace, who has lately called attention to the subject, has shown that the females of certain species of butterflies, in the Malayan Archipelago, regularly appear under two or even three conspicuously distinct forms, not connected by intermediate varieties. Fritz Muller has described analogous but more extraordinary cases with the males of certain Brazilian Crustaceans: thus, the male of a Tanais regularly occurs under two distinct forms; one of these has strong and differently shaped pincers, and the other has antennae much more abundantly furnished with smelling-hairs. Although in most of these cases, the two or three forms, both with animals and plants, are not now connected by intermediate gradations, it is possible that they were once thus connected. <abbr>Mr.</abbr> Wallace, for instance, describes a certain butterfly which presents in the same island a great range of varieties connected by intermediate links, and the extreme links of the chain closely resemble the two forms of an allied dimorphic species inhabiting another part of the Malay Archipelago. Thus also with ants, the several worker-castes are generally quite distinct; but in some cases, as we shall hereafter see, the castes are connected together by finely graduated varieties. So it is, as I have myself observed, with some dimorphic plants. It certainly at first appears a highly remarkable fact that the same female butterfly should have the power of producing at the same time three distinct female forms and a male; and that an hermaphrodite plant should produce from the same seed-capsule three distinct hermaphrodite forms, bearing three different kinds of females and three or even six different kinds of males. Nevertheless these cases are only exaggerations of the common fact that the female produces offspring of two sexes which sometimes differ from each other in a wonderful manner.</p>
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 			<p><abbr>Mr.</abbr> <abbr class="name">B. D.</abbr> Walsh, a distinguished entomologist of the United States, has described what he calls Phytophagic varieties and Phytophagic species. Most vegetable-feeding insects live on one kind of plant or on one group of plants; some feed indiscriminately on many kinds, but do not in consequence vary. In several cases, however, insects found living on different plants, have been observed by <abbr>Mr.</abbr> Walsh to present in their larval or mature state, or in both states, slight, though constant differences in colour, size, or in the nature of their secretions. In some instances the males alone, in other instances, both males and females, have been observed thus to differ in a slight degree. When the differences are rather more strongly marked, and when both sexes and all ages are affected, the forms are ranked by all entomologists as good species. But no observer can determine for another, even if he can do so for himself, which of these Phytophagic forms ought to be called species and which varieties. <abbr>Mr.</abbr> Walsh ranks the forms which it may be supposed would freely intercross, as varieties; and those which appear to have lost this power, as species. As the differences depend on the insects having long fed on distinct plants, it cannot be expected that intermediate links connecting the several forms should now be found. The naturalist thus loses his best guide in determining whether to rank doubtful forms as varieties or species. This likewise necessarily occurs with closely allied organisms, which inhabit distinct continents or islands. When, on the other hand, an animal or plant ranges over the same continent, or inhabits many islands in the same archipelago, and presents different forms in the different areas, there is always a good chance that intermediate forms will be discovered which will link together the extreme states; and these are then degraded to the rank of varieties.</p>
 			<p>Some few naturalists maintain that animals never present varieties; but then these same naturalists rank the slightest difference as of specific value; and when the same identical form is met with in two distant countries, or in two geological formations, they believe that two distinct species are hidden under the same dress. The term species thus comes to be a mere useless abstraction, implying and assuming a separate act of creation. It is certain that many forms, considered by highly competent judges to be varieties, resemble species so completely in character that they have been thus ranked by other highly competent judges. But to discuss whether they ought to be called species or varieties, before any definition of these terms has been generally accepted, is vainly to beat the air.</p>
 			<p>Many of the cases of strongly marked varieties or doubtful species well deserve consideration; for several interesting lines of argument, from geographical distribution, analogical variation, hybridism, <abbr>etc.</abbr>, have been brought to bear in the attempt to determine their rank; but space does not here permit me to discuss them. Close investigation, in many cases, will no doubt bring naturalists to agree how to rank doubtful forms. Yet it must be confessed that it is in the best known countries that we find the greatest number of them. I have been struck with the fact that if any animal or plant in a state of nature be highly useful to man, or from any cause closely attracts his attention, varieties of it will almost universally be found recorded. These varieties, moreover, will often be ranked by some authors as species. Look at the common oak, how closely it has been studied; yet a German author makes more than a dozen species out of forms, which are almost universally considered by other botanists to be varieties; and in this country the highest botanical authorities and practical men can be quoted to show that the sessile and pedunculated oaks are either good and distinct species or mere varieties.</p>
-			<p>I may here allude to a remarkable memoir lately published by A. de Candolle, on the oaks of the whole world. No one ever had more ample materials for the discrimination of the species, or could have worked on them with more zeal and sagacity. He first gives in detail all the many points of structure which vary in the several species, and estimates numerically the relative frequency of the variations. He specifies above a dozen characters which may be found varying even on the same branch, sometimes according to age or development, sometimes without any assignable reason. Such characters are not of course of specific value, but they are, as Asa Gray has remarked in commenting on this memoir, such as generally enter into specific definitions. De Candolle then goes on to say that he gives the rank of species to the forms that differ by characters never varying on the same tree, and never found connected by intermediate states. After this discussion, the result of so much labour, he emphatically remarks: “They are mistaken, who repeat that the greater part of our species are clearly limited, and that the doubtful species are in a feeble minority. This seemed to be true, so long as a genus was imperfectly known, and its species were founded upon a few specimens, that is to say, were provisional. Just as we come to know them better, intermediate forms flow in, and doubts as to specific limits augment.” He also adds that it is the best known species which present the greatest number of spontaneous varieties and sub-varieties. Thus Quercus robur has twenty-eight varieties, all of which, excepting six, are clustered round three subspecies, namely Q. pedunculata, sessiliflora and pubescens. The forms which connect these three subspecies are comparatively rare; and, as Asa Gray again remarks, if these connecting forms which are now rare were to become totally extinct the three subspecies would hold exactly the same relation to each other as do the four or five provisionally admitted species which closely surround the typical Quercus robur. Finally, De Candolle admits that out of the 300 species, which will be enumerated in his Prodromus as belonging to the oak family, at least two-thirds are provisional species, that is, are not known strictly to fulfil the definition above given of a true species. It should be added that De Candolle no longer believes that species are immutable creations, but concludes that the derivative theory is the most natural one, “and the most accordant with the known facts in palaeontology, geographical botany and zoology, of anatomical structure and classification.”</p>
+			<p>I may here allude to a remarkable memoir lately published by <abbr class="name">A.</abbr> de Candolle, on the oaks of the whole world. No one ever had more ample materials for the discrimination of the species, or could have worked on them with more zeal and sagacity. He first gives in detail all the many points of structure which vary in the several species, and estimates numerically the relative frequency of the variations. He specifies above a dozen characters which may be found varying even on the same branch, sometimes according to age or development, sometimes without any assignable reason. Such characters are not of course of specific value, but they are, as Asa Gray has remarked in commenting on this memoir, such as generally enter into specific definitions. De Candolle then goes on to say that he gives the rank of species to the forms that differ by characters never varying on the same tree, and never found connected by intermediate states. After this discussion, the result of so much labour, he emphatically remarks: “They are mistaken, who repeat that the greater part of our species are clearly limited, and that the doubtful species are in a feeble minority. This seemed to be true, so long as a genus was imperfectly known, and its species were founded upon a few specimens, that is to say, were provisional. Just as we come to know them better, intermediate forms flow in, and doubts as to specific limits augment.” He also adds that it is the best known species which present the greatest number of spontaneous varieties and sub-varieties. Thus Quercus robur has twenty-eight varieties, all of which, excepting six, are clustered round three subspecies, namely <i epub:type="z3998:taxonomy">Q. pedunculata</i>, <i epub:type="z3998:taxonomy">sessiliflora</i> and <i epub:type="z3998:taxonomy">pubescens</i>. The forms which connect these three subspecies are comparatively rare; and, as Asa Gray again remarks, if these connecting forms which are now rare were to become totally extinct the three subspecies would hold exactly the same relation to each other as do the four or five provisionally admitted species which closely surround the typical Quercus robur. Finally, De Candolle admits that out of the 300 species, which will be enumerated in his Prodromus as belonging to the oak family, at least two-thirds are provisional species, that is, are not known strictly to fulfil the definition above given of a true species. It should be added that De Candolle no longer believes that species are immutable creations, but concludes that the derivative theory is the most natural one, “and the most accordant with the known facts in palaeontology, geographical botany and zoology, of anatomical structure and classification.”</p>
 			<p>When a young naturalist commences the study of a group of organisms quite unknown to him he is at first much perplexed in determining what differences to consider as specific and what as varietal; for he knows nothing of the amount and kind of variation to which the group is subject; and this shows, at least, how very generally there is some variation. But if he confine his attention to one class within one country he will soon make up his mind how to rank most of the doubtful forms. His general tendency will be to make many species, for he will become impressed, just like the pigeon or poultry fancier before alluded to, with the amount of difference in the forms which he is continually studying; and he has little general knowledge of analogical variation in other groups and in other countries by which to correct his first impressions. As he extends the range of his observations he will meet with more cases of difficulty; for he will encounter a greater number of closely-allied forms. But if his observations be widely extended he will in the end generally be able to make up his own mind; but he will succeed in this at the expense of admitting much variation, and the truth of this admission will often be disputed by other naturalists. When he comes to study allied forms brought from countries not now continuous, in which case he cannot hope to find intermediate links, he will be compelled to trust almost entirely to analogy, and his difficulties will rise to a climax.</p>
 			<p>Certainly no clear line of demarcation has as yet been drawn between species and subspecies⁠—that is, the forms which in the opinion of some naturalists come very near to, but do not quite arrive at, the rank of species; or, again, between subspecies and well-marked varieties, or between lesser varieties and individual differences. These differences blend into each other by an insensible series; and a series impresses the mind with the idea of an actual passage.</p>
 			<p>Hence I look at individual differences, though of small interest to the systematist, as of the highest importance for us, as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. And I look at varieties which are in any degree more distinct and permanent, as steps toward more strongly marked and permanent varieties; and at the latter, as leading to subspecies, and then to species. The passage from one stage of difference to another may, in many cases, be the simple result of the nature of the organism and of the different physical conditions to which it has long been exposed; but with respect to the more important and adaptive characters, the passage from one stage of difference to another may be safely attributed to the cumulative action of natural selection, hereafter to be explained, and to the effects of the increased use or disuse of parts. A well-marked variety may therefore be called an incipient species; but whether this belief is justifiable must be judged by the weight of the various facts and considerations to be given throughout this work.</p>

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 			<p>Before entering on the subject of this chapter I must make a few preliminary remarks to show how the struggle for existence bears on natural selection. It has been seen in the last chapter that among organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or subspecies or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life and of one organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.</p>
 			<p>Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow from the struggle for life. Owing to this struggle, variations, however slight and from whatever cause proceeding, if they be in any degree profitable to the individuals of a species, in their infinitely complex relations to other organic beings and to their physical conditions of life, will tend to the preservation of such individuals, and will generally be inherited by the offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term natural selection, in order to mark its relation to man’s power of selection. But the expression often used by <abbr>Mr.</abbr> Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man’s feeble efforts, as the works of Nature are to those of Art.</p>
-			<p>We will now discuss in a little more detail the struggle for existence. In my future work this subject will be treated, as it well deserves, at greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult⁠—at least I found it so⁠—than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that, though food may be now superabundant, it is not so at all seasons of each recurring year.</p>
+			<p>We will now discuss in a little more detail the struggle for existence. In my future work this subject will be treated, as it well deserves, at greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than <abbr class="name">W.</abbr> Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult⁠—at least I found it so⁠—than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that, though food may be now superabundant, it is not so at all seasons of each recurring year.</p>
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 			<h3 epub:type="title">The Term, Struggle for Existence, Used in a Large Sense</h3>
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 			<h3 epub:type="title">Complex Relations of All Animals and Plants to Each Other in the Struggle for Existence</h3>
 			<p>Many cases are on record showing how complex and unexpected are the checks and relations between organic beings, which have to struggle together in the same country. I will give only a single instance, which, though a simple one, interested me. In Staffordshire, on the estate of a relation, where I had ample means of investigation, there was a large and extremely barren heath, which had never been touched by the hand of man; but several hundred acres of exactly the same nature had been enclosed twenty-five years previously and planted with Scotch fir. The change in the native vegetation of the planted part of the heath was most remarkable, more than is generally seen in passing from one quite different soil to another: not only the proportional numbers of the heath-plants were wholly changed, but twelve species of plants (not counting grasses and carices) flourished in the plantations, which could not be found on the heath. The effect on the insects must have been still greater, for six insectivorous birds were very common in the plantations, which were not to be seen on the heath; and the heath was frequented by two or three distinct insectivorous birds. Here we see how potent has been the effect of the introduction of a single tree, nothing whatever else having been done, with the exception of the land having been enclosed, so that cattle could not enter. But how important an element enclosure is, I plainly saw near Farnham, in Surrey. Here there are extensive heaths, with a few clumps of old Scotch firs on the distant hilltops: within the last ten years large spaces have been enclosed, and self-sown firs are now springing up in multitudes, so close together that all cannot live. When I ascertained that these young trees had not been sown or planted I was so much surprised at their numbers that I went to several points of view, whence I could examine hundreds of acres of the unenclosed heath, and literally I could not see a single Scotch fir, except the old planted clumps. But on looking closely between the stems of the heath, I found a multitude of seedlings and little trees, which had been perpetually browsed down by the cattle. In one square yard, at a point some hundred yards distant from one of the old clumps, I counted thirty-two little trees; and one of them, with twenty-six rings of growth, had, during many years tried to raise its head above the stems of the heath, and had failed. No wonder that, as soon as the land was enclosed, it became thickly clothed with vigorously growing young firs. Yet the heath was so extremely barren and so extensive that no one would ever have imagined that cattle would have so closely and effectually searched it for food.</p>
 			<p>Here we see that cattle absolutely determine the existence of the Scotch fir; but in several parts of the world insects determine the existence of cattle. Perhaps Paraguay offers the most curious instance of this; for here neither cattle nor horses nor dogs have ever run wild, though they swarm southward and northward in a feral state; and Azara and Rengger have shown that this is caused by the greater number in Paraguay of a certain fly, which lays its eggs in the navels of these animals when first born. The increase of these flies, numerous as they are, must be habitually checked by some means, probably by other parasitic insects. Hence, if certain insectivorous birds were to decrease in Paraguay, the parasitic insects would probably increase; and this would lessen the number of the navel-frequenting flies⁠—then cattle and horses would become feral, and this would certainly greatly alter (as indeed I have observed in parts of South America) the vegetation: this again would largely affect the insects; and this, as we have just seen in Staffordshire, the insectivorous birds, and so onwards in ever-increasing circles of complexity. Not that under nature the relations will ever be as simple as this. Battle within battle must be continually recurring with varying success; and yet in the long-run the forces are so nicely balanced that the face of nature remains for long periods of time uniform, though assuredly the merest trifle would give the victory to one organic being over another. Nevertheless, so profound is our ignorance, and so high our presumption, that we marvel when we hear of the extinction of an organic being; and as we do not see the cause, we invoke cataclysms to desolate the world, or invent laws on the duration of the forms of life!</p>
-			<p>I am tempted to give one more instance showing how plants and animals, remote in the scale of nature, are bound together by a web of complex relations. I shall hereafter have occasion to show that the exotic Lobelia fulgens is never visited in my garden by insects, and consequently, from its peculiar structure, never sets a seed. Nearly all our orchidaceous plants absolutely require the visits of insects to remove their pollen-masses and thus to fertilise them. I find from experiments that humblebees are almost indispensable to the fertilisation of the heartsease (Viola tricolor), for other bees do not visit this flower. I have also found that the visits of bees are necessary for the fertilisation of some kinds of clover; for instance twenty heads of Dutch clover (Trifolium repens) yielded 2,290 seeds, but twenty other heads, protected from bees, produced not one. Again, 100 heads of red clover (T. pratense) produced 2,700 seeds, but the same number of protected heads produced not a single seed. Humble bees alone visit red clover, as other bees cannot reach the nectar. It has been suggested that moths may fertilise the clovers; but I doubt whether they could do so in the case of the red clover, from their weight not being sufficient to depress the wing petals. Hence we may infer as highly probable that, if the whole genus of humblebees became extinct or very rare in England, the heartsease and red clover would become very rare, or wholly disappear. The number of humblebees in any district depends in a great measure upon the number of field-mice, which destroy their combs and nests; and Colonel Newman, who has long attended to the habits of humblebees, believes that “more than two-thirds of them are thus destroyed all over England.” Now the number of mice is largely dependent, as everyone knows, on the number of cats; and Colonel Newman says, “Near villages and small towns I have found the nests of humblebees more numerous than elsewhere, which I attribute to the number of cats that destroy the mice.” Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine, through the intervention first of mice and then of bees, the frequency of certain flowers in that district!</p>
+			<p>I am tempted to give one more instance showing how plants and animals, remote in the scale of nature, are bound together by a web of complex relations. I shall hereafter have occasion to show that the exotic <i epub:type="z3998:taxonomy">Lobelia fulgens</i> is never visited in my garden by insects, and consequently, from its peculiar structure, never sets a seed. Nearly all our orchidaceous plants absolutely require the visits of insects to remove their pollen-masses and thus to fertilise them. I find from experiments that humblebees are almost indispensable to the fertilisation of the heartsease (<i epub:type="z3998:taxonomy">Viola tricolor</i>), for other bees do not visit this flower. I have also found that the visits of bees are necessary for the fertilisation of some kinds of clover; for instance twenty heads of Dutch clover (<i epub:type="z3998:taxonomy">Trifolium repens</i>) yielded 2,290 seeds, but twenty other heads, protected from bees, produced not one. Again, 100 heads of red clover (<i epub:type="z3998:taxonomy">T. pratense</i>) produced 2,700 seeds, but the same number of protected heads produced not a single seed. Humblebees alone visit red clover, as other bees cannot reach the nectar. It has been suggested that moths may fertilise the clovers; but I doubt whether they could do so in the case of the red clover, from their weight not being sufficient to depress the wing petals. Hence we may infer as highly probable that, if the whole genus of humblebees became extinct or very rare in England, the heartsease and red clover would become very rare, or wholly disappear. The number of humblebees in any district depends in a great measure upon the number of field-mice, which destroy their combs and nests; and Colonel Newman, who has long attended to the habits of humblebees, believes that “more than two-thirds of them are thus destroyed all over England.” Now the number of mice is largely dependent, as everyone knows, on the number of cats; and Colonel Newman says, “Near villages and small towns I have found the nests of humblebees more numerous than elsewhere, which I attribute to the number of cats that destroy the mice.” Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine, through the intervention first of mice and then of bees, the frequency of certain flowers in that district!</p>
 			<p>In the case of every species, many different checks, acting at different periods of life, and during different seasons or years, probably come into play; some one check or some few being generally the most potent, but all will concur in determining the average number, or even the existence of the species. In some cases it can be shown that widely-different checks act on the same species in different districts. When we look at the plants and bushes clothing an entangled bank, we are tempted to attribute their proportional numbers and kinds to what we call chance. But how false a view is this! Everyone has heard that when an American forest is cut down, a very different vegetation springs up; but it has been observed that ancient Indian ruins in the Southern United States, which must formerly have been cleared of trees, now display the same beautiful diversity and proportion of kinds as in the surrounding virgin forests. What a struggle must have gone on during long centuries between the several kinds of trees, each annually scattering its seeds by the thousand; what war between insect and insect⁠—between insects, snails, and other animals with birds and beasts of prey⁠—all striving to increase, all feeding on each other, or on the trees, their seeds and seedlings, or on the other plants which first clothed the ground and thus checked the growth of the trees. Throw up a handful of feathers, and all fall to the ground according to definite laws; but how simple is the problem where each shall fall compared to that of the action and reaction of the innumerable plants and animals which have determined, in the course of centuries, the proportional numbers and kinds of trees now growing on the old Indian ruins!</p>
 			<p>The dependency of one organic being on another, as of a parasite on its prey, lies generally between beings remote in the scale of nature. This is likewise sometimes the case with those which may strictly be said to struggle with each other for existence, as in the case of locusts and grass-feeding quadrupeds. But the struggle will almost invariably be most severe between the individuals of the same species, for they frequent the same districts, require the same food, and are exposed to the same dangers. In the case of varieties of the same species, the struggle will generally be almost equally severe, and we sometimes see the contest soon decided: for instance, if several varieties of wheat be sown together, and the mixed seed be resown, some of the varieties which best suit the soil or climate, or are naturally the most fertile, will beat the others and so yield more seed, and will consequently in a few years supplant the other varieties. To keep up a mixed stock of even such extremely close varieties as the variously coloured sweet-peas, they must be each year harvested separately, and the seed then mixed in due proportion, otherwise the weaker kinds will steadily decrease in number and disappear. So again with the varieties of sheep: it has been asserted that certain mountain-varieties will starve out other mountain-varieties, so that they cannot be kept together. The same result has followed from keeping together different varieties of the medicinal leech. It may even be doubted whether the varieties of any of our domestic plants or animals have so exactly the same strength, habits, and constitution, that the original proportions of a mixed stock (crossing being prevented) could be kept up for half-a-dozen generations, if they were allowed to struggle together, in the same manner as beings in a state of nature, and if the seed or young were not annually preserved in due proportion.</p>
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 			<h3 epub:type="title">Sexual Selection</h3>
-			<p>Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, so no doubt it will be under nature. Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life, as is sometimes the case; or for one sex to be modified in relation to the other sex, as commonly occurs. This leads me to say a few words on what I have called sexual selection. This form of selection depends, not on a struggle for existence in relation to other organic beings or to external conditions, but on a struggle between the individuals of one sex, generally the males, for the possession of the other sex. The result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection. Generally, the most vigorous males, those which are best fitted for their places in nature, will leave most progeny. But in many cases victory depends not so much on general vigour, but on having special weapons, confined to the male sex. A hornless stag or spurless cock would have a poor chance of leaving numerous offspring. Sexual selection, by always allowing the victor to breed, might surely give indomitable courage, length of spur, and strength to the wing to strike in the spurred leg, in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks. How low in the scale of nature the law of battle descends I know not; male alligators have been described as fighting, bellowing, and whirling round, like Indians in a war-dance, for the possession of the females; male salmons have been observed fighting all day long; male stag-beetles sometimes bear wounds from the huge mandibles of other males; the males of certain hymenopterous insects have been frequently seen by that inimitable observer M. Fabre, fighting for a particular female who sits by, an apparently unconcerned beholder of the struggle, and then retires with the conqueror. The war is, perhaps, severest between the males of polygamous animals, and these seem oftenest provided with special weapons. The males of carnivorous animals are already well armed; though to them and to others, special means of defence may be given through means of sexual selection, as the mane of the lion, and the hooked jaw to the male salmon; for the shield may be as important for victory as the sword or spear.</p>
-			<p>Among birds, the contest is often of a more peaceful character. All those who have attended to the subject, believe that there is the severest rivalry between the males of many species to attract, by singing, the females. The rock-thrush of Guiana, birds of paradise, and some others, congregate, and successive males display with the most elaborate care, and show off in the best manner, their gorgeous plumage; they likewise perform strange antics before the females, which, standing by as spectators, at last choose the most attractive partner. Those who have closely attended to birds in confinement well know that they often take individual preferences and dislikes: thus Sir R. Heron has described how a pied peacock was eminently attractive to all his hen birds. I cannot here enter on the necessary details; but if man can in a short time give beauty and an elegant carriage to his bantams, according to his standard of beauty, I can see no good reason to doubt that female birds, by selecting, during thousands of generations, the most melodious or beautiful males, according to their standard of beauty, might produce a marked effect. Some well-known laws, with respect to the plumage of male and female birds, in comparison with the plumage of the young, can partly be explained through the action of sexual selection on variations occurring at different ages, and transmitted to the males alone or to both sexes at corresponding ages; but I have not space here to enter on this subject.</p>
+			<p>Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, so no doubt it will be under nature. Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life, as is sometimes the case; or for one sex to be modified in relation to the other sex, as commonly occurs. This leads me to say a few words on what I have called sexual selection. This form of selection depends, not on a struggle for existence in relation to other organic beings or to external conditions, but on a struggle between the individuals of one sex, generally the males, for the possession of the other sex. The result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection. Generally, the most vigorous males, those which are best fitted for their places in nature, will leave most progeny. But in many cases victory depends not so much on general vigour, but on having special weapons, confined to the male sex. A hornless stag or spurless cock would have a poor chance of leaving numerous offspring. Sexual selection, by always allowing the victor to breed, might surely give indomitable courage, length of spur, and strength to the wing to strike in the spurred leg, in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks. How low in the scale of nature the law of battle descends I know not; male alligators have been described as fighting, bellowing, and whirling round, like Indians in a war-dance, for the possession of the females; male salmons have been observed fighting all day long; male stag-beetles sometimes bear wounds from the huge mandibles of other males; the males of certain hymenopterous insects have been frequently seen by that inimitable observer <abbr class="name">M.</abbr> Fabre, fighting for a particular female who sits by, an apparently unconcerned beholder of the struggle, and then retires with the conqueror. The war is, perhaps, severest between the males of polygamous animals, and these seem oftenest provided with special weapons. The males of carnivorous animals are already well armed; though to them and to others, special means of defence may be given through means of sexual selection, as the mane of the lion, and the hooked jaw to the male salmon; for the shield may be as important for victory as the sword or spear.</p>
+			<p>Among birds, the contest is often of a more peaceful character. All those who have attended to the subject, believe that there is the severest rivalry between the males of many species to attract, by singing, the females. The rock-thrush of Guiana, birds of paradise, and some others, congregate, and successive males display with the most elaborate care, and show off in the best manner, their gorgeous plumage; they likewise perform strange antics before the females, which, standing by as spectators, at last choose the most attractive partner. Those who have closely attended to birds in confinement well know that they often take individual preferences and dislikes: thus Sir <abbr class="name">R.</abbr> Heron has described how a pied peacock was eminently attractive to all his hen birds. I cannot here enter on the necessary details; but if man can in a short time give beauty and an elegant carriage to his bantams, according to his standard of beauty, I can see no good reason to doubt that female birds, by selecting, during thousands of generations, the most melodious or beautiful males, according to their standard of beauty, might produce a marked effect. Some well-known laws, with respect to the plumage of male and female birds, in comparison with the plumage of the young, can partly be explained through the action of sexual selection on variations occurring at different ages, and transmitted to the males alone or to both sexes at corresponding ages; but I have not space here to enter on this subject.</p>
 			<p>Thus it is, as I believe, that when the males and females of any animal have the same general habits of life, but differ in structure, colour, or ornament, such differences have been mainly caused by sexual selection: that is, by individual males having had, in successive generations, some slight advantage over other males, in their weapons, means of defence, or charms; which they have transmitted to their male offspring alone. Yet, I would not wish to attribute all sexual differences to this agency: for we see in our domestic animals peculiarities arising and becoming attached to the male sex, which apparently have not been augmented through selection by man. The tuft of hair on the breast of the wild turkey-cock cannot be of any use, and it is doubtful whether it can be ornamental in the eyes of the female bird; indeed, had the tuft appeared under domestication it would have been called a monstrosity.</p>
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 		<section id="chapter-4-2" epub:type="z3998:subchapter">
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 			<p>To the effects of intercrossing in eliminating variations of all kinds, I shall have to recur; but it may be here remarked that most animals and plants keep to their proper homes, and do not needlessly wander about; we see this even with migratory birds, which almost always return to the same spot. Consequently each newly-formed variety would generally be at first local, as seems to be the common rule with varieties in a state of nature; so that similarly modified individuals would soon exist in a small body together, and would often breed together. If the new variety were successful in its battle for life, it would slowly spread from a central district, competing with and conquering the unchanged individuals on the margins of an ever-increasing circle.</p>
 			<p>It may be worth while to give another and more complex illustration of the action of natural selection. Certain plants excrete sweet juice, apparently for the sake of eliminating something injurious from the sap: this is effected, for instance, by glands at the base of the stipules in some Leguminosae, and at the backs of the leaves of the common laurel. This juice, though small in quantity, is greedily sought by insects; but their visits do not in any way benefit the plant. Now, let us suppose that the juice or nectar was excreted from the inside of the flowers of a certain number of plants of any species. Insects in seeking the nectar would get dusted with pollen, and would often transport it from one flower to another. The flowers of two distinct individuals of the same species would thus get crossed; and the act of crossing, as can be fully proved, gives rise to vigorous seedlings, which consequently would have the best chance of flourishing and surviving. The plants which produced flowers with the largest glands or nectaries, excreting most nectar, would oftenest be visited by insects, and would oftenest be crossed; and so in the long-run would gain the upper hand and form a local variety. The flowers, also, which had their stamens and pistils placed, in relation to the size and habits of the particular insect which visited them, so as to favour in any degree the transportal of the pollen, would likewise be favoured. We might have taken the case of insects visiting flowers for the sake of collecting pollen instead of nectar; and as pollen is formed for the sole purpose of fertilisation, its destruction appears to be a simple loss to the plant; yet if a little pollen were carried, at first occasionally and then habitually, by the pollen-devouring insects from flower to flower, and a cross thus effected, although nine-tenths of the pollen were destroyed it might still be a great gain to the plant to be thus robbed; and the individuals which produced more and more pollen, and had larger anthers, would be selected.</p>
 			<p>When our plant, by the above process long continued, had been rendered highly attractive to insects, they would, unintentionally on their part, regularly carry pollen from flower to flower; and that they do this effectually I could easily show by many striking facts. I will give only one, as likewise illustrating one step in the separation of the sexes of plants. Some holly-trees bear only male flowers, which have four stamens producing a rather small quantity of pollen, and a rudimentary pistil; other holly-trees bear only female flowers; these have a full-sized pistil, and four stamens with shrivelled anthers, in which not a grain of pollen can be detected. Having found a female tree exactly sixty yards from a male tree, I put the stigmas of twenty flowers, taken from different branches, under the microscope, and on all, without exception, there were a few pollen-grains, and on some a profusion. As the wind had set for several days from the female to the male tree, the pollen could not thus have been carried. The weather had been cold and boisterous and therefore not favourable to bees, nevertheless every female flower which I examined had been effectually fertilised by the bees, which had flown from tree to tree in search of nectar. But to return to our imaginary case; as soon as the plant had been rendered so highly attractive to insects that pollen was regularly carried from flower to flower, another process might commence. No naturalist doubts the advantage of what has been called the “physiological division of labour;” hence we may believe that it would be advantageous to a plant to produce stamens alone in one flower or on one whole plant, and pistils alone in another flower or on another plant. In plants under culture and placed under new conditions of life, sometimes the male organs and sometimes the female organs become more or less impotent; now if we suppose this to occur in ever so slight a degree under nature, then, as pollen is already carried regularly from flower to flower, and as a more complete separation of the sexes of our plant would be advantageous on the principle of the division of labour, individuals with this tendency more and more increased, would be continually favoured or selected, until at last a complete separation of the sexes might be effected. It would take up too much space to show the various steps, through dimorphism and other means, by which the separation of the sexes in plants of various kinds is apparently now in progress; but I may add that some of the species of holly in North America are, according to Asa Gray, in an exactly intermediate condition, or, as he expresses it, are more or less dioeciously polygamous.</p>
-			<p>Let us now turn to the nectar-feeding insects; we may suppose the plant of which we have been slowly increasing the nectar by continued selection, to be a common plant; and that certain insects depended in main part on its nectar for food. I could give many facts showing how anxious bees are to save time: for instance, their habit of cutting holes and sucking the nectar at the bases of certain flowers, which with a very little more trouble they can enter by the mouth. Bearing such facts in mind, it may be believed that under certain circumstances individual differences in the curvature or length of the proboscis, <abbr>etc.</abbr>, too slight to be appreciated by us, might profit a bee or other insect, so that certain individuals would be able to obtain their food more quickly than others; and thus the communities to which they belonged would flourish and throw off many swarms inheriting the same peculiarities. The tubes of the corolla of the common red or incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty glance appear to differ in length; yet the hive-bee can easily suck the nectar out of the incarnate clover, but not out of the common red clover, which is visited by humblebees alone; so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. That this nectar is much liked by the hive-bee is certain; for I have repeatedly seen, but only in the autumn, many hive-bees sucking the flowers through holes bitten in the base of the tube by humble bees. The difference in the length of the corolla in the two kinds of clover, which determines the visits of the hive-bee, must be very trifling; for I have been assured that when red clover has been mown, the flowers of the second crop are somewhat smaller, and that these are visited by many hive-bees. I do not know whether this statement is accurate; nor whether another published statement can be trusted, namely, that the Ligurian bee, which is generally considered a mere variety of the common hive-bee, and which freely crosses with it, is able to reach and suck the nectar of the red clover. Thus, in a country where this kind of clover abounded, it might be a great advantage to the hive-bee to have a slightly longer or differently constructed proboscis. On the other hand, as the fertility of this clover absolutely depends on bees visiting the flowers, if humblebees were to become rare in any country, it might be a great advantage to the plant to have a shorter or more deeply divided corolla, so that the hive-bees should be enabled to suck its flowers. Thus I can understand how a flower and a bee might slowly become, either simultaneously or one after the other, modified and adapted to each other in the most perfect manner, by the continued preservation of all the individuals which presented slight deviations of structure mutually favourable to each other.</p>
+			<p>Let us now turn to the nectar-feeding insects; we may suppose the plant of which we have been slowly increasing the nectar by continued selection, to be a common plant; and that certain insects depended in main part on its nectar for food. I could give many facts showing how anxious bees are to save time: for instance, their habit of cutting holes and sucking the nectar at the bases of certain flowers, which with a very little more trouble they can enter by the mouth. Bearing such facts in mind, it may be believed that under certain circumstances individual differences in the curvature or length of the proboscis, <abbr>etc.</abbr>, too slight to be appreciated by us, might profit a bee or other insect, so that certain individuals would be able to obtain their food more quickly than others; and thus the communities to which they belonged would flourish and throw off many swarms inheriting the same peculiarities. The tubes of the corolla of the common red or incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty glance appear to differ in length; yet the hive-bee can easily suck the nectar out of the incarnate clover, but not out of the common red clover, which is visited by humblebees alone; so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. That this nectar is much liked by the hive-bee is certain; for I have repeatedly seen, but only in the autumn, many hive-bees sucking the flowers through holes bitten in the base of the tube by humblebees. The difference in the length of the corolla in the two kinds of clover, which determines the visits of the hive-bee, must be very trifling; for I have been assured that when red clover has been mown, the flowers of the second crop are somewhat smaller, and that these are visited by many hive-bees. I do not know whether this statement is accurate; nor whether another published statement can be trusted, namely, that the Ligurian bee, which is generally considered a mere variety of the common hive-bee, and which freely crosses with it, is able to reach and suck the nectar of the red clover. Thus, in a country where this kind of clover abounded, it might be a great advantage to the hive-bee to have a slightly longer or differently constructed proboscis. On the other hand, as the fertility of this clover absolutely depends on bees visiting the flowers, if humblebees were to become rare in any country, it might be a great advantage to the plant to have a shorter or more deeply divided corolla, so that the hive-bees should be enabled to suck its flowers. Thus I can understand how a flower and a bee might slowly become, either simultaneously or one after the other, modified and adapted to each other in the most perfect manner, by the continued preservation of all the individuals which presented slight deviations of structure mutually favourable to each other.</p>
 			<p>I am well aware that this doctrine of natural selection, exemplified in the above imaginary instances, is open to the same objections which were first urged against Sir Charles Lyell’s noble views on “the modern changes of the earth, as illustrative of geology;” but we now seldom hear the agencies which we see still at work, spoken of as trifling and insignificant, when used in explaining the excavation of the deepest valleys or the formation of long lines of inland cliffs. Natural selection acts only by the preservation and accumulation of small inherited modifications, each profitable to the preserved being; and as modern geology has almost banished such views as the excavation of a great valley by a single diluvial wave, so will natural selection banish the belief of the continued creation of new organic beings, or of any great and sudden modification in their structure.</p>
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 		<section id="chapter-4-3" epub:type="z3998:subchapter">

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 				<p epub:type="bridgehead">Effects of Changed Conditions⁠—Use and Disuse, Combined with Natural Selection; organs of Flight and of Vision⁠—Acclimatisation⁠—Correlated Variation⁠—Compensation and Economy of Growth⁠—False Correlations⁠—Multiple, Rudimentary, and Lowly Organised Structures Variable⁠—Parts Developed in an Unusual Manner Are Highly Variable: Specific Characters More Variable Than Generic: Secondary Sexual Characters Variable⁠—Species of the Same Genus Vary in an Analogous Manner⁠—Reversions to Long-Lost Characters⁠—Summary.</p>
 			</header>
 			<p>I have hitherto sometimes spoken as if the variations⁠—so common and multiform with organic beings under domestication, and in a lesser degree with those under nature⁠—were due to chance. This, of course is a wholly incorrect expression, but it serves to acknowledge plainly our ignorance of the cause of each particular variation. Some authors believe it to be as much the function of the reproductive system to produce individual differences, or slight deviations of structure, as to make the child like its parents. But the fact of variations and monstrosities occurring much more frequently under domestication than under nature, and the greater variability of species having wide ranges than of those with restricted ranges, lead to the conclusion that variability is generally related to the conditions of life to which each species has been exposed during several successive generations. In the first chapter I attempted to show that changed conditions act in two ways, directly on the whole organisation or on certain parts alone, and indirectly through the reproductive system. In all cases there are two factors, the nature of the organism, which is much the most important of the two, and the nature of the conditions. The direct action of changed conditions leads to definite or indefinite results. In the latter case the organisation seems to become plastic, and we have much fluctuating variability. In the former case the nature of the organism is such that it yields readily, when subjected to certain conditions, and all, or nearly all, the individuals become modified in the same way.</p>
-			<p>It is very difficult to decide how far changed conditions, such as of climate, food, <abbr>etc.</abbr>, have acted in a definite manner. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence. But we may safely conclude that the innumerable complex co-adaptations of structure, which we see throughout nature between various organic beings, cannot be attributed simply to such action. In the following cases the conditions seem to have produced some slight definite effect: E. Forbes asserts that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species from further north or from a greater depth; but this certainly does not always hold good. <abbr>Mr.</abbr> Gould believes that birds of the same species are more brightly coloured under a clear atmosphere, than when living near the coast or on islands; and Wollaston is convinced that residence near the sea affects the colours of insects. Moquin-Tandon gives a list of plants which, when growing near the seashore, have their leaves in some degree fleshy, though not elsewhere fleshy. These slightly varying organisms are interesting in as far as they present characters analogous to those possessed by the species which are confined to similar conditions.</p>
+			<p>It is very difficult to decide how far changed conditions, such as of climate, food, <abbr>etc.</abbr>, have acted in a definite manner. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence. But we may safely conclude that the innumerable complex co-adaptations of structure, which we see throughout nature between various organic beings, cannot be attributed simply to such action. In the following cases the conditions seem to have produced some slight definite effect: <abbr class="name">E.</abbr> Forbes asserts that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species from further north or from a greater depth; but this certainly does not always hold good. <abbr>Mr.</abbr> Gould believes that birds of the same species are more brightly coloured under a clear atmosphere, than when living near the coast or on islands; and Wollaston is convinced that residence near the sea affects the colours of insects. Moquin-Tandon gives a list of plants which, when growing near the seashore, have their leaves in some degree fleshy, though not elsewhere fleshy. These slightly varying organisms are interesting in as far as they present characters analogous to those possessed by the species which are confined to similar conditions.</p>
 			<p>When a variation is of the slightest use to any being, we cannot tell how much to attribute to the accumulative action of natural selection, and how much to the definite action of the conditions of life. Thus, it is well known to furriers that animals of the same species have thicker and better fur the further north they live; but who can tell how much of this difference may be due to the warmest-clad individuals having been favoured and preserved during many generations, and how much to the action of the severe climate? For it would appear that climate has some direct action on the hair of our domestic quadrupeds.</p>
 			<p>Instances could be given of similar varieties being produced from the same species under external conditions of life as different as can well be conceived; and, on the other hand, of dissimilar varieties being produced under apparently the same external conditions. Again, innumerable instances are known to every naturalist, of species keeping true, or not varying at all, although living under the most opposite climates. Such considerations as these incline me to lay less weight on the direct action of the surrounding conditions, than on a tendency to vary, due to causes of which we are quite ignorant.</p>
 			<p>In one sense the conditions of life may be said, not only to cause variability, either directly or indirectly, but likewise to include natural selection, for the conditions determine whether this or that variety shall survive. But when man is the selecting agent, we clearly see that the two elements of change are distinct; variability is in some manner excited, but it is the will of man which accumulates the variations in certain direction; and it is this latter agency which answers to the survival of the fittest under nature.</p>
@@ -41,7 +41,7 @@
 			<h3 epub:type="title">Correlated Variation</h3>
 			<p>I mean by this expression that the whole organisation is so tied together, during its growth and development, that when slight variations in any one part occur and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood, and no doubt wholly different classes of facts may be here easily confounded together. We shall presently see that simple inheritance often gives the false appearance of correlation. One of the most obvious real cases is, that variations of structure arising in the young or larvae naturally tend to affect the structure of the mature animal. The several parts which are homologous, and which, at an early embryonic period, are identical in structure, and which are necessarily exposed to similar conditions, seem eminently liable to vary in a like manner: we see this in the right and left sides of the body varying in the same manner; in the front and hind legs, and even in the jaws and limbs, varying together, for the lower jaw is believed by some anatomists to be homologous with the limbs. These tendencies, I do not doubt, may be mastered more or less completely by natural selection: thus a family of stags once existed with an antler only on one side; and if this had been of any great use to the breed, it might probably have been rendered permanent by natural selection.</p>
 			<p>Homologous parts, as has been remarked by some authors, tend to cohere; this is often seen in monstrous plants: and nothing is more common than the union of homologous parts in normal structures, as in the union of the petals into a tube. Hard parts seem to affect the form of adjoining soft parts; it is believed by some authors that with birds the diversity in the shape of the pelvis causes the remarkable diversity in the shape of the kidneys. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. In snakes, according to Schlegel, the shape of the body and the manner of swallowing determine the position and form of several of the most important viscera.</p>
-			<p>The nature of the bond is frequently quite obscure. M. Is. Geoffroy <abbr class="name">St.</abbr> Hilaire has forcibly remarked that certain malconformations frequently, and that others rarely, coexist without our being able to assign any reason. What can be more singular than the relation in cats between complete whiteness and blue eyes with deafness, or between the tortoiseshell colour and the female sex; or in pigeons, between their feathered feet and skin betwixt the outer toes, or between the presence of more or less down on the young pigeon when first hatched, with the future colour of its plumage; or, again, the relation between the hair and the teeth in the naked Turkish dog, though here no doubt homology comes into play? With respect to this latter case of correlation, I think it can hardly be accidental that the two orders of mammals which are most abnormal in their dermal covering, <abbr>viz.</abbr>, Cetacea (whales) and Edentata (armadilloes, scaly anteaters, <abbr>etc.</abbr>), are likewise on the whole the most abnormal in their teeth, but there are so many exceptions to this rule, as <abbr>Mr.</abbr> Mivart has remarked, that it has little value.</p>
+			<p>The nature of the bond is frequently quite obscure. <abbr>M.</abbr> <abbr class="name">Is.</abbr> Geoffroy <abbr class="name">St.</abbr> Hilaire has forcibly remarked that certain malconformations frequently, and that others rarely, coexist without our being able to assign any reason. What can be more singular than the relation in cats between complete whiteness and blue eyes with deafness, or between the tortoiseshell colour and the female sex; or in pigeons, between their feathered feet and skin betwixt the outer toes, or between the presence of more or less down on the young pigeon when first hatched, with the future colour of its plumage; or, again, the relation between the hair and the teeth in the naked Turkish dog, though here no doubt homology comes into play? With respect to this latter case of correlation, I think it can hardly be accidental that the two orders of mammals which are most abnormal in their dermal covering, <abbr>viz.</abbr>, Cetacea (whales) and Edentata (armadilloes, scaly anteaters, <abbr>etc.</abbr>), are likewise on the whole the most abnormal in their teeth, but there are so many exceptions to this rule, as <abbr>Mr.</abbr> Mivart has remarked, that it has little value.</p>
 			<p>I know of no case better adapted to show the importance of the laws of correlation and variation, independently of utility, and therefore of natural selection, than that of the difference between the outer and inner flowers in some Compositous and Umbelliferous plants. Everyone is familiar with the difference between the ray and central florets of, for instance, the daisy, and this difference is often accompanied with the partial or complete abortion of the reproductive organs. But in some of these plants the seeds also differ in shape and sculpture. These differences have sometimes been attributed to the pressure of the involucra on the florets, or to their mutual pressure, and the shape of the seeds in the ray-florets of some Compositae countenances this idea; but with the Umbelliferae it is by no means, as <abbr>Dr.</abbr> Hooker informs me, the species with the densest heads which most frequently differ in their inner and outer flowers. It might have been thought that the development of the ray-petals, by drawing nourishment from the reproductive organs causes their abortion; but this can hardly be the sole case, for in some Compositae the seeds of the outer and inner florets differ, without any difference in the corolla. Possibly these several differences may be connected with the different flow of nutriment towards the central and external flowers. We know, at least, that with irregular flowers those nearest to the axis are most subject to peloria, that is to become abnormally symmetrical. I may add, as an instance of this fact, and as a striking case of correlation, that in many pelargoniums the two upper petals in the central flower of the truss often lose their patches of darker colour; and when this occurs, the adherent nectary is quite aborted, the central flower thus becoming peloric or regular. When the colour is absent from only one of the two upper petals, the nectary is not quite aborted but is much shortened.</p>
 			<p>With respect to the development of the corolla, Sprengel’s idea that the ray-florets serve to attract insects, whose agency is highly advantageous, or necessary for the fertilisation of these plants, is highly probable; and if so, natural selection may have come into play. But with respect to the seeds, it seems impossible that their differences in shape, which are not always correlated with any difference in the corolla, can be in any way beneficial; yet in the Umbelliferae these differences are of such apparent importance⁠—the seeds being sometimes orthospermous in the exterior flowers and coelospermous in the central flowers⁠—that the elder De Candolle founded his main divisions in the order on such characters. Hence modifications of structure, viewed by systematists as of high value, may be wholly due to the laws of variation and correlation, without being, as far as we can judge, of the slightest service to the species.</p>
 			<p>We may often falsely attribute to correlated variation structures which are common to whole groups of species, and which in truth are simply due to inheritance; for an ancient progenitor may have acquired through natural selection some one modification in structure, and, after thousands of generations, some other and independent modification; and these two modifications, having been transmitted to a whole group of descendants with diverse habits, would naturally be thought to be in some necessary manner correlated. Some other correlations are apparently due to the manner in which natural selection can alone act. For instance, Alph. De Candolle has remarked that winged seeds are never found in fruits which do not open; I should explain this rule by the impossibility of seeds gradually becoming winged through natural selection, unless the capsules were open; for in this case alone could the seeds, which were a little better adapted to be wafted by the wind, gain an advantage over others less well fitted for wide dispersal.</p>
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 		<section id="chapter-5-8" epub:type="z3998:subchapter">
 			<h3 epub:type="title">Secondary Sexual Characters Variable</h3>
 			<p>I think it will be admitted by naturalists, without my entering on details, that secondary sexual characters are highly variable. It will also be admitted that species of the same group differ from each other more widely in their secondary sexual characters, than in other parts of their organisation; compare, for instance, the amount of difference between the males of gallinaceous birds, in which secondary sexual characters are strongly displayed, with the amount of difference between the females. The cause of the original variability of these characters is not manifest; but we can see why they should not have been rendered as constant and uniform as others, for they are accumulated by sexual selection, which is less rigid in its action than ordinary selection, as it does not entail death, but only gives fewer offspring to the less favoured males. Whatever the cause may be of the variability of secondary sexual characters, as they are highly variable, sexual selection will have had a wide scope for action, and may thus have succeeded in giving to the species of the same group a greater amount of difference in these than in other respects.</p>
-			<p>It is a remarkable fact, that the secondary differences between the two sexes of the same species are generally displayed in the very same parts of the organisation in which the species of the same genus differ from each other. Of this fact I will give in illustration the first two instances which happen to stand on my list; and as the differences in these cases are of a very unusual nature, the relation can hardly be accidental. The same number of joints in the tarsi is a character common to very large groups of beetles, but in the Engidae, as Westwood has remarked, the number varies greatly and the number likewise differs in the two sexes of the same species. Again in the fossorial hymenoptera, the neuration of the wings is a character of the highest importance, because common to large groups; but in certain genera the neuration differs in the different species, and likewise in the two sexes of the same species. Sir J. Lubbock has recently remarked, that several minute crustaceans offer excellent illustrations of this law. “In Pontella, for instance, the sexual characters are afforded mainly by the anterior antennae and by the fifth pair of legs: the specific differences also are principally given by these organs.” This relation has a clear meaning on my view: I look at all the species of the same genus as having as certainly descended from the same progenitor, as have the two sexes of any one species. Consequently, whatever part of the structure of the common progenitor, or of its early descendants, became variable; variations of this part would, it is highly probable, be taken advantage of by natural and sexual selection, in order to fit the several places in the economy of nature, and likewise to fit the two sexes of the same species to each other, or to fit the males to struggle with other males for the possession of the females.</p>
+			<p>It is a remarkable fact, that the secondary differences between the two sexes of the same species are generally displayed in the very same parts of the organisation in which the species of the same genus differ from each other. Of this fact I will give in illustration the first two instances which happen to stand on my list; and as the differences in these cases are of a very unusual nature, the relation can hardly be accidental. The same number of joints in the tarsi is a character common to very large groups of beetles, but in the Engidae, as Westwood has remarked, the number varies greatly and the number likewise differs in the two sexes of the same species. Again in the fossorial hymenoptera, the neuration of the wings is a character of the highest importance, because common to large groups; but in certain genera the neuration differs in the different species, and likewise in the two sexes of the same species. Sir <abbr class="name">J.</abbr> Lubbock has recently remarked, that several minute crustaceans offer excellent illustrations of this law. “In Pontella, for instance, the sexual characters are afforded mainly by the anterior antennae and by the fifth pair of legs: the specific differences also are principally given by these organs.” This relation has a clear meaning on my view: I look at all the species of the same genus as having as certainly descended from the same progenitor, as have the two sexes of any one species. Consequently, whatever part of the structure of the common progenitor, or of its early descendants, became variable; variations of this part would, it is highly probable, be taken advantage of by natural and sexual selection, in order to fit the several places in the economy of nature, and likewise to fit the two sexes of the same species to each other, or to fit the males to struggle with other males for the possession of the females.</p>
 			<p>Finally, then, I conclude that the greater variability of specific characters, or those which distinguish species from species, than of generic characters, or those which are possessed by all the species; that the frequent extreme variability of any part which is developed in a species in an extraordinary manner in comparison with the same part in its congeners; and the slight degree of variability in a part, however extraordinarily it may be developed, if it be common to a whole group of species; that the great variability of secondary sexual characters and their great difference in closely allied species; that secondary sexual and ordinary specific differences are generally displayed in the same parts of the organisation, are all principles closely connected together. All being mainly due to the species of the same group being the descendants of a common progenitor, from whom they have inherited much in common, to parts which have recently and largely varied being more likely still to go on varying than parts which have long been inherited and have not varied, to natural selection having more or less completely, according to the lapse of time, overmastered the tendency to reversion and to further variability, to sexual selection being less rigid than ordinary selection, and to variations in the same parts having been accumulated by natural and sexual selection, and thus having been adapted for secondary sexual, and for ordinary purposes.</p>
 		</section>
 		<section id="chapter-5-9" epub:type="z3998:subchapter">

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@@ -27,7 +27,7 @@
 			<p>But, as by this theory innumerable transitional forms must have existed, why do we not find them embedded in countless numbers in the crust of the earth? It will be more convenient to discuss this question in the chapter on the imperfection of the geological record; and I will here only state that I believe the answer mainly lies in the record being incomparably less perfect than is generally supposed. The crust of the earth is a vast museum; but the natural collections have been imperfectly made, and only at long intervals of time.</p>
 			<p>But it may be urged that when several closely allied species inhabit the same territory, we surely ought to find at the present time many transitional forms. Let us take a simple case: in travelling from north to south over a continent, we generally meet at successive intervals with closely allied or representative species, evidently filling nearly the same place in the natural economy of the land. These representative species often meet and interlock; and as the one becomes rarer and rarer, the other becomes more and more frequent, till the one replaces the other. But if we compare these species where they intermingle, they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis inhabited by each. By my theory these allied species are descended from a common parent; and during the process of modification, each has become adapted to the conditions of life of its own region, and has supplanted and exterminated its original parent-form and all the transitional varieties between its past and present states. Hence we ought not to expect at the present time to meet with numerous transitional varieties in each region, though they must have existed there, and may be embedded there in a fossil condition. But in the intermediate region, having intermediate conditions of life, why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. But I think it can be in large part explained.</p>
 			<p>In the first place we should be extremely cautious in inferring, because an area is now continuous, that it has been continuous during a long period. Geology would lead us to believe that most continents have been broken up into islands even during the later tertiary periods; and in such islands distinct species might have been separately formed without the possibility of intermediate varieties existing in the intermediate zones. By changes in the form of the land and of climate, marine areas now continuous must often have existed within recent times in a far less continuous and uniform condition than at present. But I will pass over this way of escaping from the difficulty; for I believe that many perfectly defined species have been formed on strictly continuous areas; though I do not doubt that the formerly broken condition of areas now continuous, has played an important part in the formation of new species, more especially with freely-crossing and wandering animals.</p>
-			<p>In looking at species as they are now distributed over a wide area, we generally find them tolerably numerous over a large territory, then becoming somewhat abruptly rarer and rarer on the confines, and finally disappearing. Hence the neutral territory between two representative species is generally narrow in comparison with the territory proper to each. We see the same fact in ascending mountains, and sometimes it is quite remarkable how abruptly, as Alph. De Candolle has observed, a common alpine species disappears. The same fact has been noticed by E. Forbes in sounding the depths of the sea with the dredge. To those who look at climate and the physical conditions of life as the all-important elements of distribution, these facts ought to cause surprise, as climate and height or depth graduate away insensibly. But when we bear in mind that almost every species, even in its metropolis, would increase immensely in numbers, were it not for other competing species; that nearly all either prey on or serve as prey for others; in short, that each organic being is either directly or indirectly related in the most important manner to other organic beings⁠—we see that the range of the inhabitants of any country by no means exclusively depends on insensibly changing physical conditions, but in large part on the presence of other species, on which it lives, or by which it is destroyed, or with which it comes into competition; and as these species are already defined objects, not blending one into another by insensible gradations, the range of any one species, depending as it does on the range of others, will tend to be sharply defined. Moreover, each species on the confines of its range, where it exists in lessened numbers, will, during fluctuations in the number of its enemies or of its prey, or in the nature of the seasons, be extremely liable to utter extermination; and thus its geographical range will come to be still more sharply defined.</p>
+			<p>In looking at species as they are now distributed over a wide area, we generally find them tolerably numerous over a large territory, then becoming somewhat abruptly rarer and rarer on the confines, and finally disappearing. Hence the neutral territory between two representative species is generally narrow in comparison with the territory proper to each. We see the same fact in ascending mountains, and sometimes it is quite remarkable how abruptly, as Alph. De Candolle has observed, a common alpine species disappears. The same fact has been noticed by <abbr class="name">E.</abbr> Forbes in sounding the depths of the sea with the dredge. To those who look at climate and the physical conditions of life as the all-important elements of distribution, these facts ought to cause surprise, as climate and height or depth graduate away insensibly. But when we bear in mind that almost every species, even in its metropolis, would increase immensely in numbers, were it not for other competing species; that nearly all either prey on or serve as prey for others; in short, that each organic being is either directly or indirectly related in the most important manner to other organic beings⁠—we see that the range of the inhabitants of any country by no means exclusively depends on insensibly changing physical conditions, but in large part on the presence of other species, on which it lives, or by which it is destroyed, or with which it comes into competition; and as these species are already defined objects, not blending one into another by insensible gradations, the range of any one species, depending as it does on the range of others, will tend to be sharply defined. Moreover, each species on the confines of its range, where it exists in lessened numbers, will, during fluctuations in the number of its enemies or of its prey, or in the nature of the seasons, be extremely liable to utter extermination; and thus its geographical range will come to be still more sharply defined.</p>
 			<p>As allied or representative species, when inhabiting a continuous area, are generally distributed in such a manner that each has a wide range, with a comparatively narrow neutral territory between them, in which they become rather suddenly rarer and rarer; then, as varieties do not essentially differ from species, the same rule will probably apply to both; and if we take a varying species inhabiting a very large area, we shall have to adapt two varieties to two large areas, and a third variety to a narrow intermediate zone. The intermediate variety, consequently, will exist in lesser numbers from inhabiting a narrow and lesser area; and practically, as far as I can make out, this rule holds good with varieties in a state of nature. I have met with striking instances of the rule in the case of varieties intermediate between well-marked varieties in the genus Balanus. And it would appear from information given me by <abbr>Mr.</abbr> Watson, <abbr>Dr.</abbr> Asa Gray, and <abbr>Mr.</abbr> Wollaston, that generally, when varieties intermediate between two other forms occur, they are much rarer numerically than the forms which they connect. Now, if we may trust these facts and inferences, and conclude that varieties linking two other varieties together generally have existed in lesser numbers than the forms which they connect, then we can understand why intermediate varieties should not endure for very long periods: why, as a general rule, they should be exterminated and disappear, sooner than the forms which they originally linked together.</p>
 			<p>For any form existing in lesser numbers would, as already remarked, run a greater chance of being exterminated than one existing in large numbers; and in this particular case the intermediate form would be eminently liable to the inroads of closely allied forms existing on both sides of it. But it is a far more important consideration, that during the process of further modification, by which two varieties are supposed to be converted and perfected into two distinct species, the two which exist in larger numbers, from inhabiting larger areas, will have a great advantage over the intermediate variety, which exists in smaller numbers in a narrow and intermediate zone. For forms existing in larger numbers will have a better chance, within any given period, of presenting further favourable variations for natural selection to seize on, than will the rarer forms which exist in lesser numbers. Hence, the more common forms, in the race for life, will tend to beat and supplant the less common forms, for these will be more slowly modified and improved. It is the same principle which, as I believe, accounts for the common species in each country, as shown in the second chapter, presenting on an average a greater number of well-marked varieties than do the rarer species. I may illustrate what I mean by supposing three varieties of sheep to be kept, one adapted to an extensive mountainous region; a second to a comparatively narrow, hilly tract; and a third to the wide plains at the base; and that the inhabitants are all trying with equal steadiness and skill to improve their stocks by selection; the chances in this case will be strongly in favour of the great holders on the mountains or on the plains improving their breeds more quickly than the small holders on the intermediate narrow, hilly tract; and consequently the improved mountain or plain breed will soon take the place of the less improved hill breed; and thus the two breeds, which originally existed in greater numbers, will come into close contact with each other, without the interposition of the supplanted, intermediate hill variety.</p>
 			<p>To sum up, I believe that species come to be tolerably well-defined objects, and do not at any one period present an inextricable chaos of varying and intermediate links: first, because new varieties are very slowly formed, for variation is a slow process, and natural selection can do nothing until favourable individual differences or variations occur, and until a place in the natural polity of the country can be better filled by some modification of some one or more of its inhabitants. And such new places will depend on slow changes of climate, or on the occasional immigration of new inhabitants, and, probably, in a still more important degree, on some of the old inhabitants becoming slowly modified, with the new forms thus produced and the old ones acting and reacting on each other. So that, in any one region and at any one time, we ought to see only a few species presenting slight modifications of structure in some degree permanent; and this assuredly we do see.</p>
@@ -39,7 +39,7 @@
 			<h3 epub:type="title">On the Origin and Transition of Organic Beings with Peculiar Habits and Structure</h3>
 			<p>It has been asked by the opponents of such views as I hold, how, for instance, could a land carnivorous animal have been converted into one with aquatic habits; for how could the animal in its transitional state have subsisted? It would be easy to show that there now exist carnivorous animals presenting close intermediate grades from strictly terrestrial to aquatic habits; and as each exists by a struggle for life, it is clear that each must be well adapted to its place in nature. Look at the Mustela vison of North America, which has webbed feet, and which resembles an otter in its fur, short legs, and form of tail; during summer this animal dives for and preys on fish, but during the long winter it leaves the frozen waters, and preys, like other polecats on mice and land animals. If a different case had been taken, and it had been asked how an insectivorous quadruped could possibly have been converted into a flying bat, the question would have been far more difficult to answer. Yet I think such difficulties have little weight.</p>
 			<p>Here, as on other occasions, I lie under a heavy disadvantage, for, out of the many striking cases which I have collected, I can give only one or two instances of transitional habits and structures in allied species; and of diversified habits, either constant or occasional, in the same species. And it seems to me that nothing less than a long list of such cases is sufficient to lessen the difficulty in any particular case like that of the bat.</p>
-			<p>Look at the family of squirrels; here we have the finest gradation from animals with their tails only slightly flattened, and from others, as Sir J. Richardson has remarked, with the posterior part of their bodies rather wide and with the skin on their flanks rather full, to the so-called flying squirrels; and flying squirrels have their limbs and even the base of the tail united by a broad expanse of skin, which serves as a parachute and allows them to glide through the air to an astonishing distance from tree to tree. We cannot doubt that each structure is of use to each kind of squirrel in its own country, by enabling it to escape birds or beasts of prey, or to collect food more quickly, or, as there is reason to believe, to lessen the danger from occasional falls. But it does not follow from this fact that the structure of each squirrel is the best that it is possible to conceive under all possible conditions. Let the climate and vegetation change, let other competing rodents or new beasts of prey immigrate, or old ones become modified, and all analogy would lead us to believe that some, at least, of the squirrels would decrease in numbers or become exterminated, unless they also become modified and improved in structure in a corresponding manner. Therefore, I can see no difficulty, more especially under changing conditions of life, in the continued preservation of individuals with fuller and fuller flank-membranes, each modification being useful, each being propagated, until, by the accumulated effects of this process of natural selection, a perfect so-called flying squirrel was produced.</p>
+			<p>Look at the family of squirrels; here we have the finest gradation from animals with their tails only slightly flattened, and from others, as Sir <abbr class="name">J.</abbr> Richardson has remarked, with the posterior part of their bodies rather wide and with the skin on their flanks rather full, to the so-called flying squirrels; and flying squirrels have their limbs and even the base of the tail united by a broad expanse of skin, which serves as a parachute and allows them to glide through the air to an astonishing distance from tree to tree. We cannot doubt that each structure is of use to each kind of squirrel in its own country, by enabling it to escape birds or beasts of prey, or to collect food more quickly, or, as there is reason to believe, to lessen the danger from occasional falls. But it does not follow from this fact that the structure of each squirrel is the best that it is possible to conceive under all possible conditions. Let the climate and vegetation change, let other competing rodents or new beasts of prey immigrate, or old ones become modified, and all analogy would lead us to believe that some, at least, of the squirrels would decrease in numbers or become exterminated, unless they also become modified and improved in structure in a corresponding manner. Therefore, I can see no difficulty, more especially under changing conditions of life, in the continued preservation of individuals with fuller and fuller flank-membranes, each modification being useful, each being propagated, until, by the accumulated effects of this process of natural selection, a perfect so-called flying squirrel was produced.</p>
 			<p>Now look at the Galeopithecus or so-called flying lemur, which was formerly ranked among bats, but is now believed to belong to the Insectivora. An extremely wide flank-membrane stretches from the corners of the jaw to the tail, and includes the limbs with the elongated fingers. This flank-membrane is furnished with an extensor muscle. Although no graduated links of structure, fitted for gliding through the air, now connect the Galeopithecus with the other Insectivora, yet there is no difficulty in supposing that such links formerly existed, and that each was developed in the same manner as with the less perfectly gliding squirrels; each grade of structure having been useful to its possessor. Nor can I see any insuperable difficulty in further believing it possible that the membrane-connected fingers and forearm of the Galeopithecus might have been greatly lengthened by natural selection; and this, as far as the organs of flight are concerned, would have converted the animal into a bat. In certain bats in which the wing-membrane extends from the top of the shoulder to the tail and includes the hind-legs, we perhaps see traces of an apparatus originally fitted for gliding through the air rather than for flight.</p>
 			<p>If about a dozen genera of birds were to become extinct, who would have ventured to surmise that birds might have existed which used their wings solely as flappers, like the logger headed duck (Micropterus of Eyton); as fins in the water and as front legs on the land, like the penguin; as sails, like the ostrich; and functionally for no purpose, like the apteryx? Yet the structure of each of these birds is good for it, under the conditions of life to which it is exposed, for each has to live by a struggle: but it is not necessarily the best possible under all possible conditions. It must not be inferred from these remarks that any of the grades of wing-structure here alluded to, which perhaps may all be the result of disuse, indicate the steps by which birds actually acquired their perfect power of flight; but they serve to show what diversified means of transition are at least possible.</p>
 			<p>Seeing that a few members of such water-breathing classes as the Crustacea and Mollusca are adapted to live on the land; and seeing that we have flying birds and mammals, flying insects of the most diversified types, and formerly had flying reptiles, it is conceivable that flying-fish, which now glide far through the air, slightly rising and turning by the aid of their fluttering fins, might have been modified into perfectly winged animals. If this had been effected, who would have ever imagined that in an early transitional state they had been inhabitants of the open ocean, and had used their incipient organs of flight exclusively, so far as we know, to escape being devoured by other fish?</p>
@@ -54,7 +54,7 @@
 			<h3 epub:type="title">Organs of Extreme Perfection and Complication</h3>
 			<p>To suppose that the eye with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest degree. When it was first said that the sun stood still and the world turned round, the common sense of mankind declared the doctrine false; but the old saying of Vox populi, vox Dei, as every philosopher knows, cannot be trusted in science. Reason tells me, that if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist, each grade being useful to its possessor, as is certainly the case; if further, the eye ever varies and the variations be inherited, as is likewise certainly the case; and if such variations should be useful to any animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, should not be considered as subversive of the theory. How a nerve comes to be sensitive to light, hardly concerns us more than how life itself originated; but I may remark that, as some of the lowest organisms in which nerves cannot be detected, are capable of perceiving light, it does not seem impossible that certain sensitive elements in their sarcode should become aggregated and developed into nerves, endowed with this special sensibility.</p>
 			<p>In searching for the gradations through which an organ in any species has been perfected, we ought to look exclusively to its lineal progenitors; but this is scarcely ever possible, and we are forced to look to other species and genera of the same group, that is to the collateral descendants from the same parent-form, in order to see what gradations are possible, and for the chance of some gradations having been transmitted in an unaltered or little altered condition. But the state of the same organ in distinct classes may incidentally throw light on the steps by which it has been perfected.</p>
-			<p>The simplest organ which can be called an eye consists of an optic nerve, surrounded by pigment-cells and covered by translucent skin, but without any lens or other refractive body. We may, however, according to M. Jourdain, descend even a step lower and find aggregates of pigment-cells, apparently serving as organs of vision, without any nerves, and resting merely on sarcodic tissue. Eyes of the above simple nature are not capable of distinct vision, and serve only to distinguish light from darkness. In certain starfishes, small depressions in the layer of pigment which surrounds the nerve are filled, as described by the author just quoted, with transparent gelatinous matter, projecting with a convex surface, like the cornea in the higher animals. He suggests that this serves not to form an image, but only to concentrate the luminous rays and render their perception more easy. In this concentration of the rays we gain the first and by far the most important step towards the formation of a true, picture-forming eye; for we have only to place the naked extremity of the optic nerve, which in some of the lower animals lies deeply buried in the body, and in some near the surface, at the right distance from the concentrating apparatus, and an image will be formed on it.</p>
+			<p>The simplest organ which can be called an eye consists of an optic nerve, surrounded by pigment-cells and covered by translucent skin, but without any lens or other refractive body. We may, however, according to <abbr class="name">M.</abbr> Jourdain, descend even a step lower and find aggregates of pigment-cells, apparently serving as organs of vision, without any nerves, and resting merely on sarcodic tissue. Eyes of the above simple nature are not capable of distinct vision, and serve only to distinguish light from darkness. In certain starfishes, small depressions in the layer of pigment which surrounds the nerve are filled, as described by the author just quoted, with transparent gelatinous matter, projecting with a convex surface, like the cornea in the higher animals. He suggests that this serves not to form an image, but only to concentrate the luminous rays and render their perception more easy. In this concentration of the rays we gain the first and by far the most important step towards the formation of a true, picture-forming eye; for we have only to place the naked extremity of the optic nerve, which in some of the lower animals lies deeply buried in the body, and in some near the surface, at the right distance from the concentrating apparatus, and an image will be formed on it.</p>
 			<p>In the great class of the Articulata, we may start from an optic nerve simply coated with pigment, the latter sometimes forming a sort of pupil, but destitute of lens or other optical contrivance. With insects it is now known that the numerous facets on the cornea of their great compound eyes form true lenses, and that the cones include curiously modified nervous filaments. But these organs in the Articulata are so much diversified that Muller formerly made three main classes with seven subdivisions, besides a fourth main class of aggregated simple eyes.</p>
 			<p>When we reflect on these facts, here given much too briefly, with respect to the wide, diversified, and graduated range of structure in the eyes of the lower animals; and when we bear in mind how small the number of all living forms must be in comparison with those which have become extinct, the difficulty ceases to be very great in believing that natural selection may have converted the simple apparatus of an optic nerve, coated with pigment and invested by transparent membrane, into an optical instrument as perfect as is possessed by any member of the Articulata class.</p>
 			<p>He who will go thus far, ought not to hesitate to go one step further, if he finds on finishing this volume that large bodies of facts, otherwise inexplicable, can be explained by the theory of modification through natural selection; he ought to admit that a structure even as perfect as an eagle’s eye might thus be formed, although in this case he does not know the transitional states. It has been objected that in order to modify the eye and still preserve it as a perfect instrument, many changes would have to be effected simultaneously, which, it is assumed, could not be done through natural selection; but as I have attempted to show in my work on the variation of domestic animals, it is not necessary to suppose that the modifications were all simultaneous, if they were extremely slight and gradual. Different kinds of modification would, also, serve for the same general purpose: as <abbr>Mr.</abbr> Wallace has remarked, “If a lens has too short or too long a focus, it may be amended either by an alteration of curvature, or an alteration of density; if the curvature be irregular, and the rays do not converge to a point, then any increased regularity of curvature will be an improvement. So the contraction of the iris and the muscular movements of the eye are neither of them essential to vision, but only improvements which might have been added and perfected at any stage of the construction of the instrument.” Within the highest division of the animal kingdom, namely, the Vertebrata, we can start from an eye so simple, that it consists, as in the lancelet, of a little sack of transparent skin, furnished with a nerve and lined with pigment, but destitute of any other apparatus. In fishes and reptiles, as Owen has remarked, “The range of gradation of dioptric structures is very great.” It is a significant fact that even in man, according to the high authority of Virchow, the beautiful crystalline lens is formed in the embryo by an accumulation of epidermic cells, lying in a sack-like fold of the skin; and the vitreous body is formed from embryonic subcutaneous tissue. To arrive, however, at a just conclusion regarding the formation of the eye, with all its marvellous yet not absolutely perfect characters, it is indispensable that the reason should conquer the imagination; but I have felt the difficulty far to keenly to be surprised at others hesitating to extend the principle of natural selection to so startling a length.</p>
@@ -92,7 +92,7 @@
 			<p>Organs now of trifling importance have probably in some cases been of high importance to an early progenitor, and, after having been slowly perfected at a former period, have been transmitted to existing species in nearly the same state, although now of very slight use; but any actually injurious deviations in their structure would of course have been checked by natural selection. Seeing how important an organ of locomotion the tail is in most aquatic animals, its general presence and use for many purposes in so many land animals, which in their lungs or modified swim-bladders betray their aquatic origin, may perhaps be thus accounted for. A well-developed tail having been formed in an aquatic animal, it might subsequently come to be worked in for all sorts of purposes, as a fly-flapper, an organ of prehension, or as an aid in turning, as in the case of the dog, though the aid in this latter respect must be slight, for the hare, with hardly any tail, can double still more quickly.</p>
 			<p>In the second place, we may easily err in attributing importance to characters, and in believing that they have been developed through natural selection. We must by no means overlook the effects of the definite action of changed conditions of life, of so-called spontaneous variations, which seem to depend in a quite subordinate degree on the nature of the conditions, of the tendency to reversion to long-lost characters, of the complex laws of growth, such as of correlation, comprehension, of the pressure of one part on another, <abbr>etc.</abbr>, and finally of sexual selection, by which characters of use to one sex are often gained and then transmitted more or less perfectly to the other sex, though of no use to the sex. But structures thus indirectly gained, although at first of no advantage to a species, may subsequently have been taken advantage of by its modified descendants, under new conditions of life and newly acquired habits.</p>
 			<p>If green woodpeckers alone had existed, and we did not know that there were many black and pied kinds, I dare say that we should have thought that the green colour was a beautiful adaptation to conceal this tree-frequenting bird from its enemies; and consequently that it was a character of importance, and had been acquired through natural selection; as it is, the colour is probably in chief part due to sexual selection. A trailing palm in the Malay Archipelago climbs the loftiest trees by the aid of exquisitely constructed hooks clustered around the ends of the branches, and this contrivance, no doubt, is of the highest service to the plant; but as we see nearly similar hooks on many trees which are not climbers, and which, as there is reason to believe from the distribution of the thorn-bearing species in Africa and South America, serve as a defence against browsing quadrupeds, so the spikes on the palm may at first have been developed for this object, and subsequently have been improved and taken advantage of by the plant, as it underwent further modification and became a climber. The naked skin on the head of a vulture is generally considered as a direct adaptation for wallowing in putridity; and so it may be, or it may possibly be due to the direct action of putrid matter; but we should be very cautious in drawing any such inference, when we see that the skin on the head of the clean-feeding male turkey is likewise naked. The sutures in the skulls of young mammals have been advanced as a beautiful adaptation for aiding parturition, and no doubt they facilitate, or may be indispensable for this act; but as sutures occur in the skulls of young birds and reptiles, which have only to escape from a broken egg, we may infer that this structure has arisen from the laws of growth, and has been taken advantage of in the parturition of the higher animals.</p>
-			<p>We are profoundly ignorant of the cause of each slight variation or individual difference; and we are immediately made conscious of this by reflecting on the differences between the breeds of our domesticated animals in different countries, more especially in the less civilized countries, where there has been but little methodical selection. Animals kept by savages in different countries often have to struggle for their own subsistence, and are exposed to a certain extent to natural selection, and individuals with slightly different constitutions would succeed best under different climates. With cattle susceptibility to the attacks of flies is correlated with colour, as is the liability to be poisoned by certain plants; so that even colour would be thus subjected to the action of natural selection. Some observers are convinced that a damp climate affects the growth of the hair, and that with the hair the horns are correlated. Mountain breeds always differ from lowland breeds; and a mountainous country would probably affect the hind limbs from exercising them more, and possibly even the form of the pelvis; and then by the law of homologous variation, the front limbs and the head would probably be affected. The shape, also, of the pelvis might affect by pressure the shape of certain parts of the young in the womb. The laborious breathing necessary in high regions tends, as we have good reason to believe, to increase the size of the chest; and again correlation would come into play. The effects of lessened exercise, together with abundant food, on the whole organisation is probably still more important, and this, as H. von Nathusius has lately shown in his excellent Treatise, is apparently one chief cause of the great modification which the breeds of swine have undergone. But we are far too ignorant to speculate on the relative importance of the several known and unknown causes of variation; and I have made these remarks only to show that, if we are unable to account for the characteristic differences of our several domestic breeds, which nevertheless are generally admitted to have arisen through ordinary generation from one or a few parent-stocks, we ought not to lay too much stress on our ignorance of the precise cause of the slight analogous differences between true species.</p>
+			<p>We are profoundly ignorant of the cause of each slight variation or individual difference; and we are immediately made conscious of this by reflecting on the differences between the breeds of our domesticated animals in different countries, more especially in the less civilized countries, where there has been but little methodical selection. Animals kept by savages in different countries often have to struggle for their own subsistence, and are exposed to a certain extent to natural selection, and individuals with slightly different constitutions would succeed best under different climates. With cattle susceptibility to the attacks of flies is correlated with colour, as is the liability to be poisoned by certain plants; so that even colour would be thus subjected to the action of natural selection. Some observers are convinced that a damp climate affects the growth of the hair, and that with the hair the horns are correlated. Mountain breeds always differ from lowland breeds; and a mountainous country would probably affect the hind limbs from exercising them more, and possibly even the form of the pelvis; and then by the law of homologous variation, the front limbs and the head would probably be affected. The shape, also, of the pelvis might affect by pressure the shape of certain parts of the young in the womb. The laborious breathing necessary in high regions tends, as we have good reason to believe, to increase the size of the chest; and again correlation would come into play. The effects of lessened exercise, together with abundant food, on the whole organisation is probably still more important, and this, as <abbr class="name">H.</abbr> von Nathusius has lately shown in his excellent Treatise, is apparently one chief cause of the great modification which the breeds of swine have undergone. But we are far too ignorant to speculate on the relative importance of the several known and unknown causes of variation; and I have made these remarks only to show that, if we are unable to account for the characteristic differences of our several domestic breeds, which nevertheless are generally admitted to have arisen through ordinary generation from one or a few parent-stocks, we ought not to lay too much stress on our ignorance of the precise cause of the slight analogous differences between true species.</p>
 		</section>
 		<section id="chapter-6-7" epub:type="z3998:subchapter">
 			<h3 epub:type="title">Utilitarian Doctrine, How Far True: Beauty, How Acquired</h3>

src/epub/text/chapter-7.xhtml

@@ -25,9 +25,9 @@
 			<p>It may be worth while to illustrate some of the foregoing remarks. With respect to the assumed inutility of various parts and organs, it is hardly necessary to observe that even in the higher and best-known animals many structures exist, which are so highly developed that no one doubts that they are of importance, yet their use has not been, or has only recently been, ascertained. As Bronn gives the length of the ears and tail in the several species of mice as instances, though trifling ones, of differences in structure which can be of no special use, I may mention that, according to <abbr>Dr.</abbr> Schobl, the external ears of the common mouse are supplied in an extraordinary manner with nerves, so that they no doubt serve as tactile organs; hence the length of the ears can hardly be quite unimportant. We shall, also, presently see that the tail is a highly useful prehensile organ to some of the species; and its use would be much influence by its length.</p>
 			<p>With respect to plants, to which on account of Nageli’s essay I shall confine myself in the following remarks, it will be admitted that the flowers of the orchids present a multitude of curious structures, which a few years ago would have been considered as mere morphological differences without any special function; but they are now known to be of the highest importance for the fertilisation of the species through the aid of insects, and have probably been gained through natural selection. No one until lately would have imagined that in dimorphic and trimorphic plants the different lengths of the stamens and pistils, and their arrangement, could have been of any service, but now we know this to be the case.</p>
 			<p>In certain whole groups of plants the ovules stand erect, and in others they are suspended; and within the same ovarium of some few plants, one ovule holds the former and a second ovule the latter position. These positions seem at first purely morphological, or of no physiological signification; but <abbr>Dr.</abbr> Hooker informs me that within the same ovarium the upper ovules alone in some cases, and in others the lower ones alone are fertilised; and he suggests that this probably depends on the direction in which the pollen-tubes enter the ovarium. If so, the position of the ovules, even when one is erect and the other suspended within the same ovarium, would follow the selection of any slight deviations in position which favoured their fertilisation, and the production of seed.</p>
-			<p>Several plants belonging to distinct orders habitually produce flowers of two kinds⁠—the one open, of the ordinary structure, the other closed and imperfect. These two kinds of flowers sometimes differ wonderfully in structure, yet may be seen to graduate into each other on the same plant. The ordinary and open flowers can be intercrossed; and the benefits which certainly are derived from this process are thus secured. The closed and imperfect flowers are, however, manifestly of high importance, as they yield with the utmost safety a large stock of seed, with the expenditure of wonderfully little pollen. The two kinds of flowers often differ much, as just stated, in structure. The petals in the imperfect flowers almost always consist of mere rudiments, and the pollen-grains are reduced in diameter. In Ononis columnae five of the alternate stamens are rudimentary; and in some species of Viola three stamens are in this state, two retaining their proper function, but being of very small size. In six out of thirty of the closed flowers in an Indian violet (name unknown, for the plants have never produced with me perfect flowers), the sepals are reduced from the normal number of five to three. In one section of the Malpighiaceae the closed flowers, according to A. de Jussieu, are still further modified, for the five stamens which stand opposite to the sepals are all aborted, a sixth stamen standing opposite to a petal being alone developed; and this stamen is not present in the ordinary flowers of this species; the style is aborted; and the ovaria are reduced from three to two. Now although natural selection may well have had the power to prevent some of the flowers from expanding, and to reduce the amount of pollen, when rendered by the closure of the flowers superfluous, yet hardly any of the above special modifications can have been thus determined, but must have followed from the laws of growth, including the functional inactivity of parts, during the progress of the reduction of the pollen and the closure of the flowers.</p>
+			<p>Several plants belonging to distinct orders habitually produce flowers of two kinds⁠—the one open, of the ordinary structure, the other closed and imperfect. These two kinds of flowers sometimes differ wonderfully in structure, yet may be seen to graduate into each other on the same plant. The ordinary and open flowers can be intercrossed; and the benefits which certainly are derived from this process are thus secured. The closed and imperfect flowers are, however, manifestly of high importance, as they yield with the utmost safety a large stock of seed, with the expenditure of wonderfully little pollen. The two kinds of flowers often differ much, as just stated, in structure. The petals in the imperfect flowers almost always consist of mere rudiments, and the pollen-grains are reduced in diameter. In Ononis columnae five of the alternate stamens are rudimentary; and in some species of Viola three stamens are in this state, two retaining their proper function, but being of very small size. In six out of thirty of the closed flowers in an Indian violet (name unknown, for the plants have never produced with me perfect flowers), the sepals are reduced from the normal number of five to three. In one section of the Malpighiaceae the closed flowers, according to <abbr class="name">A.</abbr> de Jussieu, are still further modified, for the five stamens which stand opposite to the sepals are all aborted, a sixth stamen standing opposite to a petal being alone developed; and this stamen is not present in the ordinary flowers of this species; the style is aborted; and the ovaria are reduced from three to two. Now although natural selection may well have had the power to prevent some of the flowers from expanding, and to reduce the amount of pollen, when rendered by the closure of the flowers superfluous, yet hardly any of the above special modifications can have been thus determined, but must have followed from the laws of growth, including the functional inactivity of parts, during the progress of the reduction of the pollen and the closure of the flowers.</p>
 			<p>It is so necessary to appreciate the important effects of the laws of growth, that I will give some additional cases of another kind, namely of differences in the same part or organ, due to differences in relative position on the same plant. In the Spanish chestnut, and in certain fir-trees, the angles of divergence of the leaves differ, according to Schacht, in the nearly horizontal and in the upright branches. In the common rue and some other plants, one flower, usually the central or terminal one, opens first, and has five sepals and petals, and five divisions to the ovarium; while all the other flowers on the plant are tetramerous. In the British Adoxa the uppermost flower generally has two calyx-lobes with the other organs tetramerous, while the surrounding flowers generally have three calyx-lobes with the other organs pentamerous. In many Compositae and Umbelliferae (and in some other plants) the circumferential flowers have their corollas much more developed than those of the centre; and this seems often connected with the abortion of the reproductive organs. It is a more curious fact, previously referred to, that the achenes or seeds of the circumference and centre sometimes differ greatly in form, colour and other characters. In Carthamus and some other Compositae the central achenes alone are furnished with a pappus; and in Hyoseris the same head yields achenes of three different forms. In certain Umbelliferae the exterior seeds, according to Tausch, are orthospermous, and the central one coelospermous, and this is a character which was considered by De Candolle to be in other species of the highest systematic importance. Professor Braun mentions a Fumariaceous genus, in which the flowers in the lower part of the spike bear oval, ribbed, one-seeded nutlets; and in the upper part of the spike, lanceolate, two-valved and two-seeded siliques. In these several cases, with the exception of that of the well-developed ray-florets, which are of service in making the flowers conspicuous to insects, natural selection cannot, as far as we can judge, have come into play, or only in a quite subordinate manner. All these modifications follow from the relative position and interaction of the parts; and it can hardly be doubted that if all the flowers and leaves on the same plant had been subjected to the same external and internal condition, as are the flowers and leaves in certain positions, all would have been modified in the same manner.</p>
-			<p>In numerous other cases we find modifications of structure, which are considered by botanists to be generally of a highly important nature, affecting only some of the flowers on the same plant, or occurring on distinct plants, which grow close together under the same conditions. As these variations seem of no special use to the plants, they cannot have been influenced by natural selection. Of their cause we are quite ignorant; we cannot even attribute them, as in the last class of cases, to any proximate agency, such as relative position. I will give only a few instances. It is so common to observe on the same plant, flowers indifferently tetramerous, pentamerous, <abbr>etc.</abbr>, that I need not give examples; but as numerical variations are comparatively rare when the parts are few, I may mention that, according to De Candolle, the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies), or three sepals with six petals. The manner in which the petals are folded in the bud is in most groups a very constant morphological character; but Professor Asa Gray states that with some species of Mimulus, the aestivation is almost as frequently that of the Rhinanthideae as of the Antirrhinideae, to which latter tribe the genus belongs. <abbr>Aug.</abbr> <abbr class="name">St.</abbr> Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutaceae with a single ovary, but in some species flowers may be found on the same plant, and even in the same panicle, with either one or two ovaries. In Helianthemum the capsule has been described as unilocular or tri-locular; and in H. mutabile, “<i xml:lang="fr">Une lame <em>plus ou moins large</em>, s’etend entre le pericarpe et le placenta.</i>” In the flowers of Saponaria officinalis <abbr>Dr.</abbr> Masters has observed instances of both marginal and free central placentation. Lastly, <abbr class="name">St.</abbr> Hilaire found towards the southern extreme of the range of Gomphia oleaeformis two forms which he did not at first doubt were distinct species, but he subsequently saw them growing on the same bush; and he then adds, “Voila donc dans un meme individu des loges et un style qui se rattachent tantot a un axe verticale et tantot a un gynobase.”</p>
+			<p>In numerous other cases we find modifications of structure, which are considered by botanists to be generally of a highly important nature, affecting only some of the flowers on the same plant, or occurring on distinct plants, which grow close together under the same conditions. As these variations seem of no special use to the plants, they cannot have been influenced by natural selection. Of their cause we are quite ignorant; we cannot even attribute them, as in the last class of cases, to any proximate agency, such as relative position. I will give only a few instances. It is so common to observe on the same plant, flowers indifferently tetramerous, pentamerous, <abbr>etc.</abbr>, that I need not give examples; but as numerical variations are comparatively rare when the parts are few, I may mention that, according to De Candolle, the flowers of Papaver bracteatum offer either two sepals with four petals (which is the common type with poppies), or three sepals with six petals. The manner in which the petals are folded in the bud is in most groups a very constant morphological character; but Professor Asa Gray states that with some species of Mimulus, the aestivation is almost as frequently that of the Rhinanthideae as of the Antirrhinideae, to which latter tribe the genus belongs. <abbr>Aug.</abbr> <abbr class="name">St.</abbr> Hilaire gives the following cases: the genus Zanthoxylon belongs to a division of the Rutaceae with a single ovary, but in some species flowers may be found on the same plant, and even in the same panicle, with either one or two ovaries. In Helianthemum the capsule has been described as unilocular or tri-locular; and in <i epub:type="z3998:taxonomy">H. mutabile</i>, “<i xml:lang="fr">Une lame <em>plus ou moins large</em>, s’etend entre le pericarpe et le placenta.</i>” In the flowers of <i epub:type="z3998:taxonomy">Saponaria officinalis</i> <abbr>Dr.</abbr> Masters has observed instances of both marginal and free central placentation. Lastly, <abbr class="name">St.</abbr> Hilaire found towards the southern extreme of the range of <i epub:type="z3998:taxonomy">Gomphia oleaeformis</i> two forms which he did not at first doubt were distinct species, but he subsequently saw them growing on the same bush; and he then adds, “Voila donc dans un meme individu des loges et un style qui se rattachent tantot a un axe verticale et tantot a un gynobase.”</p>
 			<p>We thus see that with plants many morphological changes may be attributed to the laws of growth and the interaction of parts, independently of natural selection. But with respect to Nageli’s doctrine of an innate tendency towards perfection or progressive development, can it be said in the case of these strongly pronounced variations, that the plants have been caught in the act of progressing towards a higher state of development? On the contrary, I should infer from the mere fact of the parts in question differing or varying greatly on the same plant, that such modifications were of extremely small importance to the plants themselves, of whatever importance they may generally be to us for our classifications. The acquisition of a useless part can hardly be said to raise an organism in the natural scale; and in the case of the imperfect, closed flowers, above described, if any new principle has to be invoked, it must be one of retrogression rather than of progression; and so it must be with many parasitic and degraded animals. We are ignorant of the exciting cause of the above specified modifications; but if the unknown cause were to act almost uniformly for a length of time, we may infer that the result would be almost uniform; and in this case all the individuals of the species would be modified in the same manner.</p>
 			<p>From the fact of the above characters being unimportant for the welfare of the species, any slight variations which occurred in them would not have been accumulated and augmented through natural selection. A structure which has been developed through long-continued selection, when it ceases to be of service to a species, generally becomes variable, as we see with rudimentary organs; for it will no longer be regulated by this same power of selection. But when, from the nature of the organism and of the conditions, modifications have been induced which are unimportant for the welfare of the species, they may be, and apparently often have been, transmitted in nearly the same state to numerous, otherwise modified, descendants. It cannot have been of much importance to the greater number of mammals, birds, or reptiles, whether they were clothed with hair, feathers or scales; yet hair has been transmitted to almost all mammals, feathers to all birds, and scales to all true reptiles. A structure, whatever it may be, which is common to many allied forms, is ranked by us as of high systematic importance, and consequently is often assumed to be of high vital importance to the species. Thus, as I am inclined to believe, morphological differences, which we consider as important⁠—such as the arrangement of the leaves, the divisions of the flower or of the ovarium, the position of the ovules, <abbr>etc.</abbr>, first appeared in many cases as fluctuating variations, which sooner or later became constant through the nature of the organism and of the surrounding conditions, as well as through the intercrossing of distinct individuals, but not through natural selection; for as these morphological characters do not affect the welfare of the species, any slight deviations in them could not have been governed or accumulated through this latter agency. It is a strange result which we thus arrive at, namely, that characters of slight vital importance to the species, are the most important to the systematist; but, as we shall hereafter see when we treat of the genetic principle of classification, this is by no means so paradoxical as it may at first appear.</p>
 			<p>Although we have no good evidence of the existence in organic beings of an innate tendency towards progressive development, yet this necessarily follows, as I have attempted to show in the fourth chapter, through the continued action of natural selection. For the best definition which has ever been given of a high standard of organisation, is the degree to which the parts have been specialised or differentiated; and natural selection tends towards this end, inasmuch as the parts are thus enabled to perform their functions more efficiently.</p>
@@ -35,11 +35,11 @@
 			<p>All <abbr>Mr.</abbr> Mivart’s objections will be, or have been, considered in the present volume. The one new point which appears to have struck many readers is, “That natural selection is incompetent to account for the incipient stages of useful structures.” This subject is intimately connected with that of the gradation of the characters, often accompanied by a change of function, for instance, the conversion of a swim-bladder into lungs, points which were discussed in the last chapter under two headings. Nevertheless, I will here consider in some detail several of the cases advanced by <abbr>Mr.</abbr> Mivart, selecting those which are the most illustrative, as want of space prevents me from considering all.</p>
 			<p>The giraffe, by its lofty stature, much elongated neck, fore legs, head and tongue, has its whole frame beautifully adapted for browsing on the higher branches of trees. It can thus obtain food beyond the reach of the other Ungulata or hoofed animals inhabiting the same country; and this must be a great advantage to it during dearths. The Niata cattle in South America show us how small a difference in structure may make, during such periods, a great difference in preserving an animal’s life. These cattle can browse as well as others on grass, but from the projection of the lower jaw they cannot, during the often recurrent droughts, browse on the twigs of trees, reeds, <abbr>etc.</abbr>, to which food the common cattle and horses are then driven; so that at these times the Niatas perish, if not fed by their owners. Before coming to <abbr>Mr.</abbr> Mivart’s objections, it may be well to explain once again how natural selection will act in all ordinary cases. Man has modified some of his animals, without necessarily having attended to special points of structure, by simply preserving and breeding from the fleetest individuals, as with the racehorse and greyhound, or as with the gamecock, by breeding from the victorious birds. So under nature with the nascent giraffe, the individuals which were the highest browsers and were able during dearths to reach even an inch or two above the others, will often have been preserved; for they will have roamed over the whole country in search of food. That the individuals of the same species often differ slightly in the relative lengths of all their parts may be seen in many works of natural history, in which careful measurements are given. These slight proportional differences, due to the laws of growth and variation, are not of the slightest use or importance to most species. But it will have been otherwise with the nascent giraffe, considering its probable habits of life; for those individuals which had some one part or several parts of their bodies rather more elongated than usual, would generally have survived. These will have intercrossed and left offspring, either inheriting the same bodily peculiarities, or with a tendency to vary again in the same manner; while the individuals less favoured in the same respects will have been the most liable to perish.</p>
 			<p>We here see that there is no need to separate single pairs, as man does, when he methodically improves a breed: natural selection will preserve and thus separate all the superior individuals, allowing them freely to intercross, and will destroy all the inferior individuals. By this process long-continued, which exactly corresponds with what I have called unconscious selection by man, combined, no doubt, in a most important manner with the inherited effects of the increased use of parts, it seems to me almost certain that an ordinary hoofed quadruped might be converted into a giraffe.</p>
-			<p>To this conclusion <abbr>Mr.</abbr> Mivart brings forward two objections. One is that the increased size of the body would obviously require an increased supply of food, and he considers it as “very problematical whether the disadvantages thence arising would not, in times of scarcity, more than counterbalance the advantages.” But as the giraffe does actually exist in large numbers in Africa, and as some of the largest antelopes in the world, taller than an ox, abound there, why should we doubt that, as far as size is concerned, intermediate gradations could formerly have existed there, subjected as now to severe dearths. Assuredly the being able to reach, at each stage of increased size, to a supply of food, left untouched by the other hoofed quadrupeds of the country, would have been of some advantage to the nascent giraffe. Nor must we overlook the fact, that increased bulk would act as a protection against almost all beasts of prey excepting the lion; and against this animal, its tall neck⁠—and the taller the better⁠—would, as <abbr>Mr.</abbr> Chauncey Wright has remarked, serve as a watchtower. It is from this cause, as Sir S. Baker remarks, that no animal is more difficult to stalk than the giraffe. This animal also uses its long neck as a means of offence or defence, by violently swinging its head armed with stump-like horns. The preservation of each species can rarely be determined by any one advantage, but by the union of all, great and small.</p>
+			<p>To this conclusion <abbr>Mr.</abbr> Mivart brings forward two objections. One is that the increased size of the body would obviously require an increased supply of food, and he considers it as “very problematical whether the disadvantages thence arising would not, in times of scarcity, more than counterbalance the advantages.” But as the giraffe does actually exist in large numbers in Africa, and as some of the largest antelopes in the world, taller than an ox, abound there, why should we doubt that, as far as size is concerned, intermediate gradations could formerly have existed there, subjected as now to severe dearths. Assuredly the being able to reach, at each stage of increased size, to a supply of food, left untouched by the other hoofed quadrupeds of the country, would have been of some advantage to the nascent giraffe. Nor must we overlook the fact, that increased bulk would act as a protection against almost all beasts of prey excepting the lion; and against this animal, its tall neck⁠—and the taller the better⁠—would, as <abbr>Mr.</abbr> Chauncey Wright has remarked, serve as a watchtower. It is from this cause, as Sir <abbr class="name">S.</abbr> Baker remarks, that no animal is more difficult to stalk than the giraffe. This animal also uses its long neck as a means of offence or defence, by violently swinging its head armed with stump-like horns. The preservation of each species can rarely be determined by any one advantage, but by the union of all, great and small.</p>
 			<p><abbr>Mr.</abbr> Mivart then asks (and this is his second objection), if natural selection be so potent, and if high browsing be so great an advantage, why has not any other hoofed quadruped acquired a long neck and lofty stature, besides the giraffe, and, in a lesser degree, the camel, guanaco and macrauchenia? Or, again, why has not any member of the group acquired a long proboscis? With respect to South Africa, which was formerly inhabited by numerous herds of the giraffe, the answer is not difficult, and can best be given by an illustration. In every meadow in England, in which trees grow, we see the lower branches trimmed or planed to an exact level by the browsing of the horses or cattle; and what advantage would it be, for instance, to sheep, if kept there, to acquire slightly longer necks? In every district some one kind of animal will almost certainly be able to browse higher than the others; and it is almost equally certain that this one kind alone could have its neck elongated for this purpose, through natural selection and the effects of increased use. In South Africa the competition for browsing on the higher branches of the acacias and other trees must be between giraffe and giraffe, and not with the other ungulate animals.</p>
 			<p>Why, in other quarters of the world, various animals belonging to this same order have not acquired either an elongated neck or a proboscis, cannot be distinctly answered; but it is as unreasonable to expect a distinct answer to such a question as why some event in the history of mankind did not occur in one country while it did in another. We are ignorant with respect to the conditions which determine the numbers and range of each species, and we cannot even conjecture what changes of structure would be favourable to its increase in some new country. We can, however, see in a general manner that various causes might have interfered with the development of a long neck or proboscis. To reach the foliage at a considerable height (without climbing, for which hoofed animals are singularly ill-constructed) implies greatly increased bulk of body; and we know that some areas support singularly few large quadrupeds, for instance South America, though it is so luxuriant, while South Africa abounds with them to an unparalleled degree. Why this should be so we do not know; nor why the later tertiary periods should have been much more favourable for their existence than the present time. Whatever the causes may have been, we can see that certain districts and times would have been much more favourable than others for the development of so large a quadruped as the giraffe.</p>
 			<p>In order that an animal should acquire some structure specially and largely developed, it is almost indispensable that several other parts should be modified and coadapted. Although every part of the body varies slightly, it does not follow that the necessary parts should always vary in the right direction and to the right degree. With the different species of our domesticated animals we know that the parts vary in a different manner and degree, and that some species are much more variable than others. Even if the fitting variations did arise, it does not follow that natural selection would be able to act on them and produce a structure which apparently would be beneficial to the species. For instance, if the number of individuals existing in a country is determined chiefly through destruction by beasts of prey⁠—by external or internal parasites, <abbr>etc.</abbr>⁠—as seems often to be the case, then natural selection will be able to do little, or will be greatly retarded, in modifying any particular structure for obtaining food. Lastly, natural selection is a slow process, and the same favourable conditions must long endure in order that any marked effect should thus be produced. Except by assigning such general and vague reasons, we cannot explain why, in many quarters of the world, hoofed quadrupeds have not acquired much elongated necks or other means for browsing on the higher branches of trees.</p>
-			<p>Objections of the same nature as the foregoing have been advanced by many writers. In each case various causes, besides the general ones just indicated, have probably interfered with the acquisition through natural selection of structures, which it is thought would be beneficial to certain species. One writer asks, why has not the ostrich acquired the power of flight? But a moment’s reflection will show what an enormous supply of food would be necessary to give to this bird of the desert force to move its huge body through the air. Oceanic islands are inhabited by bats and seals, but by no terrestrial mammals; yet as some of these bats are peculiar species, they must have long inhabited their present homes. Therefore Sir C. Lyell asks, and assigns certain reasons in answer, why have not seals and bats given birth on such islands to forms fitted to live on the land? But seals would necessarily be first converted into terrestrial carnivorous animals of considerable size, and bats into terrestrial insectivorous animals; for the former there would be no prey; for the bats ground-insects would serve as food, but these would already be largely preyed on by the reptiles or birds, which first colonise and abound on most oceanic islands. Gradations of structure, with each stage beneficial to a changing species, will be favoured only under certain peculiar conditions. A strictly terrestrial animal, by occasionally hunting for food in shallow water, then in streams or lakes, might at last be converted into an animal so thoroughly aquatic as to brave the open ocean. But seals would not find on oceanic islands the conditions favourable to their gradual reconversion into a terrestrial form. Bats, as formerly shown, probably acquired their wings by at first gliding through the air from tree to tree, like the so-called flying squirrels, for the sake of escaping from their enemies, or for avoiding falls; but when the power of true flight had once been acquired, it would never be reconverted back, at least for the above purposes, into the less efficient power of gliding through the air. Bats, might, indeed, like many birds, have had their wings greatly reduced in size, or completely lost, through disuse; but in this case it would be necessary that they should first have acquired the power of running quickly on the ground, by the aid of their hind legs alone, so as to compete with birds or other ground animals; and for such a change a bat seems singularly ill-fitted. These conjectural remarks have been made merely to show that a transition of structure, with each step beneficial, is a highly complex affair; and that there is nothing strange in a transition not having occurred in any particular case.</p>
+			<p>Objections of the same nature as the foregoing have been advanced by many writers. In each case various causes, besides the general ones just indicated, have probably interfered with the acquisition through natural selection of structures, which it is thought would be beneficial to certain species. One writer asks, why has not the ostrich acquired the power of flight? But a moment’s reflection will show what an enormous supply of food would be necessary to give to this bird of the desert force to move its huge body through the air. Oceanic islands are inhabited by bats and seals, but by no terrestrial mammals; yet as some of these bats are peculiar species, they must have long inhabited their present homes. Therefore Sir <abbr class="name">C.</abbr> Lyell asks, and assigns certain reasons in answer, why have not seals and bats given birth on such islands to forms fitted to live on the land? But seals would necessarily be first converted into terrestrial carnivorous animals of considerable size, and bats into terrestrial insectivorous animals; for the former there would be no prey; for the bats ground-insects would serve as food, but these would already be largely preyed on by the reptiles or birds, which first colonise and abound on most oceanic islands. Gradations of structure, with each stage beneficial to a changing species, will be favoured only under certain peculiar conditions. A strictly terrestrial animal, by occasionally hunting for food in shallow water, then in streams or lakes, might at last be converted into an animal so thoroughly aquatic as to brave the open ocean. But seals would not find on oceanic islands the conditions favourable to their gradual reconversion into a terrestrial form. Bats, as formerly shown, probably acquired their wings by at first gliding through the air from tree to tree, like the so-called flying squirrels, for the sake of escaping from their enemies, or for avoiding falls; but when the power of true flight had once been acquired, it would never be reconverted back, at least for the above purposes, into the less efficient power of gliding through the air. Bats, might, indeed, like many birds, have had their wings greatly reduced in size, or completely lost, through disuse; but in this case it would be necessary that they should first have acquired the power of running quickly on the ground, by the aid of their hind legs alone, so as to compete with birds or other ground animals; and for such a change a bat seems singularly ill-fitted. These conjectural remarks have been made merely to show that a transition of structure, with each step beneficial, is a highly complex affair; and that there is nothing strange in a transition not having occurred in any particular case.</p>
 			<p>Lastly, more than one writer has asked why have some animals had their mental powers more highly developed than others, as such development would be advantageous to all? Why have not apes acquired the intellectual powers of man? Various causes could be assigned; but as they are conjectural, and their relative probability cannot be weighed, it would be useless to give them. A definite answer to the latter question ought not to be expected, seeing that no one can solve the simpler problem, why, of two races of savages, one has risen higher in the scale of civilisation than the other; and this apparently implies increased brain power.</p>
 			<p>We will return to <abbr>Mr.</abbr> Mivart’s other objections. Insects often resemble for the sake of protection various objects, such as green or decayed leaves, dead twigs, bits of lichen, flowers, spines, excrement of birds, and living insects; but to this latter point I shall hereafter recur. The resemblance is often wonderfully close, and is not confined to colour, but extends to form, and even to the manner in which the insects hold themselves. The caterpillars which project motionless like dead twigs from the bushes on which they feed, offer an excellent instance of a resemblance of this kind. The cases of the imitation of such objects as the excrement of birds, are rare and exceptional. On this head, <abbr>Mr.</abbr> Mivart remarks, “As, according to <abbr>Mr.</abbr> Darwin’s theory, there is a constant tendency to indefinite variation, and as the minute incipient variations will be in <em>all directions</em>, they must tend to neutralize each other, and at first to form such unstable modifications that it is difficult, if not impossible, to see how such indefinite oscillations of infinitesimal beginnings can ever build up a sufficiently appreciable resemblance to a leaf, bamboo, or other object, for natural selection to seize upon and perpetuate.”</p>
 			<p>But in all the foregoing cases the insects in their original state no doubt presented some rude and accidental resemblance to an object commonly found in the stations frequented by them. Nor is this at all improbable, considering the almost infinite number of surrounding objects and the diversity in form and colour of the hosts of insects which exist. As some rude resemblance is necessary for the first start, we can understand how it is that the larger and higher animals do not (with the exception, as far as I know, of one fish) resemble for the sake of protection special objects, but only the surface which commonly surrounds them, and this chiefly in colour. Assuming that an insect originally happened to resemble in some degree a dead twig or a decayed leaf, and that it varied slightly in many ways, then all the variations which rendered the insect at all more like any such object, and thus favoured its escape, would be preserved, while other variations would be neglected and ultimately lost; or, if they rendered the insect at all less like the imitated object, they would be eliminated. There would indeed be force in <abbr>Mr.</abbr> Mivart’s objection, if we were to attempt to account for the above resemblances, independently of natural selection, through mere fluctuating variability; but as the case stands there is none.</p>
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 			<p>The mammary glands are common to the whole class of mammals, and are indispensable for their existence; they must, therefore, have been developed at an extremely remote period, and we can know nothing positively about their manner of development. <abbr>Mr.</abbr> Mivart asks: “Is it conceivable that the young of any animal was ever saved from destruction by accidentally sucking a drop of scarcely nutritious fluid from an accidentally hypertrophied cutaneous gland of its mother? And even if one was so, what chance was there of the perpetuation of such a variation?” But the case is not here put fairly. It is admitted by most evolutionists that mammals are descended from a marsupial form; and if so, the mammary glands will have been at first developed within the marsupial sack. In the case of the fish (Hippocampus) the eggs are hatched, and the young are reared for a time, within a sack of this nature; and an American naturalist, <abbr>Mr.</abbr> Lockwood, believes from what he has seen of the development of the young, that they are nourished by a secretion from the cutaneous glands of the sack. Now, with the early progenitors of mammals, almost before they deserved to be thus designated, is it not at least possible that the young might have been similarly nourished? And in this case, the individuals which secreted a fluid, in some degree or manner the most nutritious, so as to partake of the nature of milk, would in the long run have reared a larger number of well-nourished offspring, than would the individuals which secreted a poorer fluid; and thus the cutaneous glands, which are the homologues of the mammary glands, would have been improved or rendered more effective. It accords with the widely extended principle of specialisation, that the glands over a certain space of the sack should have become more highly developed than the remainder; and they would then have formed a breast, but at first without a nipple, as we see in the Ornithorhyncus, at the base of the mammalian series. Through what agency the glands over a certain space became more highly specialised than the others, I will not pretend to decide, whether in part through compensation of growth, the effects of use, or of natural selection.</p>
 			<p>The development of the mammary glands would have been of no service, and could not have been affected through natural selection, unless the young at the same time were able to partake of the secretion. There is no greater difficulty in understanding how young mammals have instinctively learned to suck the breast, than in understanding how unhatched chickens have learned to break the eggshell by tapping against it with their specially adapted beaks; or how a few hours after leaving the shell they have learned to pick up grains of food. In such cases the most probable solution seems to be, that the habit was at first acquired by practice at a more advanced age, and afterwards transmitted to the offspring at an earlier age. But the young kangaroo is said not to suck, only to cling to the nipple of its mother, who has the power of injecting milk into the mouth of her helpless, half-formed offspring. On this head <abbr>Mr.</abbr> Mivart remarks: “Did no special provision exist, the young one must infallibly be choked by the intrusion of the milk into the windpipe. But there IS a special provision. The larynx is so elongated that it rises up into the posterior end of the nasal passage, and is thus enabled to give free entrance to the air for the lungs, while the milk passes harmlessly on each side of this elongated larynx, and so safely attains the gullet behind it.” <abbr>Mr.</abbr> Mivart then asks how did natural selection remove in the adult kangaroo (and in most other mammals, on the assumption that they are descended from a marsupial form), “this at least perfectly innocent and harmless structure?” It may be suggested in answer that the voice, which is certainly of high importance to many animals, could hardly have been used with full force as long as the larynx entered the nasal passage; and Professor Flower has suggested to me that this structure would have greatly interfered with an animal swallowing solid food.</p>
 			<p>We will now turn for a short space to the lower divisions of the animal kingdom. The Echinodermata (starfishes, sea-urchins, <abbr>etc.</abbr>) are furnished with remarkable organs, called pedicellariae, which consist, when well developed, of a tridactyle forceps⁠—that is, of one formed of three serrated arms, neatly fitting together and placed on the summit of a flexible stem, moved by muscles. These forceps can seize firmly hold of any object; and Alexander Agassiz has seen an Echinus or sea-urchin rapidly passing particles of excrement from forceps to forceps down certain lines of its body, in order that its shell should not be fouled. But there is no doubt that besides removing dirt of all kinds, they subserve other functions; and one of these apparently is defence.</p>
-			<p>With respect to these organs, <abbr>Mr.</abbr> Mivart, as on so many previous occasions, asks: “What would be the utility of the <em>first rudimentary beginnings</em> of such structures, and how could such insipient buddings have ever preserved the life of a single Echinus?” He adds, “not even the <em>sudden</em> development of the snapping action would have been beneficial without the freely movable stalk, nor could the latter have been efficient without the snapping jaws, yet no minute, nearly indefinite variations could simultaneously evolve these complex coordinations of structure; to deny this seems to do no less than to affirm a startling paradox.” Paradoxical as this may appear to <abbr>Mr.</abbr> Mivart, tridactyle forcepses, immovably fixed at the base, but capable of a snapping action, certainly exist on some starfishes; and this is intelligible if they serve, at least in part, as a means of defence. <abbr>Mr.</abbr> Agassiz, to whose great kindness I am indebted for much information on the subject, informs me that there are other starfishes, in which one of the three arms of the forceps is reduced to a support for the other two; and again, other genera in which the third arm is completely lost. In Echinoneus, the shell is described by M. Perrier as bearing two kinds of pedicellariae, one resembling those of Echinus, and the other those of Spatangus; and such cases are always interesting as affording the means of apparently sudden transitions, through the abortion of one of the two states of an organ.</p>
+			<p>With respect to these organs, <abbr>Mr.</abbr> Mivart, as on so many previous occasions, asks: “What would be the utility of the <em>first rudimentary beginnings</em> of such structures, and how could such insipient buddings have ever preserved the life of a single Echinus?” He adds, “not even the <em>sudden</em> development of the snapping action would have been beneficial without the freely movable stalk, nor could the latter have been efficient without the snapping jaws, yet no minute, nearly indefinite variations could simultaneously evolve these complex coordinations of structure; to deny this seems to do no less than to affirm a startling paradox.” Paradoxical as this may appear to <abbr>Mr.</abbr> Mivart, tridactyle forcepses, immovably fixed at the base, but capable of a snapping action, certainly exist on some starfishes; and this is intelligible if they serve, at least in part, as a means of defence. <abbr>Mr.</abbr> Agassiz, to whose great kindness I am indebted for much information on the subject, informs me that there are other starfishes, in which one of the three arms of the forceps is reduced to a support for the other two; and again, other genera in which the third arm is completely lost. In Echinoneus, the shell is described by <abbr class="name">M.</abbr> Perrier as bearing two kinds of pedicellariae, one resembling those of Echinus, and the other those of Spatangus; and such cases are always interesting as affording the means of apparently sudden transitions, through the abortion of one of the two states of an organ.</p>
 			<p>With respect to the steps by which these curious organs have been evolved, <abbr>Mr.</abbr> Agassiz infers from his own researches and those of <abbr>Mr.</abbr> Muller, that both in starfishes and sea-urchins the pedicellariae must undoubtedly be looked at as modified spines. This may be inferred from their manner of development in the individual, as well as from a long and perfect series of gradations in different species and genera, from simple granules to ordinary spines, to perfect tridactyle pedicellariae. The gradation extends even to the manner in which ordinary spines and the pedicellariae, with their supporting calcareous rods, are articulated to the shell. In certain genera of starfishes, “the very combinations needed to show that the pedicellariae are only modified branching spines” may be found. Thus we have fixed spines, with three equidistant, serrated, movable branches, articulated to near their bases; and higher up, on the same spine, three other movable branches. Now when the latter arise from the summit of a spine they form, in fact, a rude tridactyle pedicellariae, and such may be seen on the same spine together with the three lower branches. In this case the identity in nature between the arms of the pedicellariae and the movable branches of a spine, is unmistakable. It is generally admitted that the ordinary spines serve as a protection; and if so, there can be no reason to doubt that those furnished with serrated and movable branches likewise serve for the same purpose; and they would thus serve still more effectively as soon as by meeting together they acted as a prehensile or snapping apparatus. Thus every gradation, from an ordinary fixed spine to a fixed pedicellariae, would be of service.</p>
 			<p>In certain genera of starfishes these organs, instead of being fixed or borne on an immovable support, are placed on the summit of a flexible and muscular, though short, stem; and in this case they probably subserve some additional function besides defence. In the sea-urchins the steps can be followed by which a fixed spine becomes articulated to the shell, and is thus rendered movable. I wish I had space here to give a fuller abstract of <abbr>Mr.</abbr> Agassiz’s interesting observations on the development of the pedicellariae. All possible gradations, as he adds, may likewise be found between the pedicellariae of the starfishes and the hooks of the Ophiurians, another group of the Echinodermata; and again between the pedicellariae of sea-urchins and the anchors of the Holothuriae, also belonging to the same great class.</p>
 			<p>Certain compound animals, or zoophytes, as they have been termed, namely the Polyzoa, are provided with curious organs called avicularia. These differ much in structure in the different species. In their most perfect condition they curiously resemble the head and beak of a vulture in miniature, seated on a neck and capable of movement, as is likewise the lower jaw or mandible. In one species observed by me, all the avicularia on the same branch often moved simultaneously backwards and forwards, with the lower jaw widely open, through an angle of about 90 degrees, in the course of five seconds; and their movement caused the whole polyzoary to tremble. When the jaws are touched with a needle they seize it so firmly that the branch can thus be shaken.</p>

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 			<p>Many instincts are so wonderful that their development will probably appear to the reader a difficulty sufficient to overthrow my whole theory. I may here premise, that I have nothing to do with the origin of the mental powers, any more than I have with that of life itself. We are concerned only with the diversities of instinct and of the other mental faculties in animals of the same class.</p>
 			<p>I will not attempt any definition of instinct. It would be easy to show that several distinct mental actions are commonly embraced by this term; but everyone understands what is meant, when it is said that instinct impels the cuckoo to migrate and to lay her eggs in other birds’ nests. An action, which we ourselves require experience to enable us to perform, when performed by an animal, more especially by a very young one, without experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive. But I could show that none of these characters are universal. A little dose of judgment or reason, as Pierre Huber expresses it, often comes into play, even with animals low in the scale of nature.</p>
-			<p>Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. This comparison gives, I think, an accurate notion of the frame of mind under which an instinctive action is performed, but not necessarily of its origin. How unconsciously many habitual actions are performed, indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason. Habits easily become associated with other habits, with certain periods of time and states of the body. When once acquired, they often remain constant throughout life. Several other points of resemblance between instincts and habits could be pointed out. As in repeating a well-known song, so in instincts, one action follows another by a sort of rhythm; if a person be interrupted in a song, or in repeating anything by rote, he is generally forced to go back to recover the habitual train of thought: so P. Huber found it was with a caterpillar, which makes a very complicated hammock; for if he took a caterpillar which had completed its hammock up to, say, the sixth stage of construction, and put it into a hammock completed up only to the third stage, the caterpillar simply re-performed the fourth, fifth, and sixth stages of construction. If, however, a caterpillar were taken out of a hammock made up, for instance, to the third stage, and were put into one finished up to the sixth stage, so that much of its work was already done for it, far from deriving any benefit from this, it was much embarrassed, and, in order to complete its hammock, seemed forced to start from the third stage, where it had left off, and thus tried to complete the already finished work.</p>
+			<p>Frederick Cuvier and several of the older metaphysicians have compared instinct with habit. This comparison gives, I think, an accurate notion of the frame of mind under which an instinctive action is performed, but not necessarily of its origin. How unconsciously many habitual actions are performed, indeed not rarely in direct opposition to our conscious will! yet they may be modified by the will or reason. Habits easily become associated with other habits, with certain periods of time and states of the body. When once acquired, they often remain constant throughout life. Several other points of resemblance between instincts and habits could be pointed out. As in repeating a well-known song, so in instincts, one action follows another by a sort of rhythm; if a person be interrupted in a song, or in repeating anything by rote, he is generally forced to go back to recover the habitual train of thought: so <abbr class="name">P.</abbr> Huber found it was with a caterpillar, which makes a very complicated hammock; for if he took a caterpillar which had completed its hammock up to, say, the sixth stage of construction, and put it into a hammock completed up only to the third stage, the caterpillar simply re-performed the fourth, fifth, and sixth stages of construction. If, however, a caterpillar were taken out of a hammock made up, for instance, to the third stage, and were put into one finished up to the sixth stage, so that much of its work was already done for it, far from deriving any benefit from this, it was much embarrassed, and, in order to complete its hammock, seemed forced to start from the third stage, where it had left off, and thus tried to complete the already finished work.</p>
 			<p>If we suppose any habitual action to become inherited⁠—and it can be shown that this does sometimes happen⁠—then the resemblance between what originally was a habit and an instinct becomes so close as not to be distinguished. If Mozart, instead of playing the pianoforte at three years old with wonderfully little practice, had played a tune with no practice at all, be might truly be said to have done so instinctively. But it would be a serious error to suppose that the greater number of instincts have been acquired by habit in one generation, and then transmitted by inheritance to succeeding generations. It can be clearly shown that the most wonderful instincts with which we are acquainted, namely, those of the hive-bee and of many ants, could not possibly have been acquired by habit.</p>
 			<p>It will be universally admitted that instincts are as important as corporeal structures for the welfare of each species, under its present conditions of life. Under changed conditions of life, it is at least possible that slight modifications of instinct might be profitable to a species; and if it can be shown that instincts do vary ever so little, then I can see no difficulty in natural selection preserving and continually accumulating variations of instinct to any extent that was profitable. It is thus, as I believe, that all the most complex and wonderful instincts have originated. As modifications of corporeal structure arise from, and are increased by, use or habit, and are diminished or lost by disuse, so I do not doubt it has been with instincts. But I believe that the effects of habit are in many cases of subordinate importance to the effects of the natural selection of what may be called spontaneous variations of instincts;⁠—that is of variations produced by the same unknown causes which produce slight deviations of bodily structure.</p>
 			<p>No complex instinct can possibly be produced through natural selection, except by the slow and gradual accumulation of numerous, slight, yet profitable, variations. Hence, as in the case of corporeal structures, we ought to find in nature, not the actual transitional gradations by which each complex instinct has been acquired⁠—for these could be found only in the lineal ancestors of each species⁠—but we ought to find in the collateral lines of descent some evidence of such gradations; or we ought at least to be able to show that gradations of some kind are possible; and this we certainly can do. I have been surprised to find, making allowance for the instincts of animals having been but little observed, except in Europe and North America, and for no instinct being known among extinct species, how very generally gradations, leading to the most complex instincts, can be discovered. Changes of instinct may sometimes be facilitated by the same species having different instincts at different periods of life, or at different seasons of the year, or when placed under different circumstances, <abbr>etc.</abbr>; in which case either the one or the other instinct might be preserved by natural selection. And such instances of diversity of instinct in the same species can be shown to occur in nature.</p>
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 				<p>It has been objected that I have not noticed other related instincts and adaptations of structure in the cuckoo, which are spoken of as necessarily coordinated. But in all cases, speculation on an instinct known to us only in a single species, is useless, for we have hitherto had no facts to guide us. Until recently the instincts of the European and of the nonparasitic American cuckoo alone were known; now, owing to <abbr>Mr.</abbr> Ramsay’s observations, we have learned something about three Australian species, which lay their eggs in other birds’ nests. The chief points to be referred to are three: first, that the common cuckoo, with rare exceptions, lays only one egg in a nest, so that the large and voracious young bird receives ample food. Secondly, that the eggs are remarkably small, not exceeding those of the skylark⁠—a bird about one-fourth as large as the cuckoo. That the small size of the egg is a real case of adaptation we may infer from the fact of the mon-parasitic American cuckoo laying full-sized eggs. Thirdly, that the young cuckoo, soon after birth, has the instinct, the strength and a properly shaped back for ejecting its foster-brothers, which then perish from cold and hunger. This has been boldly called a beneficent arrangement, in order that the young cuckoo may get sufficient food, and that its foster-brothers may perish before they had acquired much feeling!</p>
 				<p>Turning now to the Australian species: though these birds generally lay only one egg in a nest, it is not rare to find two and even three eggs in the same nest. In the bronze cuckoo the eggs vary greatly in size, from eight to ten lines in length. Now, if it had been of an advantage to this species to have laid eggs even smaller than those now laid, so as to have deceived certain foster-parents, or, as is more probable, to have been hatched within a shorter period (for it is asserted that there is a relation between the size of eggs and the period of their incubation), then there is no difficulty in believing that a race or species might have been formed which would have laid smaller and smaller eggs; for these would have been more safely hatched and reared. <abbr>Mr.</abbr> Ramsay remarks that two of the Australian cuckoos, when they lay their eggs in an open nest, manifest a decided preference for nests containing eggs similar in colour to their own. The European species apparently manifests some tendency towards a similar instinct, but not rarely departs from it, as is shown by her laying her dull and pale-coloured eggs in the nest of the hedge-warbler with bright greenish-blue eggs. Had our cuckoo invariably displayed the above instinct, it would assuredly have been added to those which it is assumed must all have been acquired together. The eggs of the Australian bronze cuckoo vary, according to <abbr>Mr.</abbr> Ramsay, to an extraordinary degree in colour; so that in this respect, as well as in size, natural selection might have secured and fixed any advantageous variation.</p>
 				<p>In the case of the European cuckoo, the offspring of the foster-parents are commonly ejected from the nest within three days after the cuckoo is hatched; and as the latter at this age is in a most helpless condition, <abbr>Mr.</abbr> Gould was formerly inclined to believe that the act of ejection was performed by the foster-parents themselves. But he has now received a trustworthy account of a young cuckoo which was actually seen, while still blind and not able even to hold up its own head, in the act of ejecting its foster-brothers. One of these was replaced in the nest by the observer, and was again thrown out. With respect to the means by which this strange and odious instinct was acquired, if it were of great importance for the young cuckoo, as is probably the case, to receive as much food as possible soon after birth, I can see no special difficulty in its having gradually acquired, during successive generations, the blind desire, the strength, and structure necessary for the work of ejection; for those cuckoos which had such habits and structure best developed would be the most securely reared. The first step towards the acquisition of the proper instinct might have been mere unintentional restlessness on the part of the young bird, when somewhat advanced in age and strength; the habit having been afterwards improved, and transmitted to an earlier age. I can see no more difficulty in this than in the unhatched young of other birds acquiring the instinct to break through their own shells; or than in young snakes acquiring in their upper jaws, as Owen has remarked, a transitory sharp tooth for cutting through the tough eggshell. For if each part is liable to individual variations at all ages, and the variations tend to be inherited at a corresponding or earlier age⁠—propositions which cannot be disputed⁠—then the instincts and structure of the young could be slowly modified as surely as those of the adult; and both cases must stand or fall together with the whole theory of natural selection.</p>
-				<p>Some species of Molothrus, a widely distinct genus of American birds, allied to our starlings, have parasitic habits like those of the cuckoo; and the species present an interesting gradation in the perfection of their instincts. The sexes of Molothrus badius are stated by an excellent observer, <abbr>Mr.</abbr> Hudson, sometimes to live promiscuously together in flocks, and sometimes to pair. They either build a nest of their own or seize on one belonging to some other bird, occasionally throwing out the nestlings of the stranger. They either lay their eggs in the nest thus appropriated, or oddly enough build one for themselves on the top of it. They usually sit on their own eggs and rear their own young; but <abbr>Mr.</abbr> Hudson says it is probable that they are occasionally parasitic, for he has seen the young of this species following old birds of a distinct kind and clamouring to be fed by them. The parasitic habits of another species of Molothrus, the M. bonariensis, are much more highly developed than those of the last, but are still far from perfect. This bird, as far as it is known, invariably lays its eggs in the nests of strangers; but it is remarkable that several together sometimes commence to build an irregular untidy nest of their own, placed in singular ill-adapted situations, as on the leaves of a large thistle. They never, however, as far as <abbr>Mr.</abbr> Hudson has ascertained, complete a nest for themselves. They often lay so many eggs⁠—from fifteen to twenty⁠—in the same foster-nest, that few or none can possibly be hatched. They have, moreover, the extraordinary habit of pecking holes in the eggs, whether of their own species or of their foster parents, which they find in the appropriated nests. They drop also many eggs on the bare ground, which are thus wasted. A third species, the M. pecoris of North America, has acquired instincts as perfect as those of the cuckoo, for it never lays more than one egg in a foster-nest, so that the young bird is securely reared. <abbr>Mr.</abbr> Hudson is a strong disbeliever in evolution, but he appears to have been so much struck by the imperfect instincts of the Molothrus bonariensis that he quotes my words, and asks, “Must we consider these habits, not as especially endowed or created instincts, but as small consequences of one general law, namely, transition?”</p>
+				<p>Some species of Molothrus, a widely distinct genus of American birds, allied to our starlings, have parasitic habits like those of the cuckoo; and the species present an interesting gradation in the perfection of their instincts. The sexes of Molothrus badius are stated by an excellent observer, <abbr>Mr.</abbr> Hudson, sometimes to live promiscuously together in flocks, and sometimes to pair. They either build a nest of their own or seize on one belonging to some other bird, occasionally throwing out the nestlings of the stranger. They either lay their eggs in the nest thus appropriated, or oddly enough build one for themselves on the top of it. They usually sit on their own eggs and rear their own young; but <abbr>Mr.</abbr> Hudson says it is probable that they are occasionally parasitic, for he has seen the young of this species following old birds of a distinct kind and clamouring to be fed by them. The parasitic habits of another species of Molothrus, the <i epub:type="z3998:taxonomy">M. bonariensis</i>, are much more highly developed than those of the last, but are still far from perfect. This bird, as far as it is known, invariably lays its eggs in the nests of strangers; but it is remarkable that several together sometimes commence to build an irregular untidy nest of their own, placed in singular ill-adapted situations, as on the leaves of a large thistle. They never, however, as far as <abbr>Mr.</abbr> Hudson has ascertained, complete a nest for themselves. They often lay so many eggs⁠—from fifteen to twenty⁠—in the same foster-nest, that few or none can possibly be hatched. They have, moreover, the extraordinary habit of pecking holes in the eggs, whether of their own species or of their foster parents, which they find in the appropriated nests. They drop also many eggs on the bare ground, which are thus wasted. A third species, the <i epub:type="z3998:taxonomy">M. pecoris</i> of North America, has acquired instincts as perfect as those of the cuckoo, for it never lays more than one egg in a foster-nest, so that the young bird is securely reared. <abbr>Mr.</abbr> Hudson is a strong disbeliever in evolution, but he appears to have been so much struck by the imperfect instincts of the Molothrus bonariensis that he quotes my words, and asks, “Must we consider these habits, not as especially endowed or created instincts, but as small consequences of one general law, namely, transition?”</p>
 				<p>Various birds, as has already been remarked, occasionally lay their eggs in the nests of other birds. This habit is not very uncommon with the Gallinaceae, and throws some light on the singular instinct of the ostrich. In this family several hen birds unite and lay first a few eggs in one nest and then in another; and these are hatched by the males. This instinct may probably be accounted for by the fact of the hens laying a large number of eggs, but, as with the cuckoo, at intervals of two or three days. The instinct, however, of the American ostrich, as in the case of the Molothrus bonariensis, has not as yet been perfected; for a surprising number of eggs lie strewed over the plains, so that in one day’s hunting I picked up no less than twenty lost and wasted eggs.</p>
-				<p>Many bees are parasitic, and regularly lay their eggs in the nests of other kinds of bees. This case is more remarkable than that of the cuckoo; for these bees have not only had their instincts but their structure modified in accordance with their parasitic habits; for they do not possess the pollen-collecting apparatus which would have been indispensable if they had stored up food for their own young. Some species of Sphegidae (wasp-like insects) are likewise parasitic; and M. Fabre has lately shown good reason for believing that, although the Tachytes nigra generally makes its own burrow and stores it with paralysed prey for its own larvae, yet that, when this insect finds a burrow already made and stored by another sphex, it takes advantage of the prize, and becomes for the occasion parasitic. In this case, as with that of the Molothrus or cuckoo, I can see no difficulty in natural selection making an occasional habit permanent, if of advantage to the species, and if the insect whose nest and stored food are feloniously appropriated, be not thus exterminated.</p>
+				<p>Many bees are parasitic, and regularly lay their eggs in the nests of other kinds of bees. This case is more remarkable than that of the cuckoo; for these bees have not only had their instincts but their structure modified in accordance with their parasitic habits; for they do not possess the pollen-collecting apparatus which would have been indispensable if they had stored up food for their own young. Some species of Sphegidae (wasp-like insects) are likewise parasitic; and <abbr class="name">M.</abbr> Fabre has lately shown good reason for believing that, although the Tachytes nigra generally makes its own burrow and stores it with paralysed prey for its own larvae, yet that, when this insect finds a burrow already made and stored by another sphex, it takes advantage of the prize, and becomes for the occasion parasitic. In this case, as with that of the Molothrus or cuckoo, I can see no difficulty in natural selection making an occasional habit permanent, if of advantage to the species, and if the insect whose nest and stored food are feloniously appropriated, be not thus exterminated.</p>
 			</section>
 			<section id="chapter-8-4" epub:type="z3998:subchapter">
 				<h3 epub:type="title">Slave-Making Instinct</h3>
-				<p>This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work of any kind, and the workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own larvae. When the old nest is found inconvenient, and they have to migrate, it is the slaves which determine the migration, and actually carry their masters in their jaws. So utterly helpless are the masters, that when Huber shut up thirty of them without a slave, but with plenty of the food which they like best, and with their larvae and pupae to stimulate them to work, they did nothing; they could not even feed themselves, and many perished of hunger. Huber then introduced a single slave (F. fusca), and she instantly set to work, fed and saved the survivors; made some cells and tended the larvae, and put all to rights. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant, it would have been hopeless to speculate how so wonderful an instinct could have been perfected.</p>
-				<p>Another species, Formica sanguinea, was likewise first discovered by P. Huber to be a slave-making ant. This species is found in the southern parts of England, and its habits have been attended to by <abbr>Mr.</abbr> <abbr class="name">F.</abbr> Smith, of the British Museum, to whom I am much indebted for information on this and other subjects. Although fully trusting to the statements of Huber and <abbr>Mr.</abbr> Smith, I tried to approach the subject in a sceptical frame of mind, as anyone may well be excused for doubting the existence of so extraordinary an instinct as that of making slaves. Hence, I will give the observations which I made in some little detail. I opened fourteen nests of F. sanguinea, and found a few slaves in all. Males and fertile females of the slave-species (F. fusca) are found only in their own proper communities, and have never been observed in the nests of F. sanguinea. The slaves are black and not above half the size of their red masters, so that the contrast in their appearance is great. When the nest is slightly disturbed, the slaves occasionally come out, and like their masters are much agitated and defend the nest: when the nest is much disturbed, and the larvae and pupae are exposed, the slaves work energetically together with their masters in carrying them away to a place of safety. Hence, it is clear that the slaves feel quite at home. During the months of June and July, on three successive years, I watched for many hours several nests in Surrey and Sussex, and never saw a slave either leave or enter a nest. As, during these months, the slaves are very few in number, I thought that they might behave differently when more numerous; but <abbr>Mr.</abbr> Smith informs me that he has watched the nests at various hours during May, June and August, both in Surrey and Hampshire, and has never seen the slaves, though present in large numbers in August, either leave or enter the nest. Hence, he considers them as strictly household slaves. The masters, on the other hand, may be constantly seen bringing in materials for the nest, and food of all kinds. During the year 1860, however, in the month of July, I came across a community with an unusually large stock of slaves, and I observed a few slaves mingled with their masters leaving the nest, and marching along the same road to a tall Scotch-fir tree, twenty-five yards distant, which they ascended together, probably in search of aphides or cocci. According to Huber, who had ample opportunities for observation, the slaves in Switzerland habitually work with their masters in making the nest, and they alone open and close the doors in the morning and evening; and, as Huber expressly states, their principal office is to search for aphides. This difference in the usual habits of the masters and slaves in the two countries, probably depends merely on the slaves being captured in greater numbers in Switzerland than in England.</p>
-				<p>One day I fortunately witnessed a migration of F. sanguinea from one nest to another, and it was a most interesting spectacle to behold the masters carefully carrying their slaves in their jaws instead of being carried by them, as in the case of F. rufescens. Another day my attention was struck by about a score of the slave-makers haunting the same spot, and evidently not in search of food; they approached and were vigorously repulsed by an independent community of the slave species (F. fusca); sometimes as many as three of these ants clinging to the legs of the slave-making F. sanguinea. The latter ruthlessly killed their small opponents and carried their dead bodies as food to their nest, twenty-nine yards distant; but they were prevented from getting any pupae to rear as slaves. I then dug up a small parcel of the pupae of F. fusca from another nest, and put them down on a bare spot near the place of combat; they were eagerly seized and carried off by the tyrants, who perhaps fancied that, after all, they had been victorious in their late combat.</p>
-				<p>At the same time I laid on the same place a small parcel of the pupae of another species, F. flava, with a few of these little yellow ants still clinging to the fragments of their nest. This species is sometimes, though rarely, made into slaves, as has been described by <abbr>Mr.</abbr> Smith. Although so small a species, it is very courageous, and I have seen it ferociously attack other ants. In one instance I found to my surprise an independent community of F. flava under a stone beneath a nest of the slave-making F. sanguinea; and when I had accidentally disturbed both nests, the little ants attacked their big neighbours with surprising courage. Now I was curious to ascertain whether F. sanguinea could distinguish the pupae of F. fusca, which they habitually make into slaves, from those of the little and furious F. flava, which they rarely capture, and it was evident that they did at once distinguish them; for we have seen that they eagerly and instantly seized the pupae of F. fusca, whereas they were much terrified when they came across the pupae, or even the earth from the nest, of F. flava, and quickly ran away; but in about a quarter of an hour, shortly after all the little yellow ants had crawled away, they took heart and carried off the pupae.</p>
-				<p>One evening I visited another community of F. sanguinea, and found a number of these ants returning home and entering their nests, carrying the dead bodies of F. fusca (showing that it was not a migration) and numerous pupae. I traced a long file of ants burdened with booty, for about forty yards back, to a very thick clump of heath, whence I saw the last individual of F. sanguinea emerge, carrying a pupa; but I was not able to find the desolated nest in the thick heath. The nest, however, must have been close at hand, for two or three individuals of F. fusca were rushing about in the greatest agitation, and one was perched motionless with its own pupa in its mouth on the top of a spray of heath, an image of despair over its ravaged home.</p>
-				<p>Such are the facts, though they did not need confirmation by me, in regard to the wonderful instinct of making slaves. Let it be observed what a contrast the instinctive habits of F. sanguinea present with those of the continental F. rufescens. The latter does not build its own nest, does not determine its own migrations, does not collect food for itself or its young, and cannot even feed itself: it is absolutely dependent on its numerous slaves. Formica sanguinea, on the other hand, possesses much fewer slaves, and in the early part of the summer extremely few. The masters determine when and where a new nest shall be formed, and when they migrate, the masters carry the slaves. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae, and the masters alone go on slave-making expeditions. In Switzerland the slaves and masters work together, making and bringing materials for the nest: both, but chiefly the slaves, tend and milk as it may be called, their aphides; and thus both collect food for the community. In England the masters alone usually leave the nest to collect building materials and food for themselves, their slaves and larvae. So that the masters in this country receive much less service from their slaves than they do in Switzerland.</p>
-				<p>By what steps the instinct of F. sanguinea originated I will not pretend to conjecture. But as ants which are not slave-makers, will, as I have seen, carry off pupae of other species, if scattered near their nests, it is possible that such pupae originally stored as food might become developed; and the foreign ants thus unintentionally reared would then follow their proper instincts, and do what work they could. If their presence proved useful to the species which had seized them⁠—if it were more advantageous to this species, to capture workers than to procreate them⁠—the habit of collecting pupae, originally for food, might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. When the instinct was once acquired, if carried out to a much less extent even than in our British F. sanguinea, which, as we have seen, is less aided by its slaves than the same species in Switzerland, natural selection might increase and modify the instinct⁠—always supposing each modification to be of use to the species⁠—until an ant was formed as abjectly dependent on its slaves as is the Formica rufescens.</p>
+				<p>This remarkable instinct was first discovered in the Formica (Polyerges) rufescens by Pierre Huber, a better observer even than his celebrated father. This ant is absolutely dependent on its slaves; without their aid, the species would certainly become extinct in a single year. The males and fertile females do no work of any kind, and the workers or sterile females, though most energetic and courageous in capturing slaves, do no other work. They are incapable of making their own nests, or of feeding their own larvae. When the old nest is found inconvenient, and they have to migrate, it is the slaves which determine the migration, and actually carry their masters in their jaws. So utterly helpless are the masters, that when Huber shut up thirty of them without a slave, but with plenty of the food which they like best, and with their larvae and pupae to stimulate them to work, they did nothing; they could not even feed themselves, and many perished of hunger. Huber then introduced a single slave (<i epub:type="z3998:taxonomy">F. fusca</i>), and she instantly set to work, fed and saved the survivors; made some cells and tended the larvae, and put all to rights. What can be more extraordinary than these well-ascertained facts? If we had not known of any other slave-making ant, it would have been hopeless to speculate how so wonderful an instinct could have been perfected.</p>
+				<p>Another species, <i epub:type="z3998:taxonomy">Formica sanguinea</i>, was likewise first discovered by <abbr class="name">P.</abbr> Huber to be a slave-making ant. This species is found in the southern parts of England, and its habits have been attended to by <abbr>Mr.</abbr> <abbr class="name">F.</abbr> Smith, of the British Museum, to whom I am much indebted for information on this and other subjects. Although fully trusting to the statements of Huber and <abbr>Mr.</abbr> Smith, I tried to approach the subject in a sceptical frame of mind, as anyone may well be excused for doubting the existence of so extraordinary an instinct as that of making slaves. Hence, I will give the observations which I made in some little detail. I opened fourteen nests of <i epub:type="z3998:taxonomy">F. sanguinea</i>, and found a few slaves in all. Males and fertile females of the slave-species (<i epub:type="z3998:taxonomy">F. fusca</i>) are found only in their own proper communities, and have never been observed in the nests of <i epub:type="z3998:taxonomy">F. sanguinea</i>. The slaves are black and not above half the size of their red masters, so that the contrast in their appearance is great. When the nest is slightly disturbed, the slaves occasionally come out, and like their masters are much agitated and defend the nest: when the nest is much disturbed, and the larvae and pupae are exposed, the slaves work energetically together with their masters in carrying them away to a place of safety. Hence, it is clear that the slaves feel quite at home. During the months of June and July, on three successive years, I watched for many hours several nests in Surrey and Sussex, and never saw a slave either leave or enter a nest. As, during these months, the slaves are very few in number, I thought that they might behave differently when more numerous; but <abbr>Mr.</abbr> Smith informs me that he has watched the nests at various hours during May, June and August, both in Surrey and Hampshire, and has never seen the slaves, though present in large numbers in August, either leave or enter the nest. Hence, he considers them as strictly household slaves. The masters, on the other hand, may be constantly seen bringing in materials for the nest, and food of all kinds. During the year 1860, however, in the month of July, I came across a community with an unusually large stock of slaves, and I observed a few slaves mingled with their masters leaving the nest, and marching along the same road to a tall Scotch-fir tree, twenty-five yards distant, which they ascended together, probably in search of aphides or cocci. According to Huber, who had ample opportunities for observation, the slaves in Switzerland habitually work with their masters in making the nest, and they alone open and close the doors in the morning and evening; and, as Huber expressly states, their principal office is to search for aphides. This difference in the usual habits of the masters and slaves in the two countries, probably depends merely on the slaves being captured in greater numbers in Switzerland than in England.</p>
+				<p>One day I fortunately witnessed a migration of <i epub:type="z3998:taxonomy">F. sanguinea</i> from one nest to another, and it was a most interesting spectacle to behold the masters carefully carrying their slaves in their jaws instead of being carried by them, as in the case of <i epub:type="z3998:taxonomy">F. rufescens</i>. Another day my attention was struck by about a score of the slave-makers haunting the same spot, and evidently not in search of food; they approached and were vigorously repulsed by an independent community of the slave species (<i epub:type="z3998:taxonomy">F. fusca</i>); sometimes as many as three of these ants clinging to the legs of the slave-making <i epub:type="z3998:taxonomy">F. sanguinea</i>. The latter ruthlessly killed their small opponents and carried their dead bodies as food to their nest, twenty-nine yards distant; but they were prevented from getting any pupae to rear as slaves. I then dug up a small parcel of the pupae of <i epub:type="z3998:taxonomy">F. fusca</i> from another nest, and put them down on a bare spot near the place of combat; they were eagerly seized and carried off by the tyrants, who perhaps fancied that, after all, they had been victorious in their late combat.</p>
+				<p>At the same time I laid on the same place a small parcel of the pupae of another species, <i epub:type="z3998:taxonomy">F. flava</i>, with a few of these little yellow ants still clinging to the fragments of their nest. This species is sometimes, though rarely, made into slaves, as has been described by <abbr>Mr.</abbr> Smith. Although so small a species, it is very courageous, and I have seen it ferociously attack other ants. In one instance I found to my surprise an independent community of <i epub:type="z3998:taxonomy">F. flava</i> under a stone beneath a nest of the slave-making <i epub:type="z3998:taxonomy">F. sanguinea</i>; and when I had accidentally disturbed both nests, the little ants attacked their big neighbours with surprising courage. Now I was curious to ascertain whether <i epub:type="z3998:taxonomy">F. sanguinea</i> could distinguish the pupae of <i epub:type="z3998:taxonomy">F. fusca</i>, which they habitually make into slaves, from those of the little and furious <i epub:type="z3998:taxonomy">F. flava</i>, which they rarely capture, and it was evident that they did at once distinguish them; for we have seen that they eagerly and instantly seized the pupae of <i epub:type="z3998:taxonomy">F. fusca</i>, whereas they were much terrified when they came across the pupae, or even the earth from the nest, of <i epub:type="z3998:taxonomy">F. flava</i>, and quickly ran away; but in about a quarter of an hour, shortly after all the little yellow ants had crawled away, they took heart and carried off the pupae.</p>
+				<p>One evening I visited another community of <i epub:type="z3998:taxonomy">F. sanguinea</i>, and found a number of these ants returning home and entering their nests, carrying the dead bodies of <i epub:type="z3998:taxonomy">F. fusca</i> (showing that it was not a migration) and numerous pupae. I traced a long file of ants burdened with booty, for about forty yards back, to a very thick clump of heath, whence I saw the last individual of <i epub:type="z3998:taxonomy">F. sanguinea</i> emerge, carrying a pupa; but I was not able to find the desolated nest in the thick heath. The nest, however, must have been close at hand, for two or three individuals of <i epub:type="z3998:taxonomy">F. fusca</i> were rushing about in the greatest agitation, and one was perched motionless with its own pupa in its mouth on the top of a spray of heath, an image of despair over its ravaged home.</p>
+				<p>Such are the facts, though they did not need confirmation by me, in regard to the wonderful instinct of making slaves. Let it be observed what a contrast the instinctive habits of <i epub:type="z3998:taxonomy">F. sanguinea</i> present with those of the continental <i epub:type="z3998:taxonomy">F. rufescens</i>. The latter does not build its own nest, does not determine its own migrations, does not collect food for itself or its young, and cannot even feed itself: it is absolutely dependent on its numerous slaves. Formica sanguinea, on the other hand, possesses much fewer slaves, and in the early part of the summer extremely few. The masters determine when and where a new nest shall be formed, and when they migrate, the masters carry the slaves. Both in Switzerland and England the slaves seem to have the exclusive care of the larvae, and the masters alone go on slave-making expeditions. In Switzerland the slaves and masters work together, making and bringing materials for the nest: both, but chiefly the slaves, tend and milk as it may be called, their aphides; and thus both collect food for the community. In England the masters alone usually leave the nest to collect building materials and food for themselves, their slaves and larvae. So that the masters in this country receive much less service from their slaves than they do in Switzerland.</p>
+				<p>By what steps the instinct of <i epub:type="z3998:taxonomy">F. sanguinea</i> originated I will not pretend to conjecture. But as ants which are not slave-makers, will, as I have seen, carry off pupae of other species, if scattered near their nests, it is possible that such pupae originally stored as food might become developed; and the foreign ants thus unintentionally reared would then follow their proper instincts, and do what work they could. If their presence proved useful to the species which had seized them⁠—if it were more advantageous to this species, to capture workers than to procreate them⁠—the habit of collecting pupae, originally for food, might by natural selection be strengthened and rendered permanent for the very different purpose of raising slaves. When the instinct was once acquired, if carried out to a much less extent even than in our British <i epub:type="z3998:taxonomy">F. sanguinea</i>, which, as we have seen, is less aided by its slaves than the same species in Switzerland, natural selection might increase and modify the instinct⁠—always supposing each modification to be of use to the species⁠—until an ant was formed as abjectly dependent on its slaves as is the Formica rufescens.</p>
 			</section>
 			<section id="chapter-8-5" epub:type="z3998:subchapter">
 				<h3 epub:type="title">Cell-Making Instinct of the Hive-Bee</h3>
 				<p>I will not here enter on minute details on this subject, but will merely give an outline of the conclusions at which I have arrived. He must be a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. We hear from mathematicians that bees have practically solved a recondite problem, and have made their cells of the proper shape to hold the greatest possible amount of honey, with the least possible consumption of precious wax in their construction. It has been remarked that a skilful workman, with fitting tools and measures, would find it very difficult to make cells of wax of the true form, though this is effected by a crowd of bees working in a dark hive. Granting whatever instincts you please, it seems at first quite inconceivable how they can make all the necessary angles and planes, or even perceive when they are correctly made. But the difficulty is not nearly so great as at first appears: all this beautiful work can be shown, I think, to follow from a few simple instincts.</p>
-				<p>I was led to investigate this subject by <abbr>Mr.</abbr> Waterhouse, who has shown that the form of the cell stands in close relation to the presence of adjoining cells; and the following view may, perhaps, be considered only as a modification of his theory. Let us look to the great principle of gradation, and see whether Nature does not reveal to us her method of work. At one end of a short series we have humblebees, which use their old cocoons to hold honey, sometimes adding to them short tubes of wax, and likewise making separate and very irregular rounded cells of wax. At the other end of the series we have the cells of the hive-bee, placed in a double layer: each cell, as is well known, is an hexagonal prism, with the basal edges of its six sides bevelled so as to join an inverted pyramid, of three rhombs. These rhombs have certain angles, and the three which form the pyramidal base of a single cell on one side of the comb, enter into the composition of the bases of three adjoining cells on the opposite side. In the series between the extreme perfection of the cells of the hive-bee and the simplicity of those of the humblebee, we have the cells of the Mexican Melipona domestica, carefully described and figured by Pierre Huber. The Melipona itself is intermediate in structure between the hive and humble bee, but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells, in which the young are hatched, and, in addition, some large cells of wax for holding honey. These latter cells are nearly spherical and of nearly equal sizes, and are aggregated into an irregular mass. But the important point to notice is, that these cells are always made at that degree of nearness to each other that they would have intersected or broken into each other if the spheres had been completed; but this is never permitted, the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. Hence, each cell consists of an outer spherical portion, and of two, three, or more flat surfaces, according as the cell adjoins two, three or more other cells. When one cell rests on three other cells, which, from the spheres being nearly of the same size, is very frequently and necessarily the case, the three flat surfaces are united into a pyramid; and this pyramid, as Huber has remarked, is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee. As in the cells of the hive-bee, so here, the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. It is obvious that the Melipona saves wax, and what is more important, labour, by this manner of building; for the flat walls between the adjoining cells are not double, but are of the same thickness as the outer spherical portions, and yet each flat portion forms a part of two cells.</p>
+				<p>I was led to investigate this subject by <abbr>Mr.</abbr> Waterhouse, who has shown that the form of the cell stands in close relation to the presence of adjoining cells; and the following view may, perhaps, be considered only as a modification of his theory. Let us look to the great principle of gradation, and see whether Nature does not reveal to us her method of work. At one end of a short series we have humblebees, which use their old cocoons to hold honey, sometimes adding to them short tubes of wax, and likewise making separate and very irregular rounded cells of wax. At the other end of the series we have the cells of the hive-bee, placed in a double layer: each cell, as is well known, is an hexagonal prism, with the basal edges of its six sides bevelled so as to join an inverted pyramid, of three rhombs. These rhombs have certain angles, and the three which form the pyramidal base of a single cell on one side of the comb, enter into the composition of the bases of three adjoining cells on the opposite side. In the series between the extreme perfection of the cells of the hive-bee and the simplicity of those of the humblebee, we have the cells of the Mexican <i epub:type="z3998:taxonomy">Melipona domestica</i>, carefully described and figured by Pierre Huber. The <i epub:type="z3998:taxonomy">Melipona</i> itself is intermediate in structure between the hive and humblebee, but more nearly related to the latter: it forms a nearly regular waxen comb of cylindrical cells, in which the young are hatched, and, in addition, some large cells of wax for holding honey. These latter cells are nearly spherical and of nearly equal sizes, and are aggregated into an irregular mass. But the important point to notice is, that these cells are always made at that degree of nearness to each other that they would have intersected or broken into each other if the spheres had been completed; but this is never permitted, the bees building perfectly flat walls of wax between the spheres which thus tend to intersect. Hence, each cell consists of an outer spherical portion, and of two, three, or more flat surfaces, according as the cell adjoins two, three or more other cells. When one cell rests on three other cells, which, from the spheres being nearly of the same size, is very frequently and necessarily the case, the three flat surfaces are united into a pyramid; and this pyramid, as Huber has remarked, is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee. As in the cells of the hive-bee, so here, the three plane surfaces in any one cell necessarily enter into the construction of three adjoining cells. It is obvious that the Melipona saves wax, and what is more important, labour, by this manner of building; for the flat walls between the adjoining cells are not double, but are of the same thickness as the outer spherical portions, and yet each flat portion forms a part of two cells.</p>
 				<p>Reflecting on this case, it occurred to me that if the Melipona had made its spheres at some given distance from each other, and had made them of equal sizes and had arranged them symmetrically in a double layer, the resulting structure would have been as perfect as the comb of the hive-bee. Accordingly I wrote to Professor Miller, of Cambridge, and this geometer has kindly read over the following statement, drawn up from his information, and tells me that it is strictly correct:⁠—</p>
 				<p>If a number of equal spheres be described with their centres placed in two parallel layers; with the centre of each sphere at the distance of radius x sqrt(2) or radius x 1.41421 (or at some lesser distance), from the centres of the six surrounding spheres in the same layer; and at the same distance from the centres of the adjoining spheres in the other and parallel layer; then, if planes of intersection between the several spheres in both layers be formed, there will result a double layer of hexagonal prisms united together by pyramidal bases formed of three rhombs; and the rhombs and the sides of the hexagonal prisms will have every angle identically the same with the best measurements which have been made of the cells of the hive-bee. But I hear from Professor Wyman, who has made numerous careful measurements, that the accuracy of the workmanship of the bee has been greatly exaggerated; so much so, that whatever the typical form of the cell may be, it is rarely, if ever, realised.</p>
 				<p>Hence we may safely conclude that, if we could slightly modify the instincts already possessed by the Melipona, and in themselves not very wonderful, this bee would make a structure as wonderfully perfect as that of the hive-bee. We must suppose the Melipona to have the power of forming her cells truly spherical, and of equal sizes; and this would not be very surprising, seeing that she already does so to a certain extent, and seeing what perfectly cylindrical burrows many insects make in wood, apparently by turning round on a fixed point. We must suppose the Melipona to arrange her cells in level layers, as she already does her cylindrical cells; and we must further suppose, and this is the greatest difficulty, that she can somehow judge accurately at what distance to stand from her fellow-labourers when several are making their spheres; but she is already so far enabled to judge of distance, that she always describes her spheres so as to intersect to a certain extent; and then she unites the points of intersection by perfectly flat surfaces. By such modifications of instincts which in themselves are not very wonderful⁠—hardly more wonderful than those which guide a bird to make its nest⁠—I believe that the hive-bee has acquired, through natural selection, her inimitable architectural powers.</p>
@@ -78,10 +78,10 @@
 				<p>No doubt many instincts of very difficult explanation could be opposed to the theory of natural selection⁠—cases, in which we cannot see how an instinct could have originated; cases, in which no intermediate gradations are known to exist; cases of instincts of such trifling importance, that they could hardly have been acted on by natural selection; cases of instincts almost identically the same in animals so remote in the scale of nature that we cannot account for their similarity by inheritance from a common progenitor, and consequently must believe that they were independently acquired through natural selection. I will not here enter on these several cases, but will confine myself to one special difficulty, which at first appeared to me insuperable, and actually fatal to the whole theory. I allude to the neuters or sterile females in insect communities: for these neuters often differ widely in instinct and in structure from both the males and fertile females, and yet, from being sterile, they cannot propagate their kind.</p>
 				<p>The subject well deserves to be discussed at great length, but I will here take only a single case, that of working or sterile ants. How the workers have been rendered sterile is a difficulty; but not much greater than that of any other striking modification of structure; for it can be shown that some insects and other articulate animals in a state of nature occasionally become sterile; and if such insects had been social, and it had been profitable to the community that a number should have been annually born capable of work, but incapable of procreation, I can see no especial difficulty in this having been effected through natural selection. But I must pass over this preliminary difficulty. The great difficulty lies in the working ants differing widely from both the males and the fertile females in structure, as in the shape of the thorax, and in being destitute of wings and sometimes of eyes, and in instinct. As far as instinct alone is concerned, the wonderful difference in this respect between the workers and the perfect females would have been better exemplified by the hive-bee. If a working ant or other neuter insect had been an ordinary animal, I should have unhesitatingly assumed that all its characters had been slowly acquired through natural selection; namely, by individuals having been born with slight profitable modifications, which were inherited by the offspring, and that these again varied and again were selected, and so onwards. But with the working ant we have an insect differing greatly from its parents, yet absolutely sterile; so that it could never have transmitted successively acquired modifications of structure or instinct to its progeny. It may well be asked how it is possible to reconcile this case with the theory of natural selection?</p>
 				<p>First, let it be remembered that we have innumerable instances, both in our domestic productions and in those in a state of nature, of all sorts of differences of inherited structure which are correlated with certain ages and with either sex. We have differences correlated not only with one sex, but with that short period when the reproductive system is active, as in the nuptial plumage of many birds, and in the hooked jaws of the male salmon. We have even slight differences in the horns of different breeds of cattle in relation to an artificially imperfect state of the male sex; for oxen of certain breeds have longer horns than the oxen of other breeds, relatively to the length of the horns in both the bulls and cows of these same breeds. Hence, I can see no great difficulty in any character becoming correlated with the sterile condition of certain members of insect communities; the difficulty lies in understanding how such correlated modifications of structure could have been slowly accumulated by natural selection.</p>
-				<p>This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Breeders of cattle wish the flesh and fat to be well marbled together. An animal thus characterized has been slaughtered, but the breeder has gone with confidence to the same stock and has succeeded. Such faith may be placed in the power of selection that a breed of cattle, always yielding oxen with extraordinarily long horns, could, it is probable, be formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox would ever have propagated its kind. Here is a better and real illustration: According to M. Verlot, some varieties of the double annual stock, from having been long and carefully selected to the right degree, always produce a large proportion of seedlings bearing double and quite sterile flowers, but they likewise yield some single and fertile plants. These latter, by which alone the variety can be propagated, may be compared with the fertile male and female ants, and the double sterile plants with the neuters of the same community. As with the varieties of the stock, so with social insects, selection has been applied to the family, and not to the individual, for the sake of gaining a serviceable end. Hence, we may conclude that slight modifications of structure or of instinct, correlated with the sterile condition of certain members of the community, have proved advantageous; consequently the fertile males and females have flourished, and transmitted to their fertile offspring a tendency to produce sterile members with the same modifications. This process must have been repeated many times, until that prodigious amount of difference between the fertile and sterile females of the same species has been produced which we see in many social insects.</p>
+				<p>This difficulty, though appearing insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be applied to the family, as well as to the individual, and may thus gain the desired end. Breeders of cattle wish the flesh and fat to be well marbled together. An animal thus characterized has been slaughtered, but the breeder has gone with confidence to the same stock and has succeeded. Such faith may be placed in the power of selection that a breed of cattle, always yielding oxen with extraordinarily long horns, could, it is probable, be formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox would ever have propagated its kind. Here is a better and real illustration: According to <abbr class="name">M.</abbr> Verlot, some varieties of the double annual stock, from having been long and carefully selected to the right degree, always produce a large proportion of seedlings bearing double and quite sterile flowers, but they likewise yield some single and fertile plants. These latter, by which alone the variety can be propagated, may be compared with the fertile male and female ants, and the double sterile plants with the neuters of the same community. As with the varieties of the stock, so with social insects, selection has been applied to the family, and not to the individual, for the sake of gaining a serviceable end. Hence, we may conclude that slight modifications of structure or of instinct, correlated with the sterile condition of certain members of the community, have proved advantageous; consequently the fertile males and females have flourished, and transmitted to their fertile offspring a tendency to produce sterile members with the same modifications. This process must have been repeated many times, until that prodigious amount of difference between the fertile and sterile females of the same species has been produced which we see in many social insects.</p>
 				<p>But we have not as yet touched on the acme of the difficulty; namely, the fact that the neuters of several ants differ, not only from the fertile females and males, but from each other, sometimes to an almost incredible degree, and are thus divided into two or even three castes. The castes, moreover, do not generally graduate into each other, but are perfectly well defined; being as distinct from each other as are any two species of the same genus, or rather as any two genera of the same family. Thus, in Eciton, there are working and soldier neuters, with jaws and instincts extraordinarily different: in Cryptocerus, the workers of one caste alone carry a wonderful sort of shield on their heads, the use of which is quite unknown: in the Mexican Myrmecocystus, the workers of one caste never leave the nest; they are fed by the workers of another caste, and they have an enormously developed abdomen which secretes a sort of honey, supplying the place of that excreted by the aphides, or the domestic cattle as they may be called, which our European ants guard and imprison.</p>
 				<p>It will indeed be thought that I have an overweening confidence in the principle of natural selection, when I do not admit that such wonderful and well-established facts at once annihilate the theory. In the simpler case of neuter insects all of one caste, which, as I believe, have been rendered different from the fertile males and females through natural selection, we may conclude from the analogy of ordinary variations, that the successive, slight, profitable modifications did not first arise in all the neuters in the same nest, but in some few alone; and that by the survival of the communities with females which produced most neuters having the advantageous modification, all the neuters ultimately came to be thus characterized. According to this view we ought occasionally to find in the same nest neuter-insects, presenting gradations of structure; and this we do find, even not rarely, considering how few neuter-insects out of Europe have been carefully examined. <abbr>Mr.</abbr> <abbr class="name">F.</abbr> Smith has shown that the neuters of several British ants differ surprisingly from each other in size and sometimes in colour; and that the extreme forms can be linked together by individuals taken out of the same nest: I have myself compared perfect gradations of this kind. It sometimes happens that the larger or the smaller sized workers are the most numerous; or that both large and small are numerous, while those of an intermediate size are scanty in numbers. Formica flava has larger and smaller workers, with some few of intermediate size; and, in this species, as <abbr>Mr.</abbr> <abbr class="name">F.</abbr> Smith has observed, the larger workers have simple eyes (ocelli), which, though small, can be plainly distinguished, whereas the smaller workers have their ocelli rudimentary. Having carefully dissected several specimens of these workers, I can affirm that the eyes are far more rudimentary in the smaller workers than can be accounted for merely by their proportionately lesser size; and I fully believe, though I dare not assert so positively, that the workers of intermediate size have their ocelli in an exactly intermediate condition. So that here we have two bodies of sterile workers in the same nest, differing not only in size, but in their organs of vision, yet connected by some few members in an intermediate condition. I may digress by adding, that if the smaller workers had been the most useful to the community, and those males and females had been continually selected, which produced more and more of the smaller workers, until all the workers were in this condition; we should then have had a species of ant with neuters in nearly the same condition as those of Myrmica. For the workers of Myrmica have not even rudiments of ocelli, though the male and female ants of this genus have well-developed ocelli.</p>
-				<p>I may give one other case: so confidently did I expect occasionally to find gradations of important structures between the different castes of neuters in the same species, that I gladly availed myself of <abbr>Mr.</abbr> <abbr class="name">F.</abbr> Smith’s offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. The reader will perhaps best appreciate the amount of difference in these workers by my giving, not the actual measurements, but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house, of whom many were five feet four inches high, and many sixteen feet high; but we must in addition suppose that the larger workmen had heads four instead of three times as big as those of the smaller men, and jaws nearly five times as big. The jaws, moreover, of the working ants of the several sizes differed wonderfully in shape, and in the form and number of the teeth. But the important fact for us is that, though the workers can be grouped into castes of different sizes, yet they graduate insensibly into each other, as does the widely-different structure of their jaws. I speak confidently on this latter point, as Sir J. Lubbock made drawings for me, with the camera lucida, of the jaws which I dissected from the workers of the several sizes. <abbr>Mr.</abbr> Bates, in his interesting “Naturalist on the Amazons,” has described analogous cases.</p>
+				<p>I may give one other case: so confidently did I expect occasionally to find gradations of important structures between the different castes of neuters in the same species, that I gladly availed myself of <abbr>Mr.</abbr> <abbr class="name">F.</abbr> Smith’s offer of numerous specimens from the same nest of the driver ant (Anomma) of West Africa. The reader will perhaps best appreciate the amount of difference in these workers by my giving, not the actual measurements, but a strictly accurate illustration: the difference was the same as if we were to see a set of workmen building a house, of whom many were five feet four inches high, and many sixteen feet high; but we must in addition suppose that the larger workmen had heads four instead of three times as big as those of the smaller men, and jaws nearly five times as big. The jaws, moreover, of the working ants of the several sizes differed wonderfully in shape, and in the form and number of the teeth. But the important fact for us is that, though the workers can be grouped into castes of different sizes, yet they graduate insensibly into each other, as does the widely-different structure of their jaws. I speak confidently on this latter point, as Sir <abbr class="name">J.</abbr> Lubbock made drawings for me, with the camera lucida, of the jaws which I dissected from the workers of the several sizes. <abbr>Mr.</abbr> Bates, in his interesting “Naturalist on the Amazons,” has described analogous cases.</p>
 				<p>With these facts before me, I believe that natural selection, by acting on the fertile ants or parents, could form a species which should regularly produce neuters, all of large size with one form of jaw, or all of small size with widely different jaws; or lastly, and this is the greatest difficulty, one set of workers of one size and structure, and simultaneously another set of workers of a different size and structure; a graduated series having first been formed, as in the case of the driver ant, and then the extreme forms having been produced in greater and greater numbers, through the survival of the parents which generated them, until none with an intermediate structure were produced.</p>
 				<p>An analogous explanation has been given by <abbr>Mr.</abbr> Wallace, of the equally complex case, of certain Malayan butterflies regularly appearing under two or even three distinct female forms; and by Fritz Muller, of certain Brazilian crustaceans likewise appearing under two widely distinct male forms. But this subject need not here be discussed.</p>
 				<p>I have now explained how, I believe, the wonderful fact of two distinctly defined castes of sterile workers existing in the same nest, both widely different from each other and from their parents, has originated. We can see how useful their production may have been to a social community of ants, on the same principle that the division of labour is useful to civilised man. Ants, however, work by inherited instincts and by inherited organs or tools, while man works by acquired knowledge and manufactured instruments. But I must confess, that, with all my faith in natural selection, I should never have anticipated that this principle could have been efficient in so high a degree, had not the case of these neuter insects led me to this conclusion. I have, therefore, discussed this case, at some little but wholly insufficient length, in order to show the power of natural selection, and likewise because this is by far the most serious special difficulty which my theory has encountered. The case, also, is very interesting, as it proves that with animals, as with plants, any amount of modification may be effected by the accumulation of numerous, slight, spontaneous variations, which are in any way profitable, without exercise or habit having been brought into play. For peculiar habits, confined to the workers of sterile females, however long they might be followed, could not possibly affect the males and fertile females, which alone leave descendants. I am surprised that no one has advanced this demonstrative case of neuter insects, against the well-known doctrine of inherited habit, as advanced by Lamarck.</p>

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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 Kolreuter’s 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 (Anagallis arvensis and coerulea), 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 Herbert’s 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 Crinum capense fertilised by C. revolutum 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 Crinum leads me to refer to a singular fact, namely, that individual plants of certain species of Lobelia, Verbascum and Passiflora, 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 Hippeastrum, 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 Hippeastrum aulicum 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>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 Herbert’s 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">C. 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 Lobelia, Verbascum and Passiflora, 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 Hippeastrum, 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 Hippeastrum aulicum 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 Pelargonium, Fuchsia, Calceolaria, Petunia, Rhododendron, <abbr>etc.</abbr>, have been crossed, yet many of these hybrids seed freely. For instance, Herbert asserts that a hybrid from Calceolaria integrifolia and plantaginea, 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 Rhod. ponticum and catawbiense, 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>
-				<p>Although I know of hardly any thoroughly well-authenticated cases of perfectly fertile hybrid animals, I have reason to believe that the hybrids from Cervulus vaginalis and Reevesii, and from Phasianus colchicus with P. torquatus, are perfectly fertile. M. Quatrefages states that the hybrids from two moths (Bombyx cynthia and arrindia) were proved in Paris to be fertile inter se for eight generations. It has lately been asserted that two such distinct species as the hare and rabbit, when they can be got to breed together, produce offspring, which are highly fertile when crossed with one of the parent-species. The hybrids from the common and Chinese geese (A. cygnoides), species which are so different that they are generally ranked in distinct genera, have often bred in this country with either pure parent, and in one single instance they have bred inter se. This was effected by <abbr>Mr.</abbr> Eyton, who raised two hybrids from the same parents, but from different hatches; and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. In India, however, these crossbred geese must be far more fertile; for I am assured by two eminently capable judges, namely <abbr>Mr.</abbr> Blyth and Captain Hutton, that whole flocks of these crossed geese are kept in various parts of the country; and as they are kept for profit, where neither pure parent-species exists, they must certainly be highly or perfectly fertile.</p>
+				<p>Although I know of hardly any thoroughly well-authenticated cases of perfectly fertile hybrid animals, I have reason to believe that the hybrids from <i epub:type="z3998:taxonomy">Cervulus vaginalis</i> and <i epub:type="z3998:taxonomy">Reevesii</i>, and from <i epub:type="z3998:taxonomy">Phasianus colchicus</i> with <i epub:type="z3998:taxonomy">P. torquatus</i>, are perfectly fertile. <abbr class="name">M.</abbr> Quatrefages states that the hybrids from two moths (<i epub:type="z3998:taxonomy">Bombyx cynthia</i> and <i epub:type="z3998:taxonomy">arrindia</i>) were proved in Paris to be fertile inter se for eight generations. It has lately been asserted that two such distinct species as the hare and rabbit, when they can be got to breed together, produce offspring, which are highly fertile when crossed with one of the parent-species. The hybrids from the common and Chinese geese (<i epub:type="z3998:taxonomy">A. cygnoides</i>), species which are so different that they are generally ranked in distinct genera, have often bred in this country with either pure parent, and in one single instance they have bred inter se. This was effected by <abbr>Mr.</abbr> Eyton, who raised two hybrids from the same parents, but from different hatches; and from these two birds he raised no less than eight hybrids (grandchildren of the pure geese) from one nest. In India, however, these crossbred geese must be far more fertile; for I am assured by two eminently capable judges, namely <abbr>Mr.</abbr> Blyth and Captain Hutton, that whole flocks of these crossed geese are kept in various parts of the country; and as they are kept for profit, where neither pure parent-species exists, they must certainly be highly or perfectly fertile.</p>
 				<p>With our domesticated animals, the various races when crossed together are quite fertile; yet in many cases they are descended from two or more wild species. From this fact we must conclude either that the aboriginal parent-species at first produced perfectly fertile hybrids, or that the hybrids subsequently reared under domestication became quite fertile. This latter alternative, which was first propounded by Pallas, seems by far the most probable, and can, indeed, hardly be doubted. It is, for instance, almost certain that our dogs are descended from several wild stocks; yet, with perhaps the exception of certain indigenous domestic dogs of South America, all are quite fertile together; but analogy makes me greatly doubt, whether the several aboriginal species would at first have freely bred together and have produced quite fertile hybrids. So again I have lately acquired decisive evidence that the crossed offspring from the Indian humped and common cattle are inter se perfectly fertile; and from the observations by Rutimeyer on their important osteological differences, as well as from those by <abbr>Mr.</abbr> Blyth on their differences in habits, voice, constitution, <abbr>etc.</abbr>, these two forms must be regarded as good and distinct species. The same remarks may be extended to the two chief races of the pig. We must, therefore, either give up the belief of the universal sterility of species when crossed; or we must look at this sterility in animals, not as an indelible characteristic, but as one capable of being removed by domestication.</p>
 				<p>Finally, considering all the ascertained facts on the intercrossing of plants and animals, it may be concluded that some degree of sterility, both in first crosses and in hybrids, is an extremely general result; but that it cannot, under our present state of knowledge, be considered as absolutely universal.</p>
 			</section>
@@ -37,9 +37,9 @@
 				<p>It has been already remarked, that the degree of fertility, both of first crosses and of hybrids, graduates from zero to perfect fertility. It is surprising in how many curious ways this gradation can be shown; but only the barest outline of the facts can here be given. When pollen from a plant of one family is placed on the stigma of a plant of a distinct family, it exerts no more influence than so much inorganic dust. From this absolute zero of fertility, the pollen of different species applied to the stigma of some one species of the same genus, yields a perfect gradation in the number of seeds produced, up to nearly complete or even quite complete fertility; and, as we have seen, in certain abnormal cases, even to an excess of fertility, beyond that which the plant’s own pollen produces. So in hybrids themselves, there are some which never have produced, and probably never would produce, even with the pollen of the pure parents, a single fertile seed: but in some of these cases a first trace of fertility may be detected, by the pollen of one of the pure parent-species causing the flower of the hybrid to wither earlier than it otherwise would have done; and the early withering of the flower is well known to be a sign of incipient fertilisation. From this extreme degree of sterility we have self-fertilised hybrids producing a greater and greater number of seeds up to perfect fertility.</p>
 				<p>The hybrids raised from two species which are very difficult to cross, and which rarely produce any offspring, are generally very sterile; but the parallelism between the difficulty of making a first cross, and the sterility of the hybrids thus produced⁠—two classes of facts which are generally confounded together⁠—is by no means strict. There are many cases, in which two pure species, as in the genus Verbascum, can be united with unusual facility, and produce numerous hybrid offspring, yet these hybrids are remarkably sterile. On the other hand, there are species which can be crossed very rarely, or with extreme difficulty, but the hybrids, when at last produced, are very fertile. Even within the limits of the same genus, for instance in Dianthus, these two opposite cases occur.</p>
 				<p>The fertility, both of first crosses and of hybrids, is more easily affected by unfavourable conditions, than is that of pure species. But the fertility of first crosses is likewise innately variable; for it is not always the same in degree when the same two species are crossed under the same circumstances; it depends in part upon the constitution of the individuals which happen to have been chosen for the experiment. So it is with hybrids, for their degree of fertility is often found to differ greatly in the several individuals raised from seed out of the same capsule and exposed to the same conditions.</p>
-				<p>By the term systematic affinity is meant, the general resemblance between species in structure and constitution. Now the fertility of first crosses, and of the hybrids produced from them, is largely governed by their systematic affinity. This is clearly shown by hybrids never having been raised between species ranked by systematists in distinct families; and on the other hand, by very closely allied species generally uniting with facility. But the correspondence between systematic affinity and the facility of crossing is by no means strict. A multitude of cases could be given of very closely allied species which will not unite, or only with extreme difficulty; and on the other hand of very distinct species which unite with the utmost facility. In the same family there may be a genus, as Dianthus, in which very many species can most readily be crossed; and another genus, as Silene, in which the most persevering efforts have failed to produce between extremely close species a single hybrid. Even within the limits of the same genus, we meet with this same difference; for instance, the many species of Nicotiana have been more largely crossed than the species of almost any other genus; but Gartner found that N. acuminata, which is not a particularly distinct species, obstinately failed to fertilise, or to be fertilised, by no less than eight other species of Nicotiana. Many analogous facts could be given.</p>
+				<p>By the term systematic affinity is meant, the general resemblance between species in structure and constitution. Now the fertility of first crosses, and of the hybrids produced from them, is largely governed by their systematic affinity. This is clearly shown by hybrids never having been raised between species ranked by systematists in distinct families; and on the other hand, by very closely allied species generally uniting with facility. But the correspondence between systematic affinity and the facility of crossing is by no means strict. A multitude of cases could be given of very closely allied species which will not unite, or only with extreme difficulty; and on the other hand of very distinct species which unite with the utmost facility. In the same family there may be a genus, as Dianthus, in which very many species can most readily be crossed; and another genus, as Silene, in which the most persevering efforts have failed to produce between extremely close species a single hybrid. Even within the limits of the same genus, we meet with this same difference; for instance, the many species of Nicotiana have been more largely crossed than the species of almost any other genus; but Gartner found that <i epub:type="z3998:taxonomy">N. acuminata</i>, which is not a particularly distinct species, obstinately failed to fertilise, or to be fertilised, by no less than eight other species of Nicotiana. Many analogous facts could be given.</p>
 				<p>No one has been able to point out what kind or what amount of difference, in any recognisable character, is sufficient to prevent two species crossing. It can be shown that plants most widely different in habit and general appearance, and having strongly marked differences in every part of the flower, even in the pollen, in the fruit, and in the cotyledons, can be crossed. Annual and perennial plants, deciduous and evergreen trees, plants inhabiting different stations and fitted for extremely different climates, can often be crossed with ease.</p>
-				<p>By a reciprocal cross between two species, I mean the case, for instance, of a female-ass being first crossed by a stallion, and then a mare by a male-ass: these two species may then be said to have been reciprocally crossed. There is often the widest possible difference in the facility of making reciprocal crosses. Such cases are highly important, for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity, that is of any difference in their structure or constitution, excepting in their reproductive systems. The diversity of the result in reciprocal crosses between the same two species was long ago observed by Kolreuter. To give an instance: Mirabilis jalapa can easily be fertilised by the pollen of M. longiflora, and the hybrids thus produced are sufficiently fertile; but Kolreuter tried more than two hundred times, during eight following years, to fertilise reciprocally M. longiflora with the pollen of M. jalapa, and utterly failed. Several other equally striking cases could be given. Thuret has observed the same fact with certain seaweeds or Fuci. Gartner, moreover, found that this difference of facility in making reciprocal crosses is extremely common in a lesser degree. He has observed it even between closely related forms (as Matthiola annua and glabra) which many botanists rank only as varieties. It is also a remarkable fact that hybrids raised from reciprocal crosses, though of course compounded of the very same two species, the one species having first been used as the father and then as the mother, though they rarely differ in external characters, yet generally differ in fertility in a small, and occasionally in a high degree.</p>
+				<p>By a reciprocal cross between two species, I mean the case, for instance, of a female-ass being first crossed by a stallion, and then a mare by a male-ass: these two species may then be said to have been reciprocally crossed. There is often the widest possible difference in the facility of making reciprocal crosses. Such cases are highly important, for they prove that the capacity in any two species to cross is often completely independent of their systematic affinity, that is of any difference in their structure or constitution, excepting in their reproductive systems. The diversity of the result in reciprocal crosses between the same two species was long ago observed by Kolreuter. To give an instance: Mirabilis jalapa can easily be fertilised by the pollen of <i epub:type="z3998:taxonomy">M. longiflora</i>, and the hybrids thus produced are sufficiently fertile; but Kolreuter tried more than two hundred times, during eight following years, to fertilise reciprocally <i epub:type="z3998:taxonomy">M. longiflora</i> with the pollen of <i epub:type="z3998:taxonomy">M. jalapa</i>, and utterly failed. Several other equally striking cases could be given. Thuret has observed the same fact with certain seaweeds or Fuci. Gartner, moreover, found that this difference of facility in making reciprocal crosses is extremely common in a lesser degree. He has observed it even between closely related forms (as Matthiola annua and glabra) which many botanists rank only as varieties. It is also a remarkable fact that hybrids raised from reciprocal crosses, though of course compounded of the very same two species, the one species having first been used as the father and then as the mother, though they rarely differ in external characters, yet generally differ in fertility in a small, and occasionally in a high degree.</p>
 				<p>Several other singular rules could be given from Gartner: for instance, some species have a remarkable power of crossing with other species; other species of the same genus have a remarkable power of impressing their likeness on their hybrid offspring; but these two powers do not at all necessarily go together. There are certain hybrids which, instead of having, as is usual, an intermediate character between their two parents, always closely resemble one of them; and such hybrids, though externally so like one of their pure parent-species, are with rare exceptions extremely sterile. So again among hybrids which are usually intermediate in structure between their parents, exceptional and abnormal individuals sometimes are born, which closely resemble one of their pure parents; and these hybrids are almost always utterly sterile, even when the other hybrids raised from seed from the same capsule have a considerable degree of fertility. These facts show how completely the fertility of a hybrid may be independent of its external resemblance to either pure parent.</p>
 				<p>Considering the several rules now given, which govern the fertility of first crosses and of hybrids, we see that when forms, which must be considered as good and distinct species, are united, their fertility graduates from zero to perfect fertility, or even to fertility under certain conditions in excess; that their fertility, besides being eminently susceptible to favourable and unfavourable conditions, is innately variable; that it is by no means always the same in degree in the first cross and in the hybrids produced from this cross; that the fertility of hybrids is not related to the degree in which they resemble in external appearance either parent; and lastly, that the facility of making a first cross between any two species is not always governed by their systematic affinity or degree of resemblance to each other. This latter statement is clearly proved by the difference in the result of reciprocal crosses between the same two species, for, according as the one species or the other is used as the father or the mother, there is generally some difference, and occasionally the widest possible difference, in the facility of effecting an union. The hybrids, moreover, produced from reciprocal crosses often differ in fertility.</p>
 				<p>Now do these complex and singular rules indicate that species have been endowed with sterility simply to prevent their becoming confounded in nature? I think not. For why should the sterility be so extremely different in degree, when various species are crossed, all of which we must suppose it would be equally important to keep from blending together? Why should the degree of sterility be innately variable in the individuals of the same species? Why should some species cross with facility and yet produce very sterile hybrids; and other species cross with extreme difficulty, and yet produce fairly fertile hybrids? Why should there often be so great a difference in the result of a reciprocal cross between the same two species? Why, it may even be asked, has the production of hybrids been permitted? To grant to species the special power of producing hybrids, and then to stop their further propagation by different degrees of sterility, not strictly related to the facility of the first union between their parents, seems a strange arrangement.</p>
@@ -77,7 +77,7 @@
 				<p>I have as yet spoken as if the varieties of the same species were invariably fertile when intercrossed. But it is impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases, which I will briefly abstract. The evidence is at least as good as that from which we believe in the sterility of a multitude of species. The evidence is also derived from hostile witnesses, who in all other cases consider fertility and sterility as safe criterions of specific distinction. Gartner kept, during several years, a dwarf kind of maize with yellow seeds, and a tall variety with red seeds growing near each other in his garden; and although these plants have separated sexes, they never naturally crossed. He then fertilised thirteen flowers of the one kind with pollen of the other; but only a single head produced any seed, and this one head produced only five grains. Manipulation in this case could not have been injurious, as the plants have separated sexes. No one, I believe, has suspected that these varieties of maize are distinct species; and it is important to notice that the hybrid plants thus raised were themselves <em>perfectly</em> fertile; so that even Gartner did not venture to consider the two varieties as specifically distinct.</p>
 				<p>Girou de Buzareingues crossed three varieties of gourd, which like the maize has separated sexes, and he asserts that their mutual fertilisation is by so much the less easy as their differences are greater. How far these experiments may be trusted, I know not; but the forms experimented on are ranked by Sagaret, who mainly founds his classification by the test of infertility, as varieties, and Naudin has come to the same conclusion.</p>
 				<p>The following case is far more remarkable, and seems at first incredible; but it is the result of an astonishing number of experiments made during many years on nine species of Verbascum, by so good an observer and so hostile a witness as Gartner: namely, that the yellow and white varieties when crossed produce less seed than the similarly coloured varieties of the same species. Moreover, he asserts that, when yellow and white varieties of one species are crossed with yellow and white varieties of a <em>distinct</em> species, more seed is produced by the crosses between the similarly coloured flowers, than between those which are differently coloured. <abbr>Mr.</abbr> Scott also has experimented on the species and varieties of Verbascum; and although unable to confirm Gartner’s results on the crossing of the distinct species, he finds that the dissimilarly coloured varieties of the same species yield fewer seeds, in the proportion of eighty-six to 100, than the similarly coloured varieties. Yet these varieties differ in no respect, except in the colour of their flowers; and one variety can sometimes be raised from the seed of another.</p>
-				<p>Kolreuter, whose accuracy has been confirmed by every subsequent observer, has proved the remarkable fact that one particular variety of the common tobacco was more fertile than the other varieties, when crossed with a widely distinct species. He experimented on five forms which are commonly reputed to be varieties, and which he tested by the severest trial, namely, by reciprocal crosses, and he found their mongrel offspring perfectly fertile. But one of these five varieties, when used either as the father or mother, and crossed with the Nicotiana glutinosa, always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with N. glutinosa. Hence the reproductive system of this one variety must have been in some manner and in some degree modified.</p>
+				<p>Kolreuter, whose accuracy has been confirmed by every subsequent observer, has proved the remarkable fact that one particular variety of the common tobacco was more fertile than the other varieties, when crossed with a widely distinct species. He experimented on five forms which are commonly reputed to be varieties, and which he tested by the severest trial, namely, by reciprocal crosses, and he found their mongrel offspring perfectly fertile. But one of these five varieties, when used either as the father or mother, and crossed with the Nicotiana glutinosa, always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with <i epub:type="z3998:taxonomy">N. glutinosa</i>. Hence the reproductive system of this one variety must have been in some manner and in some degree modified.</p>
 				<p>From these facts it can no longer be maintained that varieties when crossed are invariably quite fertile. From the great difficulty of ascertaining the infertility of varieties in a state of nature, for a supposed variety, if proved to be infertile in any degree, would almost universally be ranked as a species; from man attending only to external characters in his domestic varieties, and from such varieties not having been exposed for very long periods to uniform conditions of life; from these several considerations we may conclude that fertility does not constitute a fundamental distinction between varieties and species when crossed. The general sterility of crossed species may safely be looked at, not as a special acquirement or endowment, but as incidental on changes of an unknown nature in their sexual elements.</p>
 			</section>
 			<section id="chapter-9-6" epub:type="z3998:subchapter">

src/epub/text/introduction.xhtml

@@ -9,7 +9,7 @@
 		<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 C. 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>
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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">Hist. Nat. des Animaux sans Vertebres</i>. In these works he <query>up holds</query> 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="../text/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">C’est donc un probleme a reserver entierement a l’avenir, suppose meme que l’avenir 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> W. 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> Matthew’s 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>
 			<p>Rafinesque, in his <i epub:type="se:name.publication.book">New Flora of North America</i>, published in <time datetime="1836">1836</time>, wrote (page 6) as follows: “All species might have been varieties once, and many varieties are gradually becoming species by assuming constant and peculiar characters;” but further on (page 18) he adds, “except the original types or ancestors of the genus.”</p>
-			<p>In <time datetime="1843">1843</time>⁠–⁠<time datetime="1844">44</time> Professor Haldeman (<i epub:type="se:name.publication.journal">Boston Journal of Nat. Hist. U. States</i>, vol. <span epub:type="z3998:roman">iv</span>, page 468) has ably given the arguments for and against the hypothesis of the development and modification of species: he seems to lean toward the side of change.</p>
+			<p>In <time datetime="1843">1843</time>⁠–⁠<time datetime="1844">44</time> Professor Haldeman (<i epub:type="se:name.publication.journal">Boston Journal of <abbr>Nat.</abbr> <abbr>Hist.</abbr> <abbr>U.</abbr> States</i>, vol. <span epub:type="z3998:roman">iv</span>, page 468) has ably given the arguments for and against the hypothesis of the development and modification of species: he seems to lean toward the side of change.</p>
 			<p>The <i epub:type="se:name.publication.book">Vestiges of Creation</i> appeared in <time datetime="1844">1844</time>. In the tenth and much improved edition (<time datetime="1853">1853</time>) the anonymous author says (page 155): “The proposition determined on after much consideration is, that the several series of animated beings, from the simplest and oldest up to the highest and most recent, are, under the providence of God, the results, <b>first</b>, of an impulse which has been imparted to the forms of life, advancing them, in definite times, by generation, through grades of organisation terminating in the highest dicotyledons and vertebrata, these grades being few in number, and generally marked by intervals of organic character, which we find to be a practical difficulty in ascertaining affinities; <b>second</b>, of another impulse connected with the vital forces, tending, in the course of generations, to modify organic structures in accordance with external circumstances, as food, the nature of the habitat, and the meteoric agencies, these being the ‘adaptations’ of the natural theologian.” The author apparently believes that organisation progresses by sudden leaps, but that the effects produced by the conditions of life are gradual. He argues with much force on general grounds that species are not immutable productions. But I cannot see how the two supposed “impulses” account in a scientific sense for the numerous and beautiful coadaptations which we see throughout nature; I cannot see that we thus gain any insight how, for instance, a woodpecker has become adapted to its peculiar habits of life. The work, from its powerful and brilliant style, though displaying in the early editions little accurate knowledge and a great want of scientific caution, immediately had a very wide circulation. In my opinion it has done excellent service in this country in calling attention to the subject, in removing prejudice, and in thus preparing the ground for the reception of analogous views.</p>
-			<p>In <time datetime="1846">1846</time> the veteran geologist M.J. d’Omalius d’Halloy published in an excellent though short paper (<i epub:type="se:name.publication.journal">Bulletins de l’Acad. Roy. Bruxelles</i>, tom. <span epub:type="z3998:roman">xiii</span>, page 581) his opinion that it is more probable that new species have been produced by descent with modification than that they have been separately created: the author first promulgated this opinion in <time datetime="1831">1831</time>.</p>
+			<p>In <time datetime="1846">1846</time> the veteran geologist <abbr class="name">M. J.</abbr> d’Omalius d’Halloy published in an excellent though short paper (<i epub:type="se:name.publication.journal">Bulletins de l’Acad. Roy. Bruxelles</i>, tom. <span epub:type="z3998:roman">xiii</span>, page 581) his opinion that it is more probable that new species have been produced by descent with modification than that they have been separately created: the author first promulgated this opinion in <time datetime="1831">1831</time>.</p>
 			<p>Professor Owen, in <time datetime="1849">1849</time> (<i epub:type="se:name.publication.book">Nature of Limbs</i>, page 86), wrote as follows: “The archetypal idea was manifested in the flesh under diverse such modifications, upon this planet, long prior to the existence of those animal species that actually exemplify it. To what natural laws or secondary causes the orderly succession and progression of such organic phenomena may have been committed, we, as yet, are ignorant.” In his address to the British Association, in <time datetime="1858">1858</time>, he speaks (page li) of “the axiom of the continuous operation of creative power, or of the ordained becoming of living things.” Further on (page xc), after referring to geographical distribution, he adds, “These phenomena shake our confidence in the conclusion that the Apteryx of New Zealand and the Red Grouse of England were distinct creations in and for those islands respectively. Always, also, it may be well to bear in mind that by the word ‘creation’ the zoologist means ‘a process he knows not what.’ ” He amplifies this idea by adding that when such cases as that of the Red Grouse are “enumerated by the zoologist as evidence of distinct creation of the bird in and for such islands, he chiefly expresses that he knows not how the Red Grouse came to be there, and there exclusively; signifying also, by this mode of expressing such ignorance, his belief that both the bird and the islands owed their origin to a great first Creative Cause.” If we interpret these sentences given in the same address, one by the other, it appears that this eminent philosopher felt in <time datetime="1858">1858</time> his confidence shaken that the Apteryx and the Red Grouse first appeared in their respective homes “he knew not how,” or by some process “he knew not what.”</p>
 			<p>This address was delivered after the papers by <abbr>Mr.</abbr> Wallace and myself on the Origin of Species, presently to be referred to, had been read before the Linnean Society. When the first edition of this work was published, I was so completely deceived, as were many others, by such expressions as “the continuous operation of creative power,” that I included Professor Owen with other palaeontologists as being firmly convinced of the immutability of species; but it appears (<i epub:type="se:name.publication.book">Anat. of Vertebrates</i>, vol. <span epub:type="z3998:roman">iii</span>, page 796) that this was on my part a preposterous error. In the last edition of this work I inferred, and the inference still seems to me perfectly just, from a passage beginning with the words “no doubt the type-form,” <abbr>etc.</abbr>(Ibid., vol. i, page <span epub:type="z3998:roman">xxxv</span>), that Professor Owen admitted that natural selection may have done something in the formation of a new species; but this it appears (Ibid., vol. <span epub:type="z3998:roman">iii</span> page 798) is inaccurate and without evidence. I also gave some extracts from a correspondence between Professor Owen and the editor of the <i epub:type="se:name.publication.journal">London Review</i>, from which it appeared manifest to the editor as well as to myself, that Professor Owen claimed to have promulgated the theory of natural selection before I had done so; and I expressed my surprise and satisfaction at this announcement; but as far as it is possible to understand certain recently published passages (Ibid., vol. <span epub:type="z3998:roman">iii</span> page 798) I have either partially or wholly again fallen into error. It is consolatory to me that others find Professor Owen’s controversial writings as difficult to understand and to reconcile with each other, as I do. As far as the mere enunciation of the principle of natural selection is concerned, it is quite immaterial whether or not Professor Owen preceded me, for both of us, as shown in this historical sketch, were long ago preceded by <abbr>Dr.</abbr> Wells and <abbr>Mr.</abbr> Matthews.</p>
 			<p>M. Isidore Geoffroy Saint-Hilaire, in his lectures delivered in <time datetime="1850">1850</time> (of which a Resume appeared in the <i epub:type="se:name.publication.journal">Revue et Mag. de Zoolog.</i>, <time datetime="1851-01"><abbr>Jan.</abbr>, 1851</time>), briefly gives his reason for believing that specific characters “<i xml:lang="fr">sont fixes, pour chaque espece, tant qu’elle se perpetue au milieu des memes circonstances: ils se modifient, si les circonstances ambiantes viennent a changer. En resume, <b>L’observation</b> des animaux sauvages demontre deja la variabilite <b>limitee</b> des especes. Les <b>experiences</b> sur les animaux sauvages devenus domestiques, et sur les animaux domestiques redevenus sauvages, la demontrent plus clairment encore. Ces memes experiences prouvent, de plus, que les differences produites peuvent etre de <b>valeur generique</b>.</i>” In his <i epub:type="se:name.publication.book">Hist. Nat. Generale</i> (tom. <span epub:type="z3998:roman">ii</span>, page 430, <time datetime="1859">1859</time>) he amplifies analogous conclusions.</p>
 			<p>From a circular lately issued it appears that <abbr>Dr.</abbr> Freke, in <time datetime="1851">1851</time> (<i epub:type="se:name.publication.journal">Dublin Medical Press</i>, page 322), propounded the doctrine that all organic beings have descended from one primordial form. His grounds of belief and treatment of the subject are wholly different from mine; but as <abbr>Dr.</abbr> Freke has now (<time datetime="1861">1861</time>) published his essay on the <i epub:type="se:name.publication.book">Origin of Species by means of Organic Affinity</i>, the difficult attempt to give any idea of his views would be superfluous on my part.</p>
 			<p><abbr>Mr.</abbr> Herbert Spencer, in an essay (originally published in the <i epub:type="se:name.publication.newspaper">Leader</i>, <time datetime="1852-03">March, 1852</time>, and republished in his <i epub:type="se:name.publication.book">Essays</i>, in <time datetime="1858">1858</time>), has contrasted the theories of the creation and the development of organic beings with remarkable skill and force. He argues from the analogy of domestic productions, from the changes which the embryos of many species undergo, from the difficulty of distinguishing species and varieties, and from the principle of general gradation, that species have been modified; and he attributes the modification to the change of circumstances. The author (<time datetime="1855">1855</time>) has also treated psychology on the principle of the necessary acquirement of each mental power and capacity by gradation.</p>
-			<p>In <time datetime="1852">1852</time> M. Naudin, a distinguished botanist, expressly stated, in an admirable paper on the Origin of Species (<i epub:type="se:name.publication.journal">Revue Horticole</i>, page 102; since partly republished in the <i epub:type="se:name.publication.journal">Nouvelles Archives du Museum</i>, tom. i, page 171), his belief that species are formed in an analogous manner as varieties are under cultivation; and the latter process he attributes to man’s power of selection. But he does not show how selection acts under nature. He believes, like Dean Herbert, that species, when nascent, were more plastic than at present. He lays weight on what he calls the principle of finality, “<i xml:lang="fr">puissance mysterieuse, indeterminee; fatalite pour les uns; pour les autres volonte providentielle, dont l’action incessante sur les etres vivantes determine, a toutes les epoques de l’existence du monde, la forme, le volume, et la duree de chacun d’eux, en raison de sa destinee dans l’ordre de choses dont il fait partie. C’est cette puissance qui harmonise chaque membre a l’ensemble, en l’appropriant à la fonction qu’il doit remplir dans l’organisme general de la nature, fonction qui est pour lui sa raison d’etre.</i>”<a href="../text/endnotes.xhtml#note-3" id="noteref-3" epub:type="noteref">3</a></p>
+			<p>In <time datetime="1852">1852</time> <abbr class="name">M.</abbr> Naudin, a distinguished botanist, expressly stated, in an admirable paper on the Origin of Species (<i epub:type="se:name.publication.journal">Revue Horticole</i>, page 102; since partly republished in the <i epub:type="se:name.publication.journal">Nouvelles Archives du Museum</i>, tom. i, page 171), his belief that species are formed in an analogous manner as varieties are under cultivation; and the latter process he attributes to man’s power of selection. But he does not show how selection acts under nature. He believes, like Dean Herbert, that species, when nascent, were more plastic than at present. He lays weight on what he calls the principle of finality, “<i xml:lang="fr">puissance mysterieuse, indeterminee; fatalite pour les uns; pour les autres volonte providentielle, dont l’action incessante sur les etres vivantes determine, a toutes les epoques de l’existence du monde, la forme, le volume, et la duree de chacun d’eux, en raison de sa destinee dans l’ordre de choses dont il fait partie. C’est cette puissance qui harmonise chaque membre a l’ensemble, en l’appropriant à la fonction qu’il doit remplir dans l’organisme general de la nature, fonction qui est pour lui sa raison d’etre.</i>”<a href="../text/endnotes.xhtml#note-3" id="noteref-3" epub:type="noteref">3</a></p>
 			<p>In <time datetime="1853">1853</time> a celebrated geologist, Count Keyserling (<i epub:type="se:name.publication.journal">Bulletin de la Soc. Geolog.</i>, 2nd Ser., tom. <span epub:type="z3998:roman">x</span>, page 357), suggested that as new diseases, supposed to have been caused by some miasma have arisen and spread over the world, so at certain periods the germs of existing species may have been chemically affected by circumambient molecules of a particular nature, and thus have given rise to new forms.</p>
 			<p>In this same year, <time datetime="1853">1853</time>, <abbr>Dr.</abbr> Schaaffhausen published an excellent pamphlet (<i epub:type="se:name.publication.book">Verhand. des Naturhist. Vereins der Preuss. Rheinlands</i>, <abbr>etc.</abbr>), in which he maintains the development of organic forms on the earth. He infers that many species have kept true for long periods, whereas a few have become modified. The distinction of species he explains by the destruction of intermediate graduated forms. “Thus living plants and animals are not separated from the extinct by new creations, but are to be regarded as their descendants through continued reproduction.”</p>
-			<p>A well-known French botanist, M. Lecoq, writes in <time datetime="1854">1854</time> (<i epub:type="se:name.publication.book">Etudes sur Geograph</i> Bot. tom. i, page 250), “<i xml:lang="fr">On voit que nos recherches sur la fixite ou la variation de l’espece, nous conduisent directement aux idees emises par deux hommes justement celebres, Geoffroy Saint-Hilaire et Goethe.</i>” Some other passages scattered through M. Lecoq’s large work make it a little doubtful how far he extends his views on the modification of species.</p>
+			<p>A well-known French botanist, <abbr class="name">M.</abbr> Lecoq, writes in <time datetime="1854">1854</time> (<i epub:type="se:name.publication.book">Etudes sur Geograph</i> Bot. tom. i, page 250), “<i xml:lang="fr">On voit que nos recherches sur la fixite ou la variation de l’espece, nous conduisent directement aux idees emises par deux hommes justement celebres, Geoffroy Saint-Hilaire et Goethe.</i>” Some other passages scattered through <abbr class="name">M.</abbr> Lecoq’s large work make it a little doubtful how far he extends his views on the modification of species.</p>
 			<p>The “Philosophy of Creation” has been treated in a masterly manner by the <abbr>Rev.</abbr> Baden Powell, in his <i epub:type="se:name.publication.book">Essays on the Unity of Worlds</i>, <time datetime="1855">1855</time>. Nothing can be more striking than the manner in which he shows that the introduction of new species is “a regular, not a casual phenomenon,” or, as Sir John Herschel expresses it, “a natural in contradistinction to a miraculous process.”</p>
 			<p>The third volume of the <i epub:type="se:name.publication.journal">Journal of the Linnean Society</i> contains papers, read <time datetime="1858-07-01">July 1, 1858</time>, by <abbr>Mr.</abbr> Wallace and myself, in which, as stated in the introductory remarks to this volume, the theory of natural selection is promulgated by <abbr>Mr.</abbr> Wallace with admirable force and clearness.</p>
 			<p>Von Baer, toward whom all zoologists feel so profound a respect, expressed about the year <time datetime="1859">1859</time> (see <abbr>Prof.</abbr> Rudolph Wagner, <i epub:type="se:name.publication.journal">Zoologisch-Anthropologische Untersuchungen</i>, 1861, s. 51) his conviction, chiefly grounded on the laws of geographical distribution, that forms now perfectly distinct have descended from a single parent-form.</p>