On the other hand, gigas and rubrinervis, oblonga and albida obviously bear the characters of progressive elementary species. They are not differentiated from lamarckiana by one or two main features. They diverge from it in nearly all organs, and in all in a definite though small degree. They may be recognized as soon as they have developed their first leaves and remain discernible throughout life. Their characters refer chiefly to the foliage, but no less to the stature, and even the seeds have peculiarities. There can be little doubt but that all the attributes of every new species are derived from one principal change. But why this should affect the foliage in one manner, the flowers in another and the fruits in a third direction, remains obscure. To gain ever so little an insight into the nature of these changes, we may best compare the differences of our evening-primroses with those between the two hundred elementary species of Draba and other similar instances. In doing so we find the same main [566] feature, the minute differences in nearly all points.

V. The same new species are produced in a large number of individuals.

This is a very curious fact. It embraces two minor points, viz: the multitude of similar mutants in the same year, and the repetition thereof in succeeding generations. Obviously there must be some common cause. This cause must be assumed to lie dormant in the Lamarckianas of my strain, and probably in all of them, as no single parent-plant proved ever to be wholly destitute of mutability. Furthermore the different causes for the sundry mutations must lie latent together in the same parent-plant. They obey the same general laws, become active under similar conditions, some of them being more easily awakened than others. The germs of the oblonga, lata and nanella are especially irritable, and are ready to spring into activity at the least summons, while those of gigas, rubrinervis and scintillans are far more difficult to arouse.

These germs must be assumed to lie dormant during many successive generations. This is especially evident in the case of lata and nanella, which appeared in the first year of the pedigree culture and which since have been repeated yearly, and have been seen to arise by mutation [567] also during the last season (1903). Only gigas appeared but once, but then there is every reason to assume that in larger sowings or by a prolongation of the experiments it might have made a second appearance.

Is the number of such germs to be supposed to be limited or unlimited? My experiment has produced about a dozen new forms. Without doubt I could easily have succeeded in getting more, if I had had any definite reason to search for them. But such figures are far from favoring the assumption of indefinite mutability. The group of possible new forms is no doubt sharply circumscribed. Partly so by the morphologic peculiarities of lamarckiana, which seem to exclude red flowers, composite leaves, etc. No doubt there are more direct reasons for these limits, some changes having taken place initially and others later, while the present mutations are only repetitions of previous ones, and do not contribute new lines of development to those already existing. This leads us to the supposition of some common original cause, which produced a number of changes, but which itself is no longer at work, but has left the affected qualities, and only these, in the state of mutability.

In nature, repeated mutations must be of far greater significance than isolated ones. How [568] great is the chance for a single individual to be destroyed in the struggle for life? Hundreds of thousands of seeds are produced by lamarckiana annually in the field, and only some slow increase of the number of specimens can be observed. Many seeds do not find the proper circumstances for germination, or the young seedlings are destroyed by lack of water, of air, or of space. Thousands of them are so crowded when becoming rosettes that only a few succeed in producing stems. Any weakness would have destroyed them. As a matter of fact they are much oftener produced in the seed than seen in the field with the usual unfavorable conditions; the careful sowing of collected seeds has given proof of this fact many times.

The experimental proof of this frequency in the origin of new types, seems to overcome many difficulties offered by the current theories on the probable origin of species at large.

VI. The relation between mutability and fluctuating variability has always been one of the chief difficulties of the followers of Darwin. The majority assumed that species arise by the slow accumulation of slight fluctuating deviations, and the mutations were only to be considered as extreme fluctuations, obtained, in the main, by a continuous selection of small differences in a constant direction.

[569] My cultures show that quite the opposite is to be regarded as fact. All organs and all qualities of lamarckiana fluctuate and vary in a more or less evident manner, and those which I had the opportunity of examining more closely were found to comply with the general laws of fluctuation. But such oscillating changes have nothing in common with the mutations. Their essential character is the heaping up of slight deviations around a mean, and the occurrence of continuous lines of increasing deviations, linking the extremes with this group. Nothing of the kind is observed in the case of mutations. There is no mean for them to be grouped around and the extreme only is to be seen, and it is wholly unconnected with the original type. It might be supposed that on closer inspection each mutation might be brought into connection with some feature of the fluctuating variability. But this is not the case. The dwarfs are not at all the extreme variants of structure, as the fluctuation of the height of the lamarckiana never decreases or even approaches that of the dwarfs. There is always a gap. The smallest specimens of the tall type are commonly the weakest, according to the general rule of the relationship between nourishment and variation, but the tallest dwarfs are of course the most robust specimens of their group. [570] Fluctuating variability, as a rule, is subject to reversion. The seeds of the extremes do not produce an offspring which fluctuates around their parents as a center, but around some point on the line which combines their attributes with the corresponding characteristic of their ancestors, as Vilmorin has put it. No reversion accompanies mutation, and this fact is perhaps the completest contrast in which these two great types of variability are opposed to each other.

The offspring of my mutants are, of course, subject to the general laws of fluctuating variability. They vary, however, around their own mean, and this mean is simply the type of the new elementary species.

VII. The mutations take place in nearly all directions.

Many authors assume that the origin of species is directed by unknown causes. These causes are assumed to work in each single case for the improvement of the animals and plants, changing them in a manner corresponding in a useful way to the changes that take place in their environment. It is not easy to imagine the nature of these influences nor how they would bring about the desired effect.

This difficulty was strongly felt by Darwin, and one of the chief purposes of his selection theory may be said to have been the attempt [571] to surmount it. Darwin tried to replace the unknown cause by natural agencies, which lie under our immediate observation. On this point Darwin was superior to his predecessors, and it is chiefly due to the clear conception of this point that his theory has gained its deserved general acceptance. According to Darwin, changes occur in all directions, quite independently of the prevailing circumstances. Some may be favorable, others detrimental, many of them without significance, neither useful nor injurious. Some of them will sooner or later be destroyed, while others will survive, but which of them will survive, is obviously dependent upon whether their particular changes agree with the existing environic conditions or not. This is what Darwin has called the struggle for life. It is a large sieve, and it only acts as such. Some fall through and are annihilated, others remain above and are selected, as the phrase goes. Many are selected, but more are destroyed; daily observation does not leave any doubt upon this point.

How the differences originate is quite another question. It has nothing to do with the theory of natural selection nor with the struggle for life. These have an active part only in the accumulation of useful qualities, and only in so [572] far as they protect the bearers of such characters against being crowded out by their more poorly constituted competitors.

However, the differentiating characteristics of elementary species are only very small. How widely distant they are from the beautiful adaptative organizations of orchids, of insectivorous plants and of so many others! Here the difference lies in the accumulation of numerous elementary characters, which all contribute to the same end. Chance must have produced them, and this would seem absolutely improbable, even impossible, were it not for Darwin's ingenious theory. Chance there is, but no more than anywhere else. It is not by mere chance that the variations move in the required direction. They do go, according to Darwin's view, in all directions, or at least in many. If these include the useful ones, and if this is repeated a number of times, cumulation is possible; if not, there is simply no progression, and the type remains stable through the ages. Natural selection is continually acting as a sieve, throwing out the useless changes and retaining the real improvements. Hence the accumulation in apparently predisposed directions, and hence the increasing adaptations to the more specialized conditions of life. It must be obvious to any one who can free himself from the current ideas, [573] that this theory of natural selection leaves the question as to how the changes themselves are brought about, quite undecided. There are two possibilities, and both have been propounded by Darwin. One is the accumulation of the slight deviations of fluctuating variability, the other consists of successive sports or leaps taking place in the same direction.

In further lectures a critical comparison of the two views will be given. Today I have only to show that the mutations of the evening-primroses, though sudden, comply with the demands made by Darwin as to the form of variability which is to be accepted as the cause of evolution and as the origin of species.

Some of my new types are stouter and others weaker than their parents, as shown by gigas and albida. Some have broader leaves and some narrower, lata and oblonga. Some have larger flowers (gigas) or deeper yellow ones (rubrinervis), or smaller blossoms (scintillans), or of a paler hue (albida). In some the capsules are longer (rubrinervis), or thicker (gigas), or more rounded (lata), or small (oblonga), and nearly destitute of seeds (brevistylis). The unevenness of the surface of the leaves may increase as in lata, or decrease as in laevifolia. The tendency to become annual prevails in rubrinervis, but gigas tends to become [574] biennial. Some are rich in pollen, while scintillans is poor. Some have large seeds, others small. Lata has become pistillate, while brevistylis has nearly lost the faculty to produce seeds. Some undescribed forms were quite sterile, and some I observed which produced no flowers at all. From this statement it may be seen that nearly all qualities vary in opposite directions and that our group of mutants affords wide material for the sifting process of natural selection. On the original field the laevifolia and brevistylis have held their own during sixteen years and probably more, without, however, being able to increase their numbers to any noticeable extent. Others perish as soon as they make their appearance, or a few individuals are allowed to bloom, but probably leave no progeny.

But perhaps the circumstances may change, or the whole strain may be dispersed and spread to new localities with different conditions. Some of the latter might be found to be favorable to the robust gigas, or to rubrinervis, which requires a drier air, with rainfall in the springtime and sunshine during the summer. It would be worth while to see whether the climate of California, where neither O. lamarckiana nor O. biennis are found wild, would not exactly [575] suit the requirements of the new species rubrinervis and gigas.

NOTE. Oenotheras are native to America and all of the species growing in Europe have escaped from gardens directly, or may have arisen by mutation, or by hybridization of introduced species. A fixed hybrid between O. cruciata and O. biennis constituting a species has been in cultivation for many years. The form known as O. biennis in Europe, and used by de Vries in all of the experiments described in these lectures, has not yet been found growing wild in America and is not identical with the species bearing that name among American botanists. Concerning this matter Professor de Vries writes under date of Sept. 12, 1905: "The 'biennis' which I collected in America has proved to be a motley collection of forms, which at that time I had no means of distinguishing. No one of them, so far as they are now growing in my garden is identical with our biennis of the sand dunes." The same appears to be the case with O. muricata. Plants from the Northeastern American seaboard, identifiable with the species do not entirely agree with those raised from seed received from Holland.

O. lamarckiana has not been found growing wild in America in recent years although the evidence at hand seems to favor the conclusion that it was seen and collected in the southern states in the last century. (See MacDougal, Vail, Shull, and Small: Mutants and Hybrids of the Oenotheras. Publication 24. Carnegie Institution. Washington, D.C., 1905.) EDITOR.



[576] LECTURE XX

THE ORIGIN OF WILD SPECIES AND VARIETIES

New species and varieties occur from time to time in the wild state. Setting aside all theoretical conceptions as to the common origin of species at large, the undoubted fact remains that new forms are sometimes met with. In the case of the peloric toad-flax the mutations are so numerous that they seem to be quite regular. The production of new species of evening-primroses was observed on the field and afterwards duplicated in the garden. There is no reason to think that these cases are isolated instances. Quite on the contrary they seem to be the prototypes of repeated occurrences in nature.

If this conception is granted, the question at once arises, how are we to deal with analogous cases, when fortune offers them, and what can we expect to learn from them?

A critical study of the existing evidence seems to be of great importance in order to ascertain the best way of dealing with new facts, and of estimating the value of the factors concerned. [577] It is manifest that we must be very careful and conservative in dealing with new facts that are brought to our attention, and every effort should be made to bring additional evidence to light. Many vegetable anomalies are so rare that they are met with only by the purest chance, and are then believed to be wholly new. When a white variety of some common plant is met with for the first time we generally assume that it originated on that very spot and only a short time previously. The discovery of a second locality for the same variety at once raises the question as to a common origin in the two instances. Could not the plants of the second locality have arisen from seeds transported from the first?

White varieties of many species of blue-bells and gentians are found not rarely, white-flowering plants of heather, both of Erica Tetralix and Calluna vulgaris occur on European heaths; white flowers of Brunella vulgaris, Ononis repens, Thymus vulgaris and others may be seen in many localities in the habitats of the colored species. Pelories of labiates seem to occur often in Austria, but are rare in Holland; white bilberries (Vaccinium Myrtillus) have many known localities throughout Europe, and nearly all the berry-bearing species in the large heath family are recorded as having white varieties.

[578] Are we to assume a single origin for all the representatives of such a variety, as we have done customarily for all the representatives of a wild species? Or can the same mutation have been repeated at different times and in distant localities? If a distinct mutation from a given species is once possible, why should it not occur twice or thrice?

A variety which seems to be new to us may only appear so, because the spot where it grows had hitherto escaped observation. Lychnis preslii is a smooth variety of Lychnis diurna and was observed for the first time in the year 1842 by Sekera. It grew abundantly in a grove near Munchengratz in southern Hungary. It was accompanied by the ordinary hairy type of the species. Since then it has been observed to be quite constant in the same locality, and some specimens have been collected for me there lately by Dr. Nemec, of Prague. No other native localities of this variety have been discovered, and there can be no doubt that it must have arisen from the ordinary campion near the spot where it still grows. But this change may have taken place some years before the first discovery, or perhaps one or more centuries ago. This could only be known if it could be proved that the locality had been satisfactorily investigated previously, and that the variety had not [579] been met with. Even in this case only something would be discovered about the time of the change, but nothing about its real nature.

So it is in many cases. If a variety is observed in a number of specimens at the time of its first discovery, and at a locality not studied previously, it takes the aspect of an old form of limited distribution, and little can be learned as to the circumstances under which it arose. If on the contrary it occurs in very small numbers or perhaps even in a single individual, and if the spot where it is found is located so that it could hardly have escaped previous observation, then the presumption of a recent origin seems justified.

What has to be ascertained on such occasions to give them scientific value? Three points strike me as being of the highest importance. First, the constancy of the new type; secondly, the occurrence or lack of intermediates, and last, but not least, the direct observation of a repeated production.

The first two points are easily ascertained. Whether the new type is linked with its more common supposed ancestor by intermediate steps is a query which at once strikes the botanist. It is usually recorded in such cases, and we may state at once that the general result is, that such intermediates do not occur. This is [580] of the highest importance and admits of only two explanations. One is that intermediates may be assumed to have preceded the existent developed form, and to have died out afterwards. But why should they have done so, especially in cases of recent changes? On the other hand the intermediates may be lacking because they have never existed, the change having taken place by a sudden leap, such as the mutations described in our former lectures. It is manifest that the assumption of hypothetical intermediates could only gain some probability if they had been found in some instance. Since they do not occur, the hypothesis seems wholly unsupported.

The second point is the constancy of the new type. Seeds should be saved and sown. If the plant fertilizes itself without the aid of insects, as do some evening-primroses, the seed saved from the native locality may prove wholly pure, and if it does give rise to a uniform progeny the constancy of the race may be assumed to be proved, provided that repeated trials do not bring to light any exceptions. If the offspring shows more than one type, cross-fertilization is always to be looked to as the most probable cause, and should be excluded, in order to sow pure seeds. Garden-experiments of this kind, and repeated trials, should always be combined [581] with the discovery of a presumed mutation. In many instances the authors have realized the importance of this point, and new types have been found constant from the very beginning. Many cases are known which show no reversions and even no partial reversions. This fact throws a distinct light on our first point, as it makes the hypothesis of a slow and gradual development still more improbable.

My third point is of quite another nature and has not as yet been dealt with. But as it appeals to me as the very soul of the problem, it seems necessary to describe it in some detail. It does not refer to the new type itself, nor to any of its morphologic or hereditary attributes, but directly concerns the presumed ancestors themselves.

The peloric toad-flax in my experiment was seen to arise thrice from the same strain. Three different individuals of my original race showed a tendency to produce peloric mutations, and they did so in a number of their seeds, exactly as the mutations of the evening-primroses were repeated nearly every year. Hence the inference, that whenever we find a novelty which is really of very recent date, the parent-strain which has produced it might still be in existence on the same spot. In the case of shrubs or perennials the very parents might yet be found. [582] But it seems probable, and is especially proved in the case of the evening-primroses, that all or the majority of the representatives of the whole strain have the same tendency to mutate. If this were a general rule, it would suffice to take some pure seeds from specimens of the presumed parents and to sow and multiply the individuals to such an extent that the mutation might have a chance to be repeated.

Unfortunately, this has not as yet been done, but in my opinion it should be the first effort of any one who has the good luck to discover a new wild mutation. Specimens of the parents should be transplanted into a garden and fertilized under isolated conditions. Seeds saved from the wild plant would have little worth, as they might have been partly fertilized by the new type itself.

After this somewhat lengthy discussion of the value of observations surrounding the discovery of new wild mutations, we now come to the description of some of the more interesting cases. As a first example, I will take the globular fruited shepherd's purse, described by Solms Laubach as Capsella heegeri. Professor Heeger discovered one plant with deviating fruits, in a group of common shepherd's purses in the market-place near Landau in Germany, in the fall of 1897. They were nearly spherical, [583] instead of flat and purse-shaped. Their valves were thick and fleshy, while those of the ordinary form are membranaceous and dry. The capsules hardly opened and therefore differed in this point from the shepherd's purse, which readily loosens both its valves as soon as it is ripe.

Only one plant was observed; whence it came could not be determined, nor whether it had arisen from the neighboring stock of Capsella or not. The discoverer took some seed to his garden and sent some to the botanical garden at Strassburg, of which Solms-Laubach is the director. The majority of the seeds of course were sowed naturally on the original spot. The following year some of the seeds germinated and repeated the novelty. The leaves, stems and flowers were those of the common shepherd's purse, but no decision could be reached concerning the type of this generation before the first flowers had faded and the rounded capsules had developed. Then it was seen that the heegeri came true from seed. It did so both in the gardens and on the market-place, where it was observed to have multiplied and spread in some small measure. The same was noted the following year, but then the place was covered with gravel and all the plants destroyed. It is not recorded to have been seen wild since.

[584] Intermediate forms have not been met with. Some slight reversions may occur in the autumn on the smallest and weakest lateral branches. Such reversions, however, seem to be very rare, as I have tried in vain to produce them on large and richly branched individuals, by applying all possible inducements in the form of manure and of cutting, to stimulate the production of successive generations of weaker side branches.

This constancy was proved by the experiments of Solms-Laubach, which I have repeated in my own garden during several years with seed received from him. No atavists or deviating specimens have been found among many hundreds of flowering plants.

It is important to note that within the family of the crucifers the form of the capsule and the attributes of the valves and seeds are usually considered to furnish the characteristics of genera, and this point has been elucidated at some length by Solms-Laubach. There is, however, no sufficient reason to construe a new genus on the ground of Heeger's globular fruited shepherd's purse; but as a true elementary species, and even as a good systematic species it has proved itself, and as such it is described by Solms-Laubach, who named it in honor of its discoverer.

Exactly analogous discoveries have been [586] instead of displaying a bright yellow cup. _O. cruciata_ grows in the Adirondack Mountains, in the states of New York and Vermont, and seems to be abundant there. It has been introduced into botanical gardens and yielded a number of hybrids, especially with _O. biennis and _O. lamarckiana_, and the narrow petals of the parent-species may be met with in combination with the stature and vegetative characteristics of these last named species. _O. cruciata_ has a purple foliage, while _biennis_ and _lamarckiana_ are green, and many of the hybrids may instantly be recognized by their purple color.

The curious attribute of the petals is not to be considered simply as a reduction in size. On anatomical inquiry it has been found that these narrow petals bear some characteristics which, on the normal plants, are limited to the calyx. Stomata and hairs, and the whole structure of the surface and inner tissues on some parts of these petals are exactly similar to those of the calyx, while on others they have retained the characteristics of petals. Sometimes there may even be seen by the naked eye green longitudinal stripes of calyx-like structure alternating with bright yellow petaloid parts. For these reasons the cruciata character may be considered as a case of sepalody of the petals, or of the petals being partly converted into sepals.

[587] It is worth while to note that as a monstrosity this occurrence is extremely rare throughout the whole vegetable kingdom, and only very few instances have been recorded.

Two cases of sudden mutations have come to my knowledge, producing this same anomaly in allied species. One has been already alluded to; it pertains to the common evening-primrose or _Oenothera biennis_, and one is a species belonging to another genus of the same family, the great hairy willow-herb or _Epilobium _hirsutum_. I propose to designate both new forms by the varietal name of _cruciata_, or _cruciatum_.

_Oenothera biennis cruciata_ was found in a native locality of the _O. biennis itself. It consisted of only one plant, showing in all its flowers the _cruciata_ marks. In all other respects it resembled wholly the _biennis_, especially in the pure green color of its foliage, which at once excluded all suspicion of hybrid origin with the purple _O. cruciata_. Moreover in our country this last occurs only in the cultivated state in botanical gardens.

Intermediates were not seen, and as the plant bore some pods, it was possible to test its constancy. I raised about 500 plants from its seeds, out of which more than 100 flowered in the first year. The others were partly kept through the winter and flowered next year. Seeds saved in [588] both seasons were sown on a large scale. Both the first and the succeeding generations of the offspring of the original plant came true without any exception. Intermediates are often found in hybrid cultures, and in them the character is a very variable one, but as yet they were not met with in progeny of this mutant. All these plants were exactly like O. biennis, with the single exception of their petals.

Epilobium hirsutum cruciatum was discovered by John Rasor near Woolpit, Bury St. Edmunds, in England. It flowered in one spot, producing about a dozen stems, among large quantities of the parent-species, which is very common there, as it is elsewhere in Europe. This species is a perennial, multiplying itself by underground runners, and the stems of the new variety were observed to stand so close to each other that they might be considered as the shoots of one individual. In this case this specimen might probably be the original mutant, as the variety had not been seen on that spot in previous years, even as it has not been found elsewhere in the vicinity.

Intermediates were not observed, though the difference is a very striking one. In the cruciate flowers the broad and bright purple petals seem at first sight to be wholly wanting. They are too weak to expand and to reflex the calyx [589] as in the normal flowers of the species. The sepals adhere to one another, and are only opened at their summit by the protruding pistils. Even the stamens hardly come to light. At the period of full bloom the flowers convey only the idea of closed buds crowned by the conspicuous white cross of the stigma. Any intermediate form would have at once betrayed itself by larger colored petals, coming out of the calyx-sheath. The cruciate petals are small and linear and greenish, recalling thereby the color of the sepals.

Mr. Rasor having sent me some flowers and some ripe capsules of his novelty, I sowed the latter in my experimental garden, where the plant flowered in large numbers and with many thousands of flowers both in 1902 and 1903. All of these plants and all of these flowers repeated the cruciate type exactly, and not the slightest impurity or tendency to partial reversion has been observed.

Thus true and constant cruciate varieties have been produced from accidentally observed initial plants, and because of their very curious characters they will no doubt be kept in botanical gardens, even if they should eventually become lost in their native localities.

At this point I might note another observation made on the wild species of Oenothera cruciata [590] from the Adirondacks. Through the kindness of Dr. MacDougal, of the New York Botanical Garden, I received seeds from Sandy Hill near Lake George. When the plants, grown from these seeds, flowered, they were not a uniform lot, but exhibited two distinct types. Some had linear petals and thin flower-buds, and in others the petals were a little broader and the buds more swollen. The difference was small, but constant on all the flowers, each single plant clearly belonging to one or the other of the two types. Probably two elementary species were intermixed here, but whether one is the systematic type and the other a mutation, remains to be seen.

Nor seem these two types to exhaust the range of variability of Oenothera cruciata. Dr. B.L. Robinson of Cambridge, Mass., had the kindness to send me seeds from another locality in the same region. The seeds were collected in New Hampshire and in my garden produced a true and constant cruciata, but with quite different secondary characters from both the aforesaid varieties. The stems and flower-spikes and even the whole foliage were much more slender, and the calyx-tubes of the flowers were noticeably more elongated. It seems not improbable that Oenothera cruciata includes a group of lesser unities, and may prove to comprise a [591] swarm of elementary species, while the original strain might even now be still in a condition of mutability. A close scrutiny in the native region is likely to reveal many unexpected features.

A very interesting novelty has already been described in a former lecture. It is the Xanthium wootoni, discovered in the region about Las Vegas, New Mexico, by T.D.A. Cockerell. It is similar in all respects to X. commune, but the burrs are more slender and the prickles much less numerous, and mostly stouter at their base. It grows in the same localities as the X. commune, and is not recorded to occur elsewhere. Whether it is an old variety or a recent mutation it is of course impossible to decide. In a culture made in my garden from the seed sent me by Mr. Cockerell, I observed (1903) that both forms had a subvariety with brownish foliage, and, besides this, one of a pure green. Possibly this species, too, is still in a mutable condition.

Perhaps the same may be asserted concerning the beautiful shrub, Hibiscus Moscheutos, observed in quite a number of divergent types by John W. Harshberger. They grew in a small meadow at Seaside Park, New Jersey, in a locality which had been undisturbed for years. They differed from each other in nearly all the [592] organs, in size, in the diameter of the stems, which were woody in some and more fleshy in others, in the shape of the foliage and in the flowers. More than twenty types could be distinguished and seeds were saved from a number of them, in order to ascertain whether they are constant, or whether perhaps a main stem in a mutating condition might be found among them. If this should prove to be the case, the relations between the observed forms would probably be analogous to those between the O. lamarckiana and its derivatives.

Many other varieties have sprung from the type-species under similar conditions from time to time. A fern-leaved mercury, Mercurialis annua laciniata, was discovered in the year 1719 by Marchant. The type was quite new at the time and maintained itself during a series of years. The yellow deadly nightshade or Atropa Belladonna lutea was found about 1850 in the Black Forest in Germany in a single spot, and has since been multiplied by seeds. It is now dispersed in botanical gardens, and seems to be quite constant. A dwarf variety of a bean, Phaseolus lunatus, was observed to spring from the ordinary type by a sudden leap about 1895 by W.W. Tracy, and many similar cases could be given.

The annual habit is not very favorable for [593] the discovery of new forms in the wild state. New varieties may appear, but may be crowded out the first year. The chances are much greater with perennials, and still greater with shrubs or trees. A single aberrant specimen may live for years and even for centuries, and under such conditions is pretty sure to be discovered sooner or later. Hence it is no wonder that many such cases are on record. They have this in common that the original plant of the variety has been found among a vast majority of representatives of the corresponding species. Nothing of course is directly known about its origin. Intermediate links have as a rule been wanting, and the seeds, which have often been sown, have not yielded reliable results, as no care was taken to preserve the blossoms from intercrossing with their parent-forms.

Stress should be laid upon one feature of these curious occurrences. Relatively often the same novelty has been found twice or thrice, or even more frequently, and under conditions which make it very improbable that any relation between such occurrences might exist. The same mutation must have taken place more than once from the same main stem.

The most interesting of these facts are connected with the origin of the purple beech, which [594] is now so universally cultivated. I take the following statements from an interesting historical essay of Prof. Jaggi. He describes three original localities. One is near the Swiss village, Buch am Irchel, and is located on the Stammberg. During the 17th century five purple beeches are recorded to have grown on this spot. Four of them have died, but one is still alive. Seedlings have germinated around this little group, and have been mostly dug up and transplanted into neighboring gardens. Nothing is known about the real origin of these plants, but according to an old document, it seems that about the year 1190 the purple beeches of Buch were already enjoying some renown, and attracting large numbers of pilgrims, owing to some old legend. The church of Embrach is said to have been built in connection with this legend, and was a goal for pilgrimages during many centuries.

A second native locality of the purple beech is found in a forest near Sondershausen in Thuringen, Germany, where a fine group of these trees is to be seen. They were mentioned for the first time in the latter half of the eighteenth century, but must have been old specimens long before that time. The third locality seems to be of much later origin. It is a forest near Roveredo in South Tyrol, where a new [595] university is being erected. It is only a century ago that the first specimens of the purple beech were discovered there.

As it is very improbable that the two last named localities should have received their purple beeches from the first named forest, it seems reasonable to assume that the variety must have been produced at least thrice.

The purple beech is now exceedingly common in cultivation. But Jaggi succeeded in showing that all the plants owe their origin to the original trees mentioned above, and are, including nearly all cultivated specimens with the sole exception of the vicinity of Buch, probably derived from the trees in Thuringen. They are easily multiplied by grafting, and come true from seed, at least often, and in a high proportion. Whether the original trees would yield a pure progeny if fertilized by their own pollen has as yet not been tested. The young seedlings have purple seed-leaves, and may easily be selected by this character, but they seem to be always subjected in a large measure to vicinism.

Many other instances of trees and shrubs, found in accidental specimens constituting a new variety in the wild state, might be given. The oak-leaved beech has been found in a forest of Lippe-Detmold in Germany and near Versailles, [596] whence it was introduced into horticulture by Carriere. Similarly divided and cleft leaves seem to have occurred more often in the wild state, and cut-leaved hazels are recorded from Rouen in France, birches and alders from Sweden and Lapland, where both are said to have been met with in several forests. The purple barberry was found about 1830 by Bertin, near Versailles. Weeping varieties of ashes were found wild in England and in Germany, and broom-like oaks, Quercus pedunculata fastigiata, are recorded from Hessen-Darmstadt, Calabria, the Pyrenees and other localities. About the real origin of all these varieties nothing is definitely known.

The "single-leaved" strawberry is a variety often seen in botanical gardens, as it is easily propagated by its runners. It was discovered wild in Lapland at the time of Linnaeus, and appeared afterwards unexpectedly in a nursery near Versailles. This happened about the year 1760 and Duchesne tested it from seeds and found it constant. This strain, however, seems to have died out before the end of the 18th century. In a picture painted by Holbein (1495-1543), strawberry leaves can be seen agreeing exactly with the monophyllous type. The variety may thus be assumed to have arisen independently [597] at least thrice, at different periods and in distant localities.

From all these statements and a good many others which can be found in horticultural and botanical literature, it may be inferred that mutations are not so very rare in nature as is often supposed. Moreover we may conclude that it is a general rule that they are neither preceded nor accompanied by intermediate steps, and that they are ordinarily constant from seed from the first.

Why then are they not met with more often? In my opinion it is the struggle for life which is the cause of this apparent rarity; which is nothing else than the premature death of all the individuals that so vary from the common type of their species as to be incapable of development under prevailing circumstances. It is obviously without consequence whether these deviations are of a fluctuating or of a mutating nature. Hence we may conclude that useless mutations will soon die out and will disappear without leaving any progeny. Even if they are produced again and again by the same strain, but under the same unfavorable conditions, there will be no appreciable result.

Thousands of mutations may perhaps take place yearly among the plants of our immediate vicinity without any chance of being discovered. [598] We are trained to the appreciation of the differentiating marks of systematic species. When we have succeeded in discerning these as given by our local flora lists, we rest content. Meeting them again we are in the habit of greeting them with their proper names. Such is the satisfaction ensuing from this knowledge that we do not feel any inclination for further inquiry. Striking deviations, such as many varietal characters, may be remarked, but then they are considered as being of only secondary interest. Our minds are turned from the delicately shaded features which differentiate elementary species.

Even in the native field of the evening-primroses, no botanist would have discovered the rosettes with smaller or paler leaves, constituting the first signs of the new species. Only by the guidance of a distinct theoretical idea were they discovered, and having once been pointed out a closer inspection soon disclosed their number.

Variability seems to us to be very general, but very limited. The limits however, are distinctly drawn by the struggle for existence. Of course the chance for useful mutations is a very small one. We have seen that the same mutations are as a rule repeated from time to time by the same species. Now, if a useful mutation, [599] or even a wholly indifferent one, might easily be produced, it would have been so, long ago, and would at the present time simply exist as a systematic variety. If produced anew somewhere the botanist, would take it for the old variety and would omit to make any inquiry as to its local origin.

Thousands of seeds with perhaps wide circles of variability are ripened each year, but only those that belong to the existing old narrow circles survive. How different would Nature appear to us if she were free to evolve all her potentialities!

Darwin himself was struck with this lack of harmony between common observations and the probable real state of things. He discussed it in connection with the cranesbill of the Pyrenees (Geranium pyrenaicum). He described how this fine little plant, which has never been extensively cultivated, had escaped from a garden in Staffordshire and had succeeded in multiplying itself so as to occupy a large area.

In doing so it had evidently found place for an uncommonly large number of plantlets from its seeds and correspondingly it had commenced to vary in almost all organs and qualities and nearly in all imaginable directions. It displayed under these exceptional circumstances a capacity which never had been exceeded and [600] which of course would have remained concealed if its multiplication had been checked in the ordinary way.

Many species have had occasion to invade new regions and cover them with hundreds of thousands of individuals. First are to be cited those species which have been introduced from America into Europe since the time of Columbus, or from Europe into this country. Some of them have become very common. In my own country the evening-primroses and Canada fleabane or are examples, and many others could be given. They should be expected to vary under these circumstances in a larger degree. Have they done so? Manifestly they have not struck out useful new characters that would enable their bearers to found new elementary species. At least none have been observed. But poor types might have been produced, and periods of mutability might have been gone through similar to that which is now under observation for Lamarck's evening primrose in Holland.

From this discussion we may infer that the chances of discovering new mutating species are great enough to justify the utmost efforts to secure them. It is only necessary to observe large numbers of plants, grown under circumstances which allow the best opportunities for [601] all the seeds. And as nature affords such opportunities only at rare intervals, we should make use of artificial methods. Large quantities of seed should be gathered from wild plants and sowed under very favorable conditions, giving all the nourishment and space required to the young seedlings. It is recommended that they be sown under glass, either in a glass-house or protected against cold and rain by glass-frames. The same lot of seed will be seen to yield twice or thrice as many seedlings if thus protected, compared with what it would have produced when sown in the field or in the garden. I have nearly wholly given up sowing seeds in my garden, as circumstances can be controlled and determined with greater exactitude when the sowing is done in a glasshouse.

The best proof perhaps, of the unfavorable influence of external conditions for slightly deteriorated deviations is afforded by variegated leaves. Many beautiful varieties are seen in our gardens and parks, and even corn has a variety with striped leaves. They are easily reproduced, both by buds and by seeds, and they are the most ordinary of all varietal deviations. They may be expected to occur wild also. But no real variegated species, nor even good varieties with this attribute occurs in nature. [602] On the other hand occasional specimens with a single variegated leaf, or with some few of them, are actually met with, and if attention is once drawn to this question, perhaps a dozen or so instances might be brought together in a summer. But they never seem to be capable of further evolution, or of reproducing themselves sufficiently and of repeating their peculiarity in their progeny. They make their appearance, are seen during a season, and then disappear. Even this slight incompleteness of some spots on one or two leaves may be enough to be their doom.

It is a common belief that new varieties owe their origin to the direct action of external conditions and moreover it is often assumed that similar deviations must have similar causes, and that these causes may act repeatedly in the same species, or in allied, or even systematically distant genera. No doubt in the end all things must have their causes, and the same causes will lead under the same circumstances to the same results. But we are not justified in deducing a direct relation between the external conditions and the internal changes of plants. These relations may be of so remote a nature that they cannot as yet be guessed at. Therefore only direct experience may be our guide. Summing up the result of our facts and discussions [603] we may state that wild new elementary species and varieties are recorded to have appeared from time to time. Invariably this happened by sudden leaps and without intermediates. The mutants are constant when propagated by seed, and at once constitute a new race. In rare instances this may be of sufficient superiority to win a place for itself in nature, but more often it has qualities which have led to its introduction into gardens as an ornamental plant or into botanical gardens by reason of the interest afforded by their novelty, or by their anomaly.

Many more mutations may be supposed to be taking place all around us, but artificial sowings on a large scale, combined with a close examination of the seedlings and a keen appreciation of the slightest indications of deviation seem required to bring them to light.

[604] LECTURE XXI

MUTATIONS IN HORTICULTURE

It is well known that Darwin based his theory of natural selection to a large extent upon the experience of breeders. Natural and artificial selection exhibit the same general features, yet it was impossible in Darwin's time to make a critical and comparative analysis of the two processes.

In accordance with our present conception there is selection of species and selection within the species. The struggle for life determines which of a group of elementary species shall survive and which shall disappear. In agricultural practice the corresponding process is usually designated by the name of variety-testing. Within the species, or within the variety, the sieve of natural selection is constantly eliminating poor specimens and preserving those that are best adapted to live under the given conditions. Some amelioration and some local races are the result, but this does not appear to be of much importance. On the contrary, the selection [605] within the race holds a prominent place in agriculture, where it is known by the imposing term, race-breeding.

Experience and methods in horticulture differ from those in agriculture in many points. Garden varieties have been tested and separated for a long time, but neither vegetables nor flowers are known to exhibit such motley groups of types as may be seen in large forage crops.

New varieties which appear from time to time may be ornamental or otherwise in flowers, and more or less profitable than their parents in vegetables and fruits. In either case the difference is usually striking, or if not, its culture would be unprofitable.

The recognition of useful new varieties being thus made easy, the whole attention of the breeder is reduced to isolating the seeds of the mutants that are to be saved and sown separately, and this process must be repeated during a few years, in order to produce the quantity of seed that is needed for a profitable introduction of the variety into commerce. In proportion to the abundance of the harvest of each year this period is shorter for some and longer for other species.

Isolation in practice is not so simple nor so easy an affair as it is in the experimental garden. Hence we have constant and nearly unavoidable [606] cross-fertilizations with the parent form or with neighboring varieties, and consequent impurity of the new strain. This impurity we have called vicinism, and in a previous lecture have shown its effects upon the horticultural races on one hand, and on the other, on the scientific value that can be ascribed to the experience of the breeder. We have established the general rule that stability is seldom met with, but that the observed instability is always open to the objection of being the result of vicinism. Often this last agency is its sole cause; or it may be complicated with other factors without our being able to discern them.

Though our assertion that the practice of the horticulturist in producing new varieties is limited to isolation, whenever chance affords them, is theoretically valid, it is not always so. We may discern between the two chief groups of varieties. The retrograde varieties are constant, the individuals not differing more from one another than those of any ordinary species. The highly variable varieties play an important part in horticulture. Double flowers, striped flowers, variegated leaves and some others yield the most striking instances. Such forms have been included in previous lectures among the ever-sporting varieties, because their peculiar characters oscillate between two extremes, viz: [607] the new one of the variety and the corresponding character of the original species.

In such cases isolation is usually accompanied by selection: rarely has the first of a double, striped or variegated race well filled or richly striped flowers or highly spotted leaves. Usually minor degrees of the anomaly are seen first, and the breeder expects the novelty to develop its features more completely and more beautifully in subsequent generations. Some varieties need selection only in the beginning, in others the most perfect specimens must be chosen every year as seed-bearers. For striped flowers, it has been prescribed by Vilmorin, that seeds should be taken only from those with the smallest stripes, because there is always reversion. Mixed seed or seed from medium types would soon yield plants with too broad stripes, and therefore less diversified flowers.

In horticulture, new varieties, both retrograde and ever-sporting, are known to occur almost yearly. Nevertheless, not every novelty of the gardener is to be considered as a mutation in the scientific sense of the word. First of all, the novelties of perennial and woody species are to be excluded. Any extreme case of fluctuating variability may be preserved and multiplied in the vegetative way. Such types are designated [608] in horticulture as varieties, though obviously they are of quite another nature than the varieties reproduced by seed. Secondly, a large number, no doubt the greater number of novelties, are of hybrid origin. Here we may discern two cases. Hybrids may be produced by the crossing of old types, either of two old cultivated forms or newly introduced species, or ordinarily between an old and an introduced variety. Such novelties are excluded from our present discussion. Secondly, hybrids may be produced between a true, new mutation and some of the already existing varieties of the same species. Examples of this obvious and usual practice will be given further on, but it must be pointed out now that by such crosses a single mutation may produce as many novelties as there are available varieties of the same species.

Summarizing these introductory remarks we must lay stress on the fact that only a small part of the horticultural novelties are real mutations, although they do occur from time to time. If useful, they are as a rule isolated and multiplied, and if necessary, improved by selection. They are in many instances, as constant from seed as the unavoidable influence of vicinism allows them to be. Exact observations on the origin, or on the degree of constancy, are usually lacking, [609] the notes being ordinarily made for commercial purposes, and often only at the date of introduction into trade, when the preceding stages of the novelty may have been partly forgotten.

With this necessary prelude I will now give a condensed survey of the historical facts relating to the origin of new horticultural varieties. An ample description has been given recently by Korshinsky, a Russian writer, who has brought together considerable historical material as evidence of the sudden appearance of novelties throughout the whole realm of garden plants.

The oldest known, and at the same time one of the most accurately described mutations is the origin of the cut-leaved variety of the greater celandine or Chelidonium majus. This variety has been described either as such, or as a distinct species, called Chelidonium laciniatum Miller.

It is distinguished from the ordinary species, by the leaves being cut into narrow lobes, with almost linear tips, a character which is, as we have seen on a previous occasion, repeated in the petals. It is at present nearly as commonly cultivated in botanical gardens as the C. majus, and has escaped in many localities and is observed to thrive as readily as the native wild [610] plants. It was not known until a few years before the close of the 16th century. Its history has been described by the French botanist, Rose. It was seen for the first time in the garden of Sprenger, an apothecary of Heidelberg, where the C. majus had been cultivated for many years. Sprenger discovered it in the year 1590, and was struck by its peculiar and sharply deviating characters. He was anxious to know whether it was a new plant and sent specimens to Clusius and to Plater, the last of whom transmitted them to Caspar Bauhin. These botanists recognized the type as quite new and Bauhin described it some years afterwards in his Phytopinax under the name of Chelidonium majus foliis quernis, or oak-leaved celandine. The new variety soon provoked general interest and was introduced into most of the botanical gardens of Europe. It was recognized as quite new, and repeated search has been made for it in a wild state, but in vain. No other origin has been discovered than that of Sprenger's garden. Afterwards it became naturalized in England and elsewhere, but there is not the least doubt as to its derivation in all the observed cases.

Hence its origin at Heidelberg is to be considered as historically proven, and it is of course only legitimate to assume that it originated in [611] the year 1590 from the seeds of the C. majus. Nevertheless, this was not ascertained by Sprenger, and some doubt as to a possible introduction from elsewhere might arise. If not, then the mutation must have been sudden, occurring without visible preparation and without the appearance of intermediates.

From the very first, the cut-leaved celandine has been constant from seed. Or at least it has been propagated by seed largely and without difficulty. Nothing, however, is known about it in the first few years of its existence. Later careful tests were made by Miller, Rose and others and later by myself, which have shown its stability to be absolute and without reversion, and it has probably been so from the beginning. The fact of its constancy has led to its specific distinction by Miller, as varieties were in his time universally, and up to the present time not rarely, though erroneously, believed to be less stable than true species.

Before leaving the laciniate celandine it is to be noted that in crosses with C. majus it follows the law of Mendel, and for this reason should be considered as a retrograde variety, the more so, as it is also treated as such from a morphological point of view by Stahl and others.

We now come to an enumeration of those cases in which the date of the first appearance [612] of a new horticultural variety has been recorded, and I must apologize for the necessity of again quoting many variations, which have previously been dealt with from another point of view. In such cases I shall limit myself as closely as possible to historical facts. They have been recorded chiefly by Verlot and Carriere, who wrote in Paris shortly after the middle of the past century, and afterwards by Darwin, Korshinsky, and others. It is from their writings and from horticultural literature at large that the following evidence is brought together.

A very well-known instance is that of the dwarf variety of Tagetes signata, which arose in the nursery of Vilmorin in the year 1860. It was observed for the first time in a single individual among a lot of the ordinary Tagetes signata. It was found impossible to isolate it, but the seeds were saved separately. The majority of the offspring returned to the parental type, but two plants were true dwarfs. From these the requisite degree of purity for commercial purposes was reached, the vicinists not being more numerous than 10% of the entire number. The same mutation had been observed a year earlier in the same nursery in a lot of Saponaria calabrica. The seeds of this dwarf repeated the variety in the next generation, but in the third none were observed. Then the variety was [613] thought to be lost, and the culture was given up, as the Mendelian law of the splitting of varietal hybrids was not known. According to our present knowledge we might expect the atavistic descendants of the first dwarf to be hybrids, and to be liable to split in their progeny into one-fourth dwarfs and three-fourths normal specimens. From this it is obvious that the dwarfs would have appeared a second time if the strain had been continued by means of the seeds of the vicinistic progeny.

In order to avoid a return to this phase of the question, another use of the vicinists should at once be pointed out. It is the possibility of increasing the yield of the new variety. If space admits of sowing the seeds of the vicinists, a quarter of the progeny may be expected to come true to the new type, and if they were partly pollinated by the dwarfs, even a larger number would do so. Hence it should be made a rule to sow these seeds also, at least when those of the true representatives of the novelty do not give seed enough for a rapid multiplication.

Other dwarfs are recorded to have sprung from species in the same sudden and unexpected manner, as for instance Ageratum coeruleum of the same nursery, further Clematis Viticella nana and Acer campestre nanum. Prunus Mahaleb nana was discovered in 1828 in one [614] specimen near Orleans by Mme. LeBrun in a large culture of Mahaleb. Lonicera tatarica nana appeared in 1825 at Fontenay-aux Roses. A tall variety of the strawberry is called "Giant of Zuidwijk" and originated at Boskoop in Holland in the nursery of Mr. van de Water, in a lot of seedlings of the ordinary strawberry. It was very large, but produced few runners, and was propagated with much difficulty, for after six years only 15 plants were available. It proved to be a late variety with abundant large fruit, and was sold at a high price. For a long time it was prominent in cultures in Holland only.

Varieties without prickles are known to have originated all of a sudden in sundry cases. Gleditschia sinensis, introduced in 1774 from China, gave two seedlings without spines in the year 1823, in the nursery of Caumzet. It is curious in being one of the rare instances where a simultaneous mutation in two specimens is acknowledged, because as a rule, such records comply with the prevailing, though inexact, belief that horticultural mutations always appear in single individuals.

From Korshinsky's survey of varieties with cut leaves or laciniate forms the following cases may be quoted. In the year 1830 a nurseryman named Jacques had sown a large lot of elms, [615] Ulmus pedunculata. One of the seedlings had cut leaves. He multiplied it by grafting and gave it to the trade under the name of U. pedunculata urticaefolia. It has since been lost.

Laciniate alders seem to have been produced by mutation at sundry times. Mirbel says that the Alnus glutinosa laciniata is found wild in Normandy and in the forests of Montmorency near Paris. A similar variety has been met with in a nursery near Orleans in the year 1855. In connection with this discovery some discussion has arisen concerning the question whether it was probable that the Orleans strain was a new mutation, or derived in some way from the trees cited by Mirbel. Of course, as always in such cases, any doubt, once pronounced, affects the importance of the observation for all time, since it is impossible to gather sufficient historical evidence to fully decide the point. The same variety had appeared under similar circumstances in a nursery at Lyons previously (1812).

Laciniated maples are said to be of relatively frequent occurrence in nurseries, among seedlings of the typical species. Loudon says that once 100 laciniated seedlings were seen to originate from seed of some normal trees. But in this case it is rather probable that the presumed [616] normal parents were in reality hybrids between the type and the laciniated form, and simply split according to Mendel's law. This hypothesis is partly founded on general considerations and partly on experiments made by myself with the cut-leaved celandine, previously alluded to, which I crossed with the type. The hybrids repeated the features of the species and showed no signs of their internal hybrid constitution. But the following year one-fourth of their progeny returned to the cut-leaved form. If the same thing has taken place in the case of Loudon's maples, but without their hybrid origin being known, the result would have been precisely what he observed.

Broussonetia papyriffera dissecta originated about 1830 at Lyons, and a second time in 1866 at Fontenay-aux-Roses. The cut-leaved hazelnuts, birches, beeches and others have mostly been found in the wild state, as I have already pointed out in a previous lecture. A similar variety of the elder, Sambucus nigra laciniata, and its near ally, Sambucus racemosa laciniata, are often to be seen in our gardens. They have been on record since 1886 and come true from seed, but their exact origin seems to have been forgotten. Cut-leaved walnuts have been known since 1812; they come true from seed, but are extremely liable to vicinism, a nuisance which is [617] ascribed by some authors to the fact that often on the same tree the male catkins flower and fall off several weeks before the ripening of the pistils of the other form of flowers.

Weeping varieties afford similar instances. Sophora japonica pendula originated about 1850, and Gleditschia triacanthos pendula some time later in a nursery at Chateau-Thierry (Aisne, France). In the year 1821 the bird's cherry, or Prunus Padus, produced a weeping variety, and in 1847 the same mutation was observed for the allied Prunus Mahaleb. Numerous other instances of the sudden origin of weeping trees, both of conifers and of others, have been brought together in Korshinsky's paper. This striking type of variation includes perhaps the best examples of the whole historical evidence. As a rule they appear in large sowings, only one, or only a few at a time. Many of them have not been observed during their youth, but only after having been planted out in parks and forests, since the weeping characters show only after several years.

The monophyllous bastard-acacia originated in the same way. Its peculiarities will be dealt with on another occasion, but the circumstances of its birth may as well be given here. In 1855 in the nursery of Deniau, at Brain-sur-l'Authion (Maine et Loire), it appeared in a lot of [618] seedlings of the typical species in a single individual. This was transplanted into the Jardin des Plantes at Paris, where it flowered and bore seeds in 1865. It must have been partly pollinated by the surrounding normal representatives of the species, since the seeds yielded only one-fourth of true offspring. This proportion, however, has varied in succeeding years. Briot remarks that the monophyllous bastard acacia is liable to petaloid alterations of its stamens, which deficiency may encroach upon its fertility and accordingly upon the purity of its offspring.

Broom-like varieties often occur among trees, and some are known for their very striking reversions by buds, as we have seen on a previous occasion. They are ordinarily called pyramidal or fastigiate forms, and as far as their history goes, they arise suddenly in large sowings of the normal species. The fastigiate birch was produced in this way by Baumann, the Abies concolor fastigiata by Thibault and Keteleer at Paris, the pyramidal cedar by Paillat, the analogous form of Wellingtonia by Otin. Other instances could easily be added, though of course some of the most highly prized broom-like trees are so old that nothing is known about their origin. This, for instance, is the case with the pyramidal yew-tree, Taxus baccata fastigiata. [619] Others have been found wild, as already mentioned in a former lecture.

An analogous case is afforded by the purpleleaved plums, of which the most known form is Prunus Pissardi. It is said to be a purple variety of Prunus cerasifera, and was introduced at the close of the seventies from Persia, where it is said to have been found in Tabris. A similar variety arose independently and unexpectedly in the nursery of Spath, near Berlin, about 1880, but it seems to differ in some minor points from the Persian prototype.

A white variety of Cyclamen vernum made its appearance in the year 1836 in Holland. A single individual was observed for the first time among a large lot of seedlings, in a nursery near Haarlem. It yielded a satisfactory amount of seed, and the progeny was true to the new type. Such plants propagate slowly, and it was only twenty-seven years later (1863) that the bulbs were offered for sale by the Haarlem firm of Krelage & Son. The price of each bulb was $5.00 in that year, but soon afterwards was reduced to $1.00 each, which was about thrice the ordinary price of the red variety.

The firm of Messrs. Krelage & Son has brought into commerce a wide range of new bulb-varieties, all due to occasional mutations, some by seed and others by buds, or to the accidental [620] transference of new qualities into the already existing varieties by cross-pollination through the agency of insects. Instead of giving long lists of these novelties, I may cite the black tulips, which cost during the first few years of their introduction about $25.00 apiece.

Horticultural mutations are as a rule very rare, especially in genera or species which have not yet been brought to a high degree of variability. In these the wide range of varieties and the large scale in which they are multiplied of course give a greater chance for new varieties. But then the possibilities of crossing are likewise much larger, and apparent changes due to this cause may easily be taken for original mutations.

The rarity of the mutations is often proved by the lapse of time between the introduction of a species and its first sport. Some instances may be given. They afford a proof of the length of the period during which the species remained unaltered, although some of these alterations may be due to a cross with an allied form. Erythrina Crista-galli was introduced about 1770, and produced its first sport in 1884, after more than a century of cultivation. Begonia semperflorens has been cultivated since 1829, and for half a century before it commenced sporting. The same length of time has elapsed [621] between the first culture and the first variation of Crambe maritima. Other cases are on record in which the variability exhibited itself much sooner, perhaps within a few years after the original discovery of the species. But such instances seem, as a rule, to be subject to doubt as to the concurrence of hybridization. So for instance the Iris lortetii, introduced in the year 1895 from the Lebanon, which produced a white variety from its very first seeds. If by chance the introduced plants were natural hybrids between the species and the white variety, this apparent and rather improbable mutation would find a very simple explanation. The length of the period preceding the first signs of variability is largely, of course, due to divergent methods of culture. Such species as Erythrina, which are perennial and only sown on a small scale, should not be expected to show varieties very soon. Annual species, which are cultivated yearly in thousands or even hundreds of thousands of individuals, have a much better chance. Perhaps the observed differences are largely due to this cause.

Monstrosities have, from time to time, given rise to cultivated races. The cockscomb or Celosia is one of the most notorious instances. Cauliflowers, turnips and varieties of cabbages are recorded by De Candolle to have arisen in [622] culture, more than a century ago, as isolated monstrous individuals. They come true from seed, but show deviations from time to time which seem to be intimately linked with their abnormal characters. Apetalous flowers may be considered as another form of monstrosity, and in Salpiglossis sinuata such a variety without a corolla made its appearance in the year 1892 in the nursery of Vilmorin. It appeared suddenly, yielded a good crop of seed and was constant from the outset, without any sign of vicinism or impurity.

In several cases the origin of a variety is obscure, while the subsequent historical evidence is such as to make an original sudden appearance quite probable. Although these instances offer but indirect evidence, and will sooner or later lose their importance, it seems desirable to lay some stress on them here, because most of these cases are very obvious and more striking than purely historical facts. Sterile varieties belong to this heading. Sometimes they bear fruit without kernels, sometimes flowers without sexual organs, or even no flowers at all. Instances have been given in the lecture on retrograde varieties; they are ordinarily assumed to have originated by a leap, because it is not quite clear how a loss of the capacity for the formation of seeds could have been slowly accumulated [623] in preceding generations. An interesting case is afforded by a sterile variety of corn, which originated some time ago in my own pedigree-cultures made for another purpose, and which had begun with an ear of 1886. The first generation from the original seeds showed nothing particular, but the second at once produced quite a number of sterile plants. The sterility was caused by the total lack of branches, including those bearing the pistillate flowers. The terminal spikes themselves were reduced to naked spindles, without branches, without flowers and even almost without bracts.

In some individuals, however, this negative character was seen to give way at the tip, showing a few small naked branches. Of course it was impossible to propagate this curious form, but my observations showed that it sprang into existence from known ancestors by a single step or sudden leap. This leap, however, was not confined to a single specimen; on the contrary it affected 40 plants out of a culture of 340 individuals. The same phenomenon was repeated from the seeds of the normal plants in the following year, but afterwards the monstrosity disappeared.

The Italian poplar affords another instance. It is considered by some authors as a distinct species, Populus italica, and by others as a [624] broom-like variety of the Populus nigra, from which it is distinguished by its erect branches and other characters of minor importance. It is often called the pyramidal or fastigiate poplar. Its origin is absolutely unknown and it occurs only in the cultivated state. In Italy it seems to have been cultivated from the earliest historical times, but it was not introduced into other countries till the eighteenth century. In 1749 it was brought into France, and in 1758 into England, and to day it may be seen along roads throughout central Europe and in a large part of Asia. But the most curious fact is that it is only observed in staminate specimens; pistillate trees have not been found, although often sought for. This circumstance makes it very probable that the origin of the broom-like poplar was a sudden mutation, producing only one individual. This being staminate, it has been propagated exclusively by cuttings. It is to be admitted, however, that no material evidence is at hand to prove that it is not an original wild species, the pistillate form of which has been lost by vegetative multiplication. One form only of many dioecious plants is to be found in cultivation, as, for instance some South American species of Ribes.

Total lack of historical evidence concerning [625] the origin of a variety has sometimes been considered as sufficient proof of a sudden origin. The best known instance is that of the renowned cactus-dahlia with its recurved instead of incurved ray-florets. It was introduced from Mexico into the Netherlands by Van den Berg of Jutphaas, under the following remarkable circumstances. In the autumn of 1872 one of his friends had sent him a small case, containing seeds, bulbs and roots from Mexico. From one of these roots a Dahlia shoot developed. It was cultivated with great care and bloomed next year. It surprised all who saw it by the unexpected peculiarity of its large rich crimson flowers, the rays of which were reversed tubular. The margins of the narrow rays were curved backwards, showing the bright color of the upper surface. It was a very showy novelty, rapidly multiplied by cuttings, and was soon introduced into commerce. It has since been crossed with nearly all other available varieties of the Dahlia, giving a large and rich group of forms, bound together by the curious curling of the petals. It has never been observed to grow in Mexico, either wild or in gardens, and thus the introduced individual has come to be considered as the first of its race.

I have already mentioned that the rapid production of large numbers of new varieties, by [626] means of the crossing of the offspring of a single mutant with previously existing sorts, is a very common feature in horticultural practice. It warns us that only a small part of the novelties introduced yearly are due to real mutations. Further instances of novelties with such a common origin are the purple-leaved dahlias, the gooseberries without prickles, the double petunias, erect gloxinias and many others. Accumulation of characters, acquired in different races of a species, may easily be effected in this way; in fact it is one of the important factors in the breeding of horticultural novelties.

I have alluded more than once in this lecture to the question, whether it is probable that mutations occur in one individual or in more. The common belief among horticulturists is that, as a rule, they appear in a single plant. This belief is so widespread that whenever a novelty is seen for the first time in two or more specimens it is at once suggested that it might have originated and been overlooked in a previous generation. Not caring to confess a lack of close observation, the number of mutants in such cases is usually kept secret. At least this statement has been made to me by some of the horticulturists at Erfurt, whom I visited some years ago in order to learn as much as [627] possible about the methods of production of their novelties. Hence it is simply impossible to decide the question on the basis of the experience of the breeders. Even in the case of the same novelty arising in sundry varieties of the same species, the question as to common origin, by means of crossing, is often hard to decide, as for instance in moss-roses and nectarines. On the other hand, instances are on record where the same novelty has appeared at different times, often at long intervals. Such is the case with the butterfly-cyclamen, a form with wide-spreading petals which originated in Martin's nursery in England. The first time it was seen it was thought to be of no value, and was thrown away, but when appearing for a second time it was multiplied and eventually placed on the market. Other varieties of Cyclamen, as for instance the crested forms, are also known to have originated repeatedly.

In concluding this series of examples of horticultural mutations, I might mention two cases, which have occurred in my own experimental garden. The first refers to a tubular Dahlia. It has ray-florets, the ligules of which have their margins grown together so as to form tubes, with the outer surface corresponding to the pale under-surface of the corolla.

This novelty originated in a single plant in a [628] culture from the seed of the dwarf variety "Jules Chretien." The seeds were taken from introduced plants in my garden, and as the sport has no ornamental value it is uncertain whether this was the first instance or whether it had previously occurred in the nursery at Lyons, from whence the bulbs were secured. Afterwards it proved true from seed, but was very variable, exhibiting rather the features of an ever-sporting variety.

Another novelty was seen the first time in several individuals. It was a pink sport of the European cranesbill, Geranium pratense. It arose quite unexpectedly in the summer of 1902 from a striped variety of the blue species. It was seen in seven specimens out of a lot of about a hundred plants. This strain was introduced into my garden in 1897, when I bought two plants under the name of Geranium pratense album, which however proved to belong to the striped variety. From their seeds I sowed in 1898 a first generation, of which a hundred plants flowered the next year, and from their seeds I sowed in 1900 the lot which produced the sport. Neither the introduced plants nor their offspring had exhibited the least sign of a color-variation, besides the blue and white stripes. Hence it is very probable that my novelty was a true first mutation, the more probably [629] so since a pink variety would without doubt have a certain horticultural value and would have been preserved if it had occurred. But as far as I have been able to ascertain, it is as yet unknown, nor has it been described until today.

Summing up the results of this long, though very incomplete, list of horticultural novelties with a more or less well-known origin, we see that sudden appearances are the rule. Having once sprung into existence the new varieties are ordinarily constant, except as affected by vicinism. Details concerning the process are mostly unavailable or at least are of very doubtful value. And to this it should be added that really progressive mutations have hardly been observed in horticulture. Hence the theoretical value of the facts is far less than might have been expected.

[630] LECTURE XXII

SYSTEMATIC ATAVISM

The steady cooperation of progression and retrogression is one of the important principles of organic evolution. I have dwelt upon this point more than once in previous lectures. I have tried to show that both in the more important lines of the general pedigree of the vegetable kingdom, and in the numerous lateral branches ending in the genera and species within the families, progression and retrogression are nearly always at work together. Your attention has been directed to the monocotyledons as an example, where retrogression is everywhere so active that it can almost be said to be the prevailing movement. Reduction in the vegetative and generative organs, in the anatomical structure and growth of the stems, and in sundry other ways is the method by which the monocotyledons have originated as a group from their supposed ancestors among the lower dicotyledonous families. Retrogression is the leading idea in the larger families of the group, [631] as for instance in the aroids and the grasses. Retrograde evolution is also typical in the highest and most highly differentiated family of the monocotyledons, the orchids, which have but one or two stamens. In the second place I have had occasion more than once to assert that retrogression, though seemingly consisting in the disappearance of some quality, need not, as a rule, be considered as a complete loss. Quite on the contrary, it is very probable that real losses are extremely rare, if not wholly lacking. Ordinarily the loss is only apparent, the capacity becomes inactive only, but is not destroyed. The character has become latent, as it is commonly stated, and therefore may return to activity and to the full display of its peculiarity, whenever occasion offers.

Such a return to activity was formerly called atavism. But as we have seen, when dealing with the phenomena of latency at large, sundry cases of latency are to be distinguished, in order to get a clear insight into these difficult processes.

So it is with atavism, too. If any plant reverts to a known ancestor, we have a positive and simple case. But ancestors with alternate specific marks are as a rule neither historically nor experimentally manifest. They are only reputed to be such, and the presumption rests [632] upon the systematic affinity between the derivative species and its nearest probable allies. Such reversions are now to be examined at some length and may be adequately treated under the head of systematic atavism. To this form of atavism pertain, on the basis of our definition, those phenomena by which species assume one or more characters of allies, from which they are understood to have descended by the loss of the character under discussion. The phenomena themselves consist in the production of anomalies and varieties, and as the genetic relation of the latter is often hardly beyond doubt, the anomalies seem to afford the best instances for the study of systematic atavism. This study has for its chief aim the demonstration of the presence of the latent characters, and to show that they return to activity suddenly and not by a slow and gradual recovery of the former features. It supports the assertion that the visible elementary characters are essentially an external display of qualities carried by the bearers of heredity, and that these bearers are separate entities, which may be mingled together, but are not fused into a chaotic primitive life-substance. Systematic atavism by this means leads us to a closer examination of the internal and concealed causes, which rule the affinities and divergencies of [633] allied species. It brings before us and emphasizes the importance of the conception of the so-called unit-characters.