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Species and Varieties, Their Origin by Mutation
by Hugo DeVries
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Species and Varieties Their Origin by Mutation

Lectures delivered at the University of California

By Hugo DeVries Professor of Botany in the University of Amsterdam

Edited by Daniel Trembly MacDougal Director Department of Botanical Research Carnegie Institution of Washington

Second Edition Corrected and Revised

CHICAGO The Open Court Publishing Company LONDON Kegan Paul, Trench, Trubner and Co., Ltd. 1906

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COPYRIGHT 1904 BY The Open Court Pub. Co. CHICAGO

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THE ORIGIN OF SPECIES

The origin of species is a natural phenomenon. LAMARCK

The origin of species is an object of inquiry. DARWIN

The origin of species is an object of experimental investigation. DeVRIES.

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PREFACE BY THE AUTHOR

THE purpose of these lectures is to point out the means and methods by which the origin of species and varieties may become an object for experimental inquiry, in the interest of agricultural and horticultural practice as well as in that of general biologic science. Comparative studies have contributed all the evidence hitherto adduced for the support of the Darwinian theory of descent and given us some general ideas about the main lines of the pedigree of the vegetable kingdom, but the way in which one species originates from another has not been adequately explained. The current belief assumes that species are slowly changed into new types. In contradiction to this conception the theory of mutation assumes that new species and varieties are produced from existing forms by sudden leaps. The parent-type itself remains unchanged throughout this process, and may repeatedly give birth to new forms. These may arise simultaneously and in groups or separately at more or less widely distant periods.

The principal features of the theory of mutation have been dealt with at length in my book "Die Mutationstheorie" (Vol. I., 1901, Vol. II., 1903. Leipsic, Veit & Co.), in which I have endeavored to present as completely as possible the detailed evidence obtained from trustworthy historical records, and from my own experimental researches, upon which the theory is based.

The University of California invited me to deliver a series of lectures on this subject, at Berkeley, during the [vii] summer of 1904, and these lectures are offered in this form to a public now thoroughly interested in the progress of modern ideas on evolution. Some of my experiments and pedigree-cultures are described here in a manner similar to that used in the "Mutationstheorie," but partly abridged and partly elaborated, in order to give a clear conception of their extent and scope. New experiments and observations have been added, and a wider choice of the material afforded by the more recent current literature has been made in the interest of a clear representation of the leading ideas, leaving the exact and detailed proofs thereof to the students of the larger book.

Scientific demonstration is often long and encumbered with difficult points of minor importance. In these lectures I have tried to devote attention to the more important phases of the subject and have avoided the details of lesser interest to the general reader.

Considerable care has been bestowed upon the indication of the lacunae in our knowledge of the subject and the methods by which they may be filled. Many interesting observations bearing upon the little known parts of the subject may be made with limited facilities, either in the garden or upon the wild flora. Accuracy and perseverance, and a warm love for Nature's children are here the chief requirements in such investigations.

In his admirable treatise on Evolution and Adaptation (New York, Macmillan & Co., 1903), Thomas Hunt Morgan has dealt in a critical manner with many of the speculations upon problems subsidiary to the theory of descent, in so convincing and complete a manner, that I think myself justified in neglecting these questions here. His book gives an accurate survey of them all, and is easily understood by the general reader.

In concluding I have to offer my thanks to Dr. D.T. MacDougal and Miss A.M. Vail of the New York Botanical Garden for their painstaking work in the preparation of the manuscript for the press. Dr. MacDougal, by [viii] his publications, has introduced my results to his American colleagues, and moreover by his cultures of the mutative species of the great evening-primrose has contributed additional proof of the validity of my views, which will go far to obviate the difficulties, which are still in the way of a more universal acceptation of the theory of mutation. My work claims to be in full accord with the principles laid down by Darwin, and to give a thorough and sharp analysis of some of the ideas of variability, inheritance, selection, and mutation, which were necessarily vague at his time. It is only just to state, that Darwin established so broad a basis for scientific research upon these subjects, that after half a century many problems of major interest remain to be taken up. The work now demanding our attention is manifestly that of the experimental observation and control of the origin of species. The principal object of these lectures is to secure a more general appreciation of this kind of work.

HUGO DE VRIES. Amsterdam, October, 1904.

[ix]

PREFACE BY THE EDITOR

PROFESSOR DE VRIES has rendered an additional service to all naturalists by the preparation of the lectures on mutation published in the present volume. A perusal of the lectures will show that the subject matter of "Die Mutationstheorie" has been presented in a somewhat condensed form, and that the time which has elapsed since the original was prepared has given opportunity for the acquisition of additional facts, and a re-examination of some of the more important conclusions with the result that a notable gain has been made in the treatment of some complicated problems.

It is hoped that the appearance of this English version of the theory of mutation will do much to stimulate investigation of the various phases of the subject. This volume, however, is by no means intended to replace, as a work of reference, the larger book with its detailed recital of facts and its comprehensive records, but it may prove a substitute for the use of the general reader.

The revision of the lectures has been a task attended with no little pleasure, especially since it has given the editor the opportunity for an advance consideration of some of the more recent results, thus materially facilitating investigations which have been in progress at the New York Botanical Garden for some time. So far as the ground has been covered the researches in question corroborate the conclusions of de Vries in all important particulars. The preparation of the manuscript for the printer has consisted chiefly in the adaptation of oral [xii] discussions and demonstrations to a form suitable for permanent record, together with certain other alterations which have been duly submitted to the author. The original phraseology has been preserved as far as possible. The editor wishes to acknowledge material assistance in this work from Miss A.M. Vail, Librarian of the New York Botanical Garden.

D.T. MacDougal. New York Botanical Garden, October, 1904.

PREFACE TO THE SECOND EDITION.

THE constantly increasing interest in all phases of evolution has made necessary the preparation of a second edition of this book within a few months after the first appeared. The opportunity has been used to eliminate typographical errors, and to make alterations in the form of a few sentences for the sake of clearness and smoothness. The subject matter remains practically unchanged. An explanatory note has been added on page 575 in order to avoid confusion as to the identity of some of the plants which figure prominently in the experimental investigations in Amsterdam and New York.

The portrait which forms the frontispiece is a reproduction of a photograph taken by Professor F.E. Lloyd and Dr. W.A. Cannon during the visit of Professor de Vries at the Desert Botanical Laboratory of the Carnegie Institution, at Tucson, Arizona, in June, 1904.

D. T. MACDOUGAL. December 15, 1905.



CONTENTS

A. INTRODUCTION.

LECTURE PAGE

I. Descent: theories of evolution and methods of investigation. 1 The theory of descent and of natural selection. Evolution and adaptation. Elementary species and varieties. Methods of scientific pedigree-culture.

B. ELEMENTARY SPECIES.

II. Elementary species in nature. 32 Viola tricolor, Draba verna, Primula acaulis, and other examples. Euphorbia pecacuanha. Prunus maritima. Taraxacum and Hieracium.

III. Elementary species of cultivated plants. 63 Beets, apples, pears, clover, flax and coconut.

IV. Selection of elementary species. 92 Cereals. Le Couteur. Running out of varieties. Rimpau and Risler, Avena fatua. Meadows. Old Egyptian cereals. Selection by the Romans. Shirreff. Hays.

C. RETROGRADE VARIETIES.

V. Characters of retrograde varieties. 121 Seed varieties of pure, not hybrid origin. Differences from elementary species. Latent characters. Ray-florets of composites. [xiii] Progressive red varieties. Apparent losses. Xanthium canadense. Correlative variability. Laciniate leaves and petals. Compound characters.

VI. Stability and real atavism. 154 Constancy of retrograde varieties. Atavism in Ribes sanguineum Albidum, in conifers, in Iris pallida. Seedlings of Acacia. Reversion by buds.

VII. Ordinary or false atavism. 185 Vicinism or variation under the influence of pollination by neighboring individuals. Vicinism in nurseries. Purifying new and old varieties. A case of running out of corn in Germany.

VIII. Latent characters. 216 Leaves of seedlings, adventitious buds, systematic latency and retrogressive evolution. Degressive evolution. Latency of specific and varietal characters in wheat-ear carnation, in the green dahlias, in white campanulas and others. Systematic latency of flower colors.

IX. Crossing of species and varieties. 247 Balanced and unbalanced, or species and variety crosses. Constant hybrids of Oenothera muricata and O. biennis. Aegilops, Medicago, brambles and other instances.

X. Mendel's law of balanced crosses. 276 Pairs of antagonistic characters, one active and one latent. Papaver somniferum. [xiv] Mephisto Danebrog. Mendel's laws. Unit- characters.

D. EVERSPORTING VARIETIES.

XI. Striped flowers. 309 Antirrhinum majus luteum rubro-striatum with pedigree. Striped flowers, fruits and radishes. Double stocks.

XII. "Five leaved" clover. 340 Origin of this variety. Periodicity of the anomaly. Pedigree- cultures. Ascidia.

XIII. Polycephalic poppies. 369 Permanency and high variability. Sensitive period of the anomaly. Dependency on external conditions.

XIV. Monstrosities. 400 Inheritance of monstrosities. Half races and middle races. Hereditary value of atavists. Twisted stems and fasciations. Middle races of tricotyls and syncotyls. Selection by the hereditary percentage among the offspring.

XV. Double adaptations. 430 Analogy between double adaptations and anomalous middle races. Polygonum amphibium. Alpine plants. Othonna crassifolia. Leaves in sunshine and shadow. Giants and dwarfs. Figs and ivy. Leaves of seedlings.

E. MUTATIONS.

XVI. Origin of the peloric toad-flax. 459 Sudden and frequent origin in the wild state. Origin in the experiment-garden. Law of repeated mutations. Probable origin of other pelories.

[xv] XVII. The production of double flowers. 488 Sudden appearance of double flowers in horticulture. Historical evidence. Experimental origin of Chrysanthemum segetum plenum. Dependency upon nourishment. Petalody of stamens.

XVIII New species of Oenothera. 516 Mutations of Oenothera lamarckiana in the wild state near Hilversum. New varieties of O. laevifolia, O. brevistylis, and O. nanella. New elementary species, O. gigas, O. rubrinervis, albida, and oblonga. O. lata, a pistillate form. Inconstancy of O. scintillans.

XIX. Experimental pedigree-cultures. 547 Pedigree of the mutative products of Oenothera lamarckiana in the Botanical Garden at Amsterdam. Laws of mutability. Sudden and repeated leaps from an unchanging main strain. Constancy of the new forms. Mutations in all directions.

XX. Origin of wild species and varieties. 576 Problems to solve. Capsella heegeri. Oenothera biennis cruciata. Epilobium hirsutum cruciatum. Hibiscus Moscheutos. Purple beech. Monophyllous strawberries. Chances of success with new mutations.

XXI. Mutations in horticulture. 604 Chelidonium majus lacinatum. Dwarf and spineless varieties. Laciniate leaves. Monophyllous and broom-like varieties. [xvi] Purple leaves. Celosia. Italian poplar. Cactus dahlia. Mutative origin of Dahlia fistulosa, and Geranium praetense in the experiment-garden.

XXII. Systematic atavism. 630 Reappearance of ancestral characters. Primula acaulis umbellata. Bracts of crucifers. Zea Mays cryptosperma. Equisetum, Dipsacus sylvestris torsus. Tomatoes.

XXIII. Taxonomic anomalies. 658 Specific characters occurring in other cases as casual anomalies. Papaver bracteatum monopetalum. Desmodium gyrans and monophyllous varieties. Peltate leaves and ascidia. Flowers on leaves. Leaves. Hordeum trifurcatum.

XXIV. Hypothesis of periodical mutations. 686 Discovering mutable strains. Periods of mutability and constancy. Periods of mutations. Genealogical trees. Limited life-time of the organic kingdom.

F. FLUCTUATIONS.

XXV. General laws of fluctuations. 715 Fluctuating variability. Quetelet's law. Individual and partial fluctuations. Linear variability. Influence of nutrition. Periodicity curves.

XXVI. Asexual multiplication of extremes. 742 Selection between species and intra-specific selection. Excluding individual [xvii] embryonic variability. Sugar-canes. Flowering cannas. Double lilacs. Other instances. Burbank's method of selection.

XXVII. Inconstancy of improved races 770 Larger variability in the case of propagation by seed, progression and regression after a single selection, and after repeated selections. Selection experiments with corn. Advantages and effect of repeated selection.

XXVIII. Artificial and natural selection. 798 Conclusions. Specific and intra-specific selection. Natural selection in the field. Acclimatization. Improvement-selection of sugar-beets by various methods. Rye. Hereditary percentage and centgener power as marks by which intraspecific selection may be guided.

Index 827

[1] A. INTRODUCTION

LECTURE I

DESCENT: THEORIES OF EVOLUTION AND METHODS OF INVESTIGATION

Newton convinced his contemporaries that natural laws rule the whole universe. Lyell showed, by his principle of slow and gradual evolution, that natural laws have reigned since the beginning of time. To Darwin we owe the almost universal acceptance of the theory of descent.

This doctrine is one of the most noted landmarks in the advance of science. It teaches the validity of natural laws of life in its broadest sense, and crowns the philosophy founded by Newton and Lyell.

Lamarck proposed the hypothesis of a common origin of all living beings and this ingenious and thoroughly philosophical conception was warmly welcomed by his partisans, but was not widely accepted owing to lack of supporting evidence. To Darwin was reserved the task of [2] bringing the theory of common descent to its present high rank in scientific and social philosophy.

Two main features in his work have contributed to this early and unexpected victory. One of them is the almost unlimited amount of comparative evidence, the other is his demonstration of the possibility of a physiological explanation of the process of descent itself.

The universal belief in the independent creation of living organisms was revised by Linnaeus and was put upon a new foundation. Before him the genera were supposed to be created, the species and minor forms having arisen from them through the agency of external conditions. In his first book Linnaeus adhered to this belief, but later changed his mind and maintained the principle of the separate creation of species. The weight of his authority soon brought this conception to universal acceptance, and up to the present time the prevailing conception of a species has been chiefly based on the definition given by Linnaeus. His species comprised subspecies and varieties, which were in their turn, supposed to have evolved from species by the common method.

Darwin tried to show that the links which bind species to genera are of the same nature as those which determine the relationship of [3] subspecies and varieties. If an origin by natural laws is conceded for the latter, it must on this ground be granted for the first also. In this discussion he simply returned to the pre-Linnean attitude. But his material was such as to allow him to go one step further, and this step was an important and decisive one. He showed that the relation between the various genera of a family does not exhibit any features of a nature other than that between the species of a genus. What has been conceded for the one must needs be accepted for the other. The same holds good for the large groups.

The conviction of the common origin of closely allied forms necessarily leads to the conception of a similar descent even in remote relationships.

The origin of subspecies and varieties as found in nature was not proved, but only generally recognized as evident. A broader knowledge has brought about the same state of opinion for greater groups of relationships. Systematic affinities find their one possible explanation by the aid of this principle; without it, all similarity is only apparent and accidental. Geographic and paleontologic facts, brought together by Darwin and others on a previously unequalled scale, point clearly in the same direction. The vast amount of evidence of all [4] comparative sciences compels us to accept the idea. To deny it, is to give up all opportunity of conceiving Nature in her true form.

The general features of the theory of descent are now accepted as the basis of all biological science. Half a century of discussion and investigation has cleared up the minor points and brought out an abundance of facts; but they have not changed the principle. Descent with modification is now universally accepted as the chief law of nature in the organic world. In honor of him, who with unsurpassed genius, and by unlimited labor has made it the basis of modern thought, this law is called the "Darwinian theory of descent."

Darwin's second contribution to this attainment was his proof of the possibility of a physiological explanation of the process of descent itself. Of this possibility he fully convinced his contemporaries, but in indicating the particular means by which the change of species has been brought about, he has not succeeded in securing universal acceptation. Quite on the contrary, objections have been raised from the very outset, and with such force as to compel Darwin himself to change his views in his later writings. This however, was of no avail, and objections and criticisms have since steadily accumulated. Physiologic facts concerning the origin of [5] species in nature were unknown in the time of Darwin. It was a happy idea to choose the experience of the breeders in the production of new varieties, as a basis on which to build an explanation of the processes of nature. In my opinion Darwin was quite right, and he has succeeded in giving the desired proof. But the basis was a frail one, and would not stand too close an examination. Of this Darwin was always well aware. He has been prudent to the utmost, leaving many points undecided, and among them especially the range of validity of his several arguments. Unfortunately this prudence has not been adopted by his followers. Without sufficient warrant they have laid stress on one phase of the problem, quite overlooking the others. Wallace has even gone so far in his zeal and ardent veneration for Darwin, as to describe as Darwinism some things, which in my opinion, had never been a part of Darwin's conceptions.

The experience of the breeders was quite inadequate to the use which Darwin made of it. It was neither scientific, nor critically accurate. Laws of variation were barely conjectured; the different types of variability were only imperfectly distinguished. The breeders' conception was fairly sufficient for practical purposes, but science needed a clear understanding of the [6] factors in the general process of variation. Repeatedly Darwin tried to formulate these causes, but the evidence available did not meet his requirements.

Quetelet's law of variation had not yet been published. Mendel's claim of hereditary units for the explanation of certain laws of hybrids discovered by him, was not yet made. The clear distinction between spontaneous and sudden changes, as compared with the ever-present fluctuating variations, is only of late coming into recognition by agriculturists. Innumerable minor points which go to elucidate the breeders' experience, and with which we are now quite familiar, were unknown in Darwin's time. No wonder that he made mistakes, and laid stress on modes of descent, which have since been proved to be of minor importance or even of doubtful validity.

Notwithstanding all these apparently unsurmountable difficulties, Darwin discovered the great principle which rules the evolution of organisms. It is the principle of natural selection. It is the sifting out of all organisms of minor worth through the struggle for life. It is only a sieve, and not a force of nature, not a direct cause of improvement, as many of Darwin's adversaries, and unfortunately many of his followers also, have so often asserted.

It is [7] only a sieve, which decides what is to live, and what is to die. But evolutionary lines are of great length, and the evolution of a flower, or of an insectivorous plant is a way with many sidepaths. It is the sieve that keeps evolution on the main line, killing all, or nearly all that try to go in other directions. By this means natural selection is the one directing cause of the broad lines of evolution.

Of course, with the single steps of evolution it has nothing to do. Only after the step has been taken, the sieve acts, eliminating the unfit. The problem, as to the manner in which the individual steps are brought about, is quite another side of the question.

On this point Darwin has recognized two possibilities. One means of change lies in the sudden and spontaneous production of new forms from the old stock. The other method is the gradual accumulation of those always present and ever fluctuating variations which are indicated by the common assertion that no two individuals of a given race are exactly alike. The first changes are what we now call "mutations," the second are designated as "individual variations," or as this term is often used in another sense, as "fluctuations." Darwin recognized both lines of evolution; Wallace disregarded the sudden changes and proposed fluctuations [8] as the exclusive factor. Of late, however, this point of view has been abandoned by many investigators, especially in America.

The actual occurrence of mutations is recognized, and the battle rages about the question, as to whether they are be regarded as the principal means of evolution, or whether slow and gradual changes have not also played a large and important part.

The defenders of the theory of evolution by slow accumulation of slight fluctuations are divided into two camps. One group is called the Neo-Lamarckians; they assume a direct modifying agency of the environment, producing a corresponding and useful change in the organization. The other group call themselves Darwinians or selectionists, but to my mind with no other right beyond the arbitrary restriction of the Darwinian principles by Wallace. They assume fluctuating variations in all directions and leave the choice between them to the sieve of natural selection.

Of course we are far from a decision between these views, on the sole ground of the facts as known at present. Mutations under observation are as yet very rare; enough to indicate the possible and most probable ways, but no more. On the other hand the accumulation of fluctuations does not transgress relatively narrow [9] limits as far as the present methods of selection go. But the question remains to be solved, whether our methods are truly the right ones, and whether by the use of new principles, new results might not cause the balance of opinion to favor the opposite side.

Of late, a thorough and detailed discussion of the opposing views has been given by Morgan in his valuable book on evolution and adaptation. He has subjected all the proposed theories to a severe criticism both on the ground of facts and on that of their innate possibility and logical value. He decides in favor of the mutation theory. His arguments are incisive and complete and wholly adapted to the comprehension of all intelligent readers, so that his book relieves me entirely of the necessity of discussing these general questions, as it could not be done in a better or in a clearer way.

I intend to give a review of the facts obtained from plants which go to prove the assertion, that species and varieties have originated by mutation, and are, at present, not known to originate in any other way. This review consists of two parts. One is a critical survey of the facts of agricultural and horticultural breeding, as they have accumulated since the time of Darwin. This body of evidence is to be combined with some corresponding experiments [10] concerning the real nature of species in the wild state. The other part rests on my own observations and experiments, made in the botanical garden of the University of Amsterdam.

For many years past I have tried to elucidate the hereditary conditions of species and varieties, and the occasional occurrence of mutations, that suddenly produce new forms.

The present discussion has a double purpose. On one side it will give the justification of the theory of mutations, as derived from the facts now at hand. On the other hand it will point out the deficiencies of available evidence, and indicate the ways by which the lacunae may gradually be filled. Experimental work on heredity does not require vast installments or costly laboratory equipment. It demands chiefly assiduity and exactitude. Any one who has these two qualities, and who has a small garden at his disposal is requested to take part in this line of investigation.

In order to observe directly the birth of new forms it is necessary, in the first place, to be fully clear concerning the question as to what forms are to be expected to arise from others, and before proceeding to a demonstration of the origin of species, it is pertinent to raise the question as to what constitutes a species.

Species is a word, which always has had a [11] double meaning. One is the systematic species, which is the unit of our system. But these units are by no means indivisible. Long ago Linnaeus knew them to be compound in a great number of instances, and increasing knowledge has shown that the same rule prevails in other instances. Today the vast majority of the old systematic species are known to consist of minor units. These minor entities are called varieties in systematic works. However, there are many objections to this usage. First, the term variety is applied in horticulture and agriculture to things so widely divergent as to convey no clear idea at all. Secondly, the subdivisions of species are by no means all of the same nature, and the systematic varieties include units the real value of which is widely different in different cases. Some of these varieties are in reality as good as species, and have been "elevated," as it is called by some writers, to this rank. This conception of the elementary species would be quite justifiable, and would at once get rid of all difficulties, were it not for one practical obstacle. The number of the species in all genera would be doubled and tripled, and as these numbers are already cumbersome in many cases, the distinction of the native species of any given country would lose most of its charm and interest.

[12] In order to meet this difficulty we must recognize two sorts of species. The systematic species are the practical units of the systematists and florists, and all friends of wild nature should do their utmost to preserve them as Linnaeus has proposed them. These units however, are not really existing entities; they have as little claim to be regarded as such as genera and families. The real units are the elementary species; their limits often apparently overlap and can only in rare cases be determined on the sole ground of field observations. Pedigree-culture is the method required and any form which remains constant and distinct from its allies in the garden is to be considered as an elementary species.

In the following lectures we shall consider this point at length, to show the compound nature of systematic species in wild and in cultivated plants. In both cases, the principle is becoming of great importance, and many papers published recently indicate its almost universal acceptation.

Among the systematic subdivisions of species, not all have the same claim to the title of elementary species. In the first place the cases in which the differences may occur between parts of the same individual are to be excluded. Dividing an alpine plant into two halves and [13] planting one in a garden, varietal differences at once arise and are often designated in systematic works under different varietal names. Secondly all individual differences which are of a fluctuating nature are to be combined into a group. But with these we shall deal later.

Apart from these minor points the subdivisions of the systematic species exhibit two widely different features. I will now try to make this clear in a few words, but will return in another lecture to a fuller discussion of this most interesting contrast.

Linnaeus himself knew that in some cases all subdivisions of a species are of equal rank, together constituting the group called species. No one of them outranks the others; it is not a species with varieties, but a group, consisting only of varieties. A closer inquiry into the cases treated in this manner by the great master of systematic science, shows that here his varieties were exactly what we now call elementary species.

In other cases the varieties are of a derivative nature. The species constitutes a type that is pure in a race which ordinarily is still growing somewhere, though in some cases it may have died out. From this type the varieties are derived, and the way of this derivation is usually quite manifest to the botanist. It is ordinarily [14] by the disappearance of some superficial character that a variety is distinguished from its species, as by the lack of color in the flowers, of hairs on stems and foliage, of the spines and thorns, &c. Such varieties are, strictly speaking, not to be treated in the same way as elementary species, though they often are. We shall designate them by the term of "retrograde varieties," which clearly indicates the nature of their relationship to the species from which they are assumed to have sprung. In order to lay more stress on the contrast between elementary species and retrograde varieties, it should be stated at once, that the first are considered to have originated from their parent-form in a progressive way. They have succeeded in attaining something quite new for themselves, while retrograde varieties have only thrown off some peculiarity, previously acquired by their ancestors.

The whole vegetable kingdom exhibits a constant struggle between progression and retrogression. Of course, the great lines of the general pedigree are due to progression, many single steps in this direction leading together to the great superiority of the flowering plants over their cryptogamous ancestors. But progression is nearly always accompanied by retrogression in the principal lines of evolution, [15] as well as in the collateral branches of the genealogical tree. Sometimes it prevails, and the monocotyledons are obviously a reduced branch of the primitive dicotyledons. In orchids and aroids, in grasses and sedges, reduction plays a most important part, leaving its traces on the flowers as well as on the embryo of the seed. Many instances could be given to prove that progression and retrogression are the two main principles of evolution at large. Hence the conclusion, that our analysis must dissect the complicated phenomena of evolution so far as to show the separate functions of these two contrasting principles. Hundreds of steps were needed to evolve the family of the orchids, but the experimenter must take the single steps for the object of his inquiry. He finds that some are progressive and others retrogressive and so his investigation falls under two heads, the origin of progressive characters, and the subsequent loss of the same. Progressive steps are the marks of elementary species, while retrograde varieties are distinguished by apparent losses. They have equal claim to our interest and our study.

As already stated I propose to deal first with the elementary species and afterwards with the retrograde varieties. I shall try to depict them to you in the first place as they are seen in [16] nature and in culture, leaving the question of their origin to a subsequent experimental treatment.

The question of the experimental origin of new species and varieties has to be taken up from two widely separated starting points. This may be inferred from what we have already seen concerning the two opposing theories, derived and isolated from Darwin's original broad conception. One of them considers mutations as the origin of new forms, while the other assumes fluctuations to be the source of all evolution.

As mentioned above, my own experience has led me to accept the first view. Therefore I shall have to show that mutations do yield new and constant forms, while fluctuations are not adequate to do so. Retrograde varieties and elementary species may both be seen to be produced by sudden mutations. Varieties have often been observed to appear at once and quite unexpectedly in horticulture and agriculture, and a survey of these historical facts will be the subject of one of my lectures. In some instances I have succeeded in repeating these observations in my garden under the strict conditions of a scientific experiment, and these instances teach us the real nature of the process of mutation in all its visible features. New elementary [17] species are far more rare, but I have discovered in the great evening-primrose, or Oenothera lamarckiana a strain which is producing them yearly in the wild state as well as in my garden. These observations and pedigree-experiments will be dealt with at due length in subsequent lectures.

Having proved the existence and importance of mutations, it remains to inquire how far the improvements may go which are due only to fluctuating variability. As the term indicates, this variability is fluctuating to and fro, oscillating around an average type. It never fails nor does it, under ordinary circumstances, depart far from the fixed average.

But the deviation may be enlarged by a choice of extremes. In sowing their seed, the average of the strain is seen to be changed, and in repeating the experiment the change may be considerable. It is not clear, whether theoretically by such an accumulation, deviations might be reached which could not be attained at once in a single sowing. This question is hardly susceptible of an experimental answer, as it would require such an enormous amount of seed from a few mother plants as can scarcely ever be produced.

The whole character of the fluctuations shows them to be of an opposite nature, contrasting [18] manifestly with specific and varietal characters. By this method they may be proved to be inadequate ever to make a single step along the great lines of evolution, in regard to progressive as well as to retrograde development.

First of all fluctuations are linear, amplifying or lessening the existing qualities, but not really changing their nature. They are not observed to produce anything quite new, and evolution of course, is not restricted to the increase of the already existing peculiarities, but depends chiefly on the continuous addition of new characters to the stock. Fluctuations always oscillate around an average, and if removed from this for some time, they show a tendency to return to it. This tendency, called retrogression, has never been observed to fail, as it should, in order to free the new strain from the links with the average, while new species and new varieties are seen to be quite free from their ancestors and not linked to them by intermediates.

The last few lectures will be devoted to questions concerning the great problem of the analogy between natural and artificial selection. As already stated, Darwin made this analogy the foundation stone of his theory of descent, and he met with the severest objections and criticisms precisely on this point. But I hope to [19] show that he was quite right, and that the cause of the divergence of opinions is due simply to the very incomplete state of knowledge concerning both processes. If both are critically analyzed they may be seen to comprise the same factors, and further discussion may be limited to the appreciation of the part which each of them has played in nature and among cultivated plants.

Both natural and artificial selection are partly specific, and partly intra-specific or individual. Nature of course, and intelligent men first chose the best elementary species from among the swarms. In cultivation this is the process of variety-testing. In nature it is the survival of the fittest species, or, as Morgan designates it, the survival of species in the struggle for existence. The species are not changed by this struggle, they are only weighed against each other, the weak being thrown aside.

Within the chosen elementary species there is also a struggle. It is obvious, that the fluctuating variability adapts some to the given circumstances, while it lessens the chances of others. A choice results, and this choice is what is often exclusively called selection, either natural or artificial. In cultivation it produces the improved and the local races; in nature little is known about improvement in this way, but [19] local adaptations with slight changes of the average character in separate localities, seem to be of quite normal occurrence.

A new method of individual selection has been used in recent years in America, especially by W.M. Hays. It consists in judging the hereditary worth of a plant by the average condition of its offspring, instead of by its own visible characters. If this determination of the "centgener power," as Hays calls it, should prove to be the true principle of selection, then indeed the analogy between natural and artificial selection would lose a large part of its importance. We will reserve this question for the last lecture, as it pertains more to the future, than to our present stock of knowledge.

Something should be said here concerning hybrids and hybridism. This problem has of late reached such large proportions that it cannot be dealt with adequately in a short survey of the phenomena of heredity in general. It requires a separate treatment. For this reason I shall limit myself to a single phase of the problem, which seems to be indispensable for a true and at the same time easy distinction between elementary species and retrograde varieties. According to accepted terminology, some crosses are to be considered as unsymmetrical, while others are symmetrical. The first are one-sided, [21] some peculiarity being found in one of the parents and lacking in the other. The second are balanced, as all the characters are present in both parents, but are found in a different condition. Active in one of them, they are concealed or inactive in the other. Hence pairs of contrasting units result, while in unbalanced crosses no pairing of the particular character under consideration is possible. This leads to the principal difference between species and varieties, and to an experimental method of deciding between them in difficult and doubtful cases.

Having thus indicated the general outlines of the subjects I shall deal with, something now may be said as to methods of investigation.

There are two points in which scientific investigation differs from ordinary pedigree-culture in practice. First the isolation of the individuals and the study of individual inheritance, instead of averages. Next comes the task of keeping records. Every individual must be entered, its ancestry must be known as completely as possible, and all its relations must be noted in such a form, that the most complete reference is always possible. Mutations may come unexpectedly, and when once arisen, their parents and grand-parents should be known. Records must be available which will allow of a most complete knowledge of the whole ancestral [22] line. This, and approximately this only, is the essential difference between experimental and accidental observation.

Mutations are occurring from time to time in the wild state as well as in horticulture and agriculture. A selection of the most interesting instances will be given later. But in all such cases the experimental proof is wanting. The observations as a rule, only began when the mutation had made its appearance. A more or less vague remembrance about the previous state of the plants in question might be available, though even this is generally absent. But on doubtful points, concerning possible crosses or possible introduction of foreign strains, mere recollection is insufficient. The fact of the mutation may be very probable, but the full proof is, of course, wanting. Such is the case with the mutative origin of Xanthium commune Wootoni from New Mexico and of Oenothera biennis cruciata from Holland. The same doubt exists as to the origin of the Capsella heegeri of Solms-Laubach, and of the oldest recorded mutation, that of Chelidonium laciniatum in Heidelberg about 1600.

First, we have doubts about the fact itself. These, however, gradually lose their importance in the increasing accumulation of evidence. Secondly, the impossibility of a closer [23] inquiry into the real nature of the change. For experimental purposes a single mutation does not suffice; it must be studied repeatedly, and be produced more or less arbitrarily, according to the nature of the problems to be solved. And in order to do this, it is evidently not enough to have in hand the mutated individual, but it is indispensable to have also the mutable parents, or the mutable strain from which it sprang.

All conditions previous to the mutation are to be considered as of far higher importance than all those subsequent to it.

Now mutations come unexpectedly, and if the ancestry of an accidental mutation is to be known, it is of course necessary to keep accounts of all the strains cultivated. It is evident that the required knowledge concerning the ancestry of a supposed mutation, must necessarily nearly all be acquired from the plants in the experimental garden.

Obviously this rule is as simple in theory, as it is difficult to carry out in practice. First of all comes the book-keeping. The parents, grandparents and previous ancestors must be known individually. Accounts of them must be kept under two headings. A full description of their individual character and peculiarities must always be available on the one hand, and on the other, all facts concerning their hereditary [24] qualities. These are to be deduced from the composition of the progeny, and in order to obtain complete evidence on this point, two successive generations are often required. The investigation must ascertain the average condition of this offspring and the occurrence of any deviating specimens, and for both purposes it is necessary to cultivate them in relatively large numbers. It is obvious that, properly speaking, the whole family of a mutated individual, including all its nearer and more remote relatives, should be known and recorded.

Hence pedigree-book-keeping must become the general rule. Subordinate to this are two further points, which should likewise be stated here. One pertains to the pure or hybrid nature of the original strain, and the other to the life-conditions and all other external influences. It is manifest that a complete understanding of a mutation depends upon full information upon these points.

All experiments must have a beginning. The starting-point may be a single individual, or a small group of plants, or a lot of seeds. In many cases the whole previous history is obscure, but sometimes a little historical evidence is at hand. Often it is evident that the initial material belongs to a pure species, but with respect to the question of elementary species it is [25] not rarely open to doubt. Large numbers of hybrid plants and hybrid races are in existence, concerning the origin of which it is impossible to decide. It is impossible in many instances to ascertain whether they are of hybrid or of pure origin. Often there is only one way of determining the matter; it is to guess at the probable parents in case of a cross and to repeat the cross. This is a point which always requires great care in the interpretation of unusual facts.

Three cases are to be distinguished as to heredity. Many plants are so constituted as to be fertilized with their own pollen. In this case the visits of insects have simply to be excluded, which may be done by covering plants with iron gauze or with bags of prepared paper. Sometimes they fertilize themselves without any aid, as for instance, the common evening-primrose; in other cases the pollen has to be placed on the stigma artificially, as with Lamarck's evening-primrose and its derivatives. Other plants need cross-fertilization in order to produce a normal yield of seeds. Here two individuals have always to be combined, and the pedigree becomes a more complicated one. Such is the case with the toad-flax, which is nearly sterile with its own pollen. But even in these cases the visits of insects bringing pollen [26] from other plants, must be carefully excluded. A special lecture will be devoted to this very interesting source of impurity and of uncertainty in ordinary cultures.

Of course, crosses may lie in the proposed line of work, and this is the third point to be alluded to. They must be surrounded with the same careful isolation and protection against bees, as any other fertilizations. And not only the seed-parent, but also the pollen must be kept pure from all possible foreign admixtures.

A pure and accurately recorded ancestry is thus to be considered as the most important condition of success in experimental plant breeding. Next to this comes the gathering of the seeds of each individual separately. Fifty or sixty, and often more, bags of seeds are by no means uncommon for a single experiment, and in ordinary years the harvest of my garden is preserved in over a thousand separate lots.

Complying with these conditions, the origin of species may be seen as easily as any other phenomenon. It is only necessary to have a plant in a mutable condition. Not all species are in such a state at present, and therefore I have begun by ascertaining which were stable and which were not. These attempts, of course, had to be made in the experimental garden, and large quantities of seed had to be procured and [27] sown. Cultivated plants of course, had only a small chance to exhibit new qualities, as they have been so strictly controlled during so many years. Moreover their purity of origin is in many cases doubtful. Among wild plants only those could be expected to reward the investigator which were of easy cultivation. For this reason I have limited myself to the trial of wild plants of Holland, and have had the good fortune to find among them at least one species in a state of mutability. It was not really a native plant, but one that had been introduced from America and belongs to an American genus. I refer to the great evening-primrose or the evening-primrose of Lamarck. A strain of this beautiful species is growing in an abandoned field in the vicinity of Hilversum, at a short distance from Amsterdam. Here it has escaped from a park and multiplied. In doing so it has produced and is still producing quite a number of new types, some of which may be considered as retrograde varieties, while others evidently are of the nature of progressive elementary species.

This interesting plant has afforded me the means of observing directly how new species originate, and of studying the laws of these changes. My researches have followed a double line of inquiry. On one side, I have limited [28] myself to direct field observations, and to tests of seed, collected from the wild plants in their native locality. Obviously the mutations are decided within the seed, and the culture of young plants from them had no other aim than that of ascertaining what had occurred in the field. And then the many chances of destruction that threaten young plants in a wild state, could be avoided in the garden, where environmental factors can be controlled.

My second line of inquiry was an experimental repetition of the phenomena which were only partly discerned at the native locality. It was not my aim to intrude into the process, nor to try to bring out new features. My only object was to submit to the precepts just given concerning pure treatment, individual seed gathering, exclusion of crosses and accurate recording of all the facts. The result has been a pedigree which now permits of stating the relation between all the descendants of my original introduced plant. This pedigree at once exhibits the laws followed by the mutating species. The main fact is, that it does not change itself gradually, but remains unaffected during all succeeding generations. It only throws off new forms, which are sharply contrasted with the parent, and which are from the very beginning as perfect and as constant, as narrowly [29] defined and as pure of type as might be expected of any species.

These new species are not produced once or in single individuals, but yearly and in large numbers. The whole phenomenon conveys the idea of a close group of mutations, all belonging to one single condition of mutability. Of course this mutable state must have had a beginning, as it must sometime come to an end. It is to be considered as a period within the life-time of the species and probably it is only a small part of it.

The detailed description of this experiment, however, I must delay to a subsequent lecture, but I may be allowed to state, that the discovery of this period of mutability is of a definite theoretical importance. One of the greatest objections to the Darwinian theory of descent arose from the length of time it would require, if all evolution was to be explained on the theory of slow and nearly invisible changes. This difficulty is at once met and fully surmounted by the hypothesis of periodical but sudden and quite noticeable steps. This assumption requires only a limited number of mutative periods, which might well occur within the time allowed by physicists and geologists for the existence of animal and vegetable life on the earth.

[30] Summing up the main points of these introductory remarks, I propose to deal with the subjects mentioned above at some length, devoting to each of them, if possible at least an entire lecture. The decisive facts and discussions upon which the conclusions are based will be given in every case. Likewise I hope to point out the weak places and the lacunae in our present knowledge, and to show the way in which each of you may try to contribute his part towards the advancement of science in this subject. Lastly I shall try to prove that sudden mutation is the normal way in which nature produces new species and new varieties. These mutations are more readily accessible to observation and experiment than the slow and gradual changes surmised by Wallace and his followers, which are entirely beyond our present and future experience.

The theory of mutations is a starting-point for direct investigation, while the general belief in slow changes has held back science from such investigations during half a century.

Coming now to the subdivisions and headings under which my material is to be presented, I propose describing first the real nature of the elementary species and retrograde varieties, both in normal form and in hybridizations. A discussion of other types of varieties, including [31] monstrosities will complete the general plan. The second subdivision will deal with the origin of species and varieties as taught by experiment and observation, treating separately the sudden variations which to my mind do produce new forms, and subsequently the fluctuations which I hold to be not adequate to this purpose.

[32] B. ELEMENTARY SPECIES

LECTURE II

ELEMENTARY SPECIES IN NATURE

What are species? Species are considered as the true units of nature by the vast majority of biologists. They have gained this high rank in our estimation principally through the influence of Linnaeus. They have supplanted the genera which were the accepted units before Linnaeus. They are now to be replaced in their turn, by smaller types, for reasons which do not rest upon comparative studies but upon direct experimental evidence.

Biological studies and practical interests alike make new demands upon systematic botany. Species are not only the subject-material of herbaria and collections, but they are living entities, and their life-history and life-conditions command a gradually increasing interest. One phase of the question is to determine the easiest manner to deal with the collected forms of a country, and another feature is the problem [33] as to what groups are real units and will remain constant and unchanged through all the years of our observations.

Before Linnaeus, the genera were the real units of the system. De Candolle pointed out that the old common names of plants, such as roses and clover, poplars and oaks, nearly all refer to genera. The type of the clovers is rich in color, and the shape of the flower-heads and the single flowers escape ordinary observation; but notwithstanding this, clovers are easily recognized, even if new types come to hand. White and red clovers and many other species are distinguished simply by adjectives, the generic name remaining the same for all.

Tournefort, who lived in the second half of the 17th century (1656-1708), is generally considered as the author of genera in systematic botany. He adopted, what was at that time the general conception and applied it throughout the vegetable kingdom. He grouped the new and the rare and the previously overlooked forms in the same manner in which the more conspicuous plants were already arranged by universal consent. Species were distinguished by minor marks and often indicated by short descriptions, but they were considered of secondary importance.

Based on the idea of a direct creation of all [34] living beings, the genera were then accepted as the created forms. They were therefore regarded as the real existing types, and it was generally surmised that species and varieties owed their origin to subsequent changes under the influence of external conditions. Even Linnaeus agreed with this view in his first treatises and in his "Philosophical Botany" he still kept to the idea that all genera had been created at once with the beginning of life.

Afterwards Linnaeus changed his opinion on this important point, and adopted species as the units of the system. He declared them to be the created forms, and by this decree, at once reduced the genera to the rank of artificial groups. Linnaeus was well aware that this conception was wholly arbitrary, and that even the species are not real indivisible entities. But he simply forbade the study of lesser subdivisions. At his time he was quite justified in doing so, because the first task of the systematic botanists was the clearing up of the chaos of forms and the bringing of them into connection with their real allies.

Linnaeus himself designated the subdivisions of the species as varieties, but in doing so he followed two clearly distinct principles. In some cases his species were real plants, and the varieties seemed to be derived from them by [35] some simple changes. They were subordinated to the parent-species. In other cases his species were groups of lesser forms of equal value, and it was not possible to discern which was the primary and which were the derivatives.

These two methods of subdivision seem in the main, and notwithstanding their relatively imperfect application in many single examples, to correspond with two really distinct cases. The derivative varieties are distinguished from the parent-species by some single, but striking mark, and often this attribute manifests itself as the loss of some apparent quality. The loss of spines and of hairs and the loss of blue and red flower-colors are the most notorious, but in rarer cases many single peculiarities may disappear, thereby constituting a variety. This relation of varieties to the parent-species is gradually increasing in importance in the estimation of botanists, sharply contrasting with those cases, in which such dependency is not to be met with.

If among the subdivisions of a species, no single one can be pointed out as playing a primary part, and the others can not be traced back to it, the relation between these lesser units is of course of another character. They are to be considered of equal importance. They are distinguished from each other by more than [36] one character, often by slight differences in nearly all their organs and qualities. Such forms have come to be designated as "elementary species." They are only varieties in a broad and vague systematic significance of the word, not in the sense accorded to this term in horticultural usage, nor in a sharper and more scientific conception.

Genera and species are, at the present time, for a large part artificial, or stated more correctly, conventional groups. Every systematist is free to delimit them in a wider or in a narrower sense, according to his judgment. The greater authorities have as a rule preferred larger genera, others of late have elevated innumerable subgenera to the rank of genera. This would work no real harm, if unfortunately, the names of the plants had not to be changed each time, according to current ideas concerning genera. Quite the same inconstancy is observed with species. In the Handbook of the British Flora, Bentham and Hooker describe the forms of brambles under 5 species, while Babington in his Manual of British Botany makes 45 species out of the same material. So also in other cases. For instance, the willows which have 13 species in one and 31 species in the other of these manuals, and the hawkweeds for which the figures are 7 and 32 [37] respectively. Other authors have made still greater numbers of species in the same groups.

It is very difficult to estimate systematic differences on the ground of comparative studies alone. All sorts of variability occur, and no individual or small group of specimens can really be considered as a reliable representative of the supposed type. Many original diagnoses of new species have been founded on divergent specimens and of course, the type can afterwards neither be derived from this individual, nor from the diagnosis given.

This chaotic state of things has brought some botanists to the conviction that even in systematic studies only direct experimental evidence can be relied upon. This conception has induced them to test the constancy of species and varieties, and to admit as real units only such groups of individuals as prove to be uniform and constant throughout succeeding generations. The late Alexis Jordan, of Lyons in France, made extensive cultures in this direction. In doing so, he discovered that systematic species, as a rule, comprise some lesser forms, which often cannot easily be distinguished when grown in different regions, or by comparing dried material. This fact was, of course, most distasteful to the systematists of his time and even for a long period afterwards [38] they attempted to discredit it. Milde and many others have opposed these new ideas with some temporary success. Only of late has the school of Jordan received due recognition, after Thuret, de Bary, Rosen and others tested its practices and openly pronounced for them. Of late Wittrock of Sweden has joined them, making extensive experimental studies concerning the real units of some of the larger species of his country.

From the evidence given by these eminent authorities, we may conclude that systematic species, as they are accepted nowadays, are as a rule compound groups. Sometimes they consist of two or three, or a few elementary types, but in other cases they comprise twenty, or fifty, or even hundreds of constant and well differentiated forms.

The inner constitution of these groups is however, not at all the same in all cases. This will be seen by the description of some of the more interesting of them. The European heartsease, from which our garden-pansies have been chiefly derived, will serve as an example. The garden-pansies are a hybrid race, won by crossing the Viola tricolor with the large flowered and bright yellow V. lutea. They combine, as everyone knows, in their wide range of [39] varieties, the attributes of the latter with the peculiarities of the former species.

Besides the lutea, there are some other species, nearly allied to tricolor, as for instance, cornuta, calcarata, and altaica, which are combined with it under the head of Melanium as a subgenus, and which together constitute a systematic unity of undoubted value, but ranging between the common conceptions of genus and species. These forms are so nearly allied to the heartsease that they have of late been made use of in crosses, in order to widen the range of variability of garden-pansies.

Viola tricolor is a common European weed. It is widely dispersed and very abundant, growing in many localities in large numbers. It is an annual and ripens its seeds freely, and if opportunity is afforded, it multiplies rapidly.

Viola tricolor has three subspecies, which have been elevated to the rank of species by some authors, and which may here be called, for brevity's sake, by their binary names. One is the typical V. tricolor, with broad flowers, variously colored and veined with yellow, purple and white. It occurs in waste places on sandy soil. The second is called V. arvensis or the field-pansy; it has small inconspicuous flowers, with pale-yellowish petals which are shorter than the sepals. It pollinates itself without the [40] aid of insects, and is widely dispersed in cultivated fields. The third form, V. alpestris, grows in the Alps, but is of lesser importance for our present discussion.

Anywhere throughout the central part of Europe V. tricolor and V. arvensis may be seen, each occupying its own locality. They may be considered as ranging among the most common native plants of the particular regions they inhabit. They vary in the color of the flowers, branching of the stems, in the foliage and other parts, but not to such an extent as to constitute distinct strains. They have been brought into cultivation by Jordan, Wittrock and others, but throughout Europe each of them constitutes a single type.

These types must be very old and constant, fluctuating always within the same distinct and narrow limits. No slow, gradual changes can have taken place. In different countries their various habitats are as old as the historical records, and probably many centuries older. They are quite independent of one another, the distance being in numerous cases far too great for the exchange of pollen or of seeds. If slow and gradual changes were the rule, the types could not have remained so uniform throughout the whole range of these two species. They would necessarily have split up into thousands [41] and thousands of minor races, which would show their peculiar characteristics if tested by cultures in adjacent beds. This however, is not what happens. As a matter of fact V. tricolor and V. arvensis are widely distributed but wholly constant types.

Besides these, there occur distinct types in numerous localities. Some of them evidently have had time and opportunity to spread more or less widely and now occupy larger regions or even whole countries. Others are narrowly limited, being restricted to a single locality. Wittrock collected seeds or plants from as many localities as possible in different parts of Sweden and neighboring states and sowed them in his garden near Stockholm. He secured seeds from his plants, and grew from them a second, and in many cases a third generation in order to estimate the amount of variability. As a rule the forms introduced into his garden proved constant, notwithstanding the new and abnormal conditions under which they were propagated.

First of all we may mention three perennial forms called by him Viola tricolor ammotropha, V. tricolor coniophila and V. stenochila. The typical V. tricolor is an annual plant; sowing itself in summer and germinating soon afterwards. The young plants thrive throughout [42] the latter part of the summer and during the fall, reaching an advanced stage of development of the branched stems before winter. Early in the spring the flowers begin to open, but after the ripening of the seeds the whole plant dies.

The three perennial species just mentioned develop in the same manner in the first year. During their flowering period, however, and afterwards, they produce new shoots from the lower parts of the stem. They prefer dry and sandy soils, often becoming covered with the sand that is blown on them by the winds. They are prepared for such seemingly adverse circumstances by the accumulation of food in the older stems and by the capacity of the new shoots to thrive on this food till they have become long enough to reach the light. V. tricolor ammotropha is native near Ystad in Sweden, and the other two forms on Gotland. All three have narrowly limited habitats.

The typical tricolored heartsease has remained annual in all its other subspecies. It may be divided into two types in the first place, _V. tricolor genuina_ and _V. tricolor versicolor_. Both of them have a wide distribution and seem to be the prototypes from which the rarer forms must have been derived. Among these latter Wittrock describes seven local types, which [43] proved to be constant in his pedigree-cultures. Some of them have produced other forms, related to them in the way of varieties. They all have nearly the same general habit and do not exhibit any marked differences in their growth, in the structure and branching of the stems, or in the character of their foliage. Differentiating points are to be found mainly in the colors and patterns of the flowers. The veins, which radiate from the centre of the corolla are branched in some and undivided in others; in one elementary species they are wholly lacking. The purple color may be absent, leaving the flowers of a pale or a deep yellow. Or the purple may be reddish or bluish. Of the petals all five may have the purple hue on their tips, or this attribute may be limited to the two upper ones. Contrasting with this wide variability is the stability of the yellow spot in the centre, which is always present and becomes inconspicuous only, when the whole petals are of the same hue. It is a general conception that colors and color-markings are liable to great variability and do not constitute reliable standards. But the cultures of Wittrock have proved the contrary, at least in the case of the violets. No pattern, however quaint, appears changeable, if one elementary species only is considered. Hundreds of plants from seeds [44] from one locality may be grown, and all will exhibit exactly the same markings. Most of these forms are of very local occurrence. The most beautiful of all, the _ornatissima_, is found only in Jemtland, the _aurobadia_ only in Sodermanland, the anopetala_ in other localities in the same country, the _roseola_ near Stockholm, and the yellow _lutescens_ in Finmarken.

The researches of Wittrock included only a small number of elementary species, but every one who has observed the violets in the central parts of Europe must be convinced that many dozens of constant forms of the typical Viola tricolor might easily be found and isolated.

We now come to the field pansy, the Viola arvensis, a very common weed in the grain-fields of central Europe. I have already mentioned its small corolla, surpassed by the lobes of the calyx and its capacity of self-fertilization. It has still other curious differentiating characters; the pollen grains, which are square in V. tricolor, are five-sided in V. arvensis. Some transgressive fluctuating variability may occur in both cases through the admixture of pollen-grains. Even three-angled pollen grains are seen sometimes. Other marks are observed in the form of the anthers and the spur.

There seem to be very many local subspecies [45] of the field-pansy. Jordan has described some from the vicinity of Lyons, and Wittrock others from the northern parts of Europe. They diverge from their common prototype in nearly all attributes, the flowers not showing the essential differentiating characters as in the V. tricolor. Some have their flower-stalks erect, and in others the flowers are held nearly at right angles to the stem. V. pallescens is a small, almost unbranched species with small pale flowers. V. segetalis is a stouter species with two dark blue spots on the tips of the upper petals. V. agrestis is a tall and branched, hairy form. V. nemausensis attains a height of only 10 cm., has rounded leaves and long flower-stalks. Even the seeds afford characters which may be made use of in isolating the various species.

The above-mentioned elementary forms belong to the flora of southern France, and Wittrock has isolated and cultivated a number of others from the fields of Sweden. A species from Stockholm is called Viola patens; V. arvensis curtisepala occurs in Gotland, and V. arvensis striolata is a distinct form, which has appeared in his cultures without its true origin being ascertained.

The alpine violets comprise a more widespread type with some local elementary species [46] derived exactly in the same way as the tricolored field pansies.

Summarizing the general result of this description we see that the original species Viola tricolor may be split up into larger and lesser groups of separate forms. These last prove to be constant in pedigree-cultures, and therefore are to be considered as really existent units. They are very numerous, comprising many dozens in each of the two larger subdivisions.

All systematic grouping of these forms, and their combination into subspecies and species rests on the comparative study of their characters. The result of such studies must necessarily depend on principles which underlie them. According to the choice of these principles, the construction of the groups will be found to be different. Wittrock trusts in the first place to morphologic characters, and considers the development as passing from the more simple to the more complex types. On the other hand the geographic distribution may be considered as an indication of the direction of evolution, the wide-spread forms being regarded as the common parents of the minor local species.

However, such considerations are only of secondary importance. It must be borne in mind that an ordinary systematic species may include [47] many dozens of elementary forms, each of which remains constant and unchanged in successive generations, even if cultivated in the same garden and under similar external conditions.

Leaving the violets, we may take the vernal whitlow-grass or Draba verna for a second illustration. This little annual cruciferous plant is common in the fields of many parts of the United States, though originally introduced from Europe. It has small basal rosettes which develop during summer and winter, and produce numerous leafless flowering stems early in the spring. It is a native of central Europe and western Asia, and may be considered as one of the most common plants, occurring anywhere in immense numbers on sandy soils. Jordan was the first to point out that it is not the same throughout its entire range. Although a hasty survey does not reveal differences, they show themselves on closer inspection. De Bary, Thuret, Rosen and many others confirmed this result, and repeated the pedigree-cultures of Jordan. Every type is constant and remains unchanged in successive generations. The anthers open in the flower-buds and pollinate the stigmas before the expansion of the flowers, thus assuring self-fertilization. Moreover, these inconspicuous little flowers are only sparingly visited by insects. Dozens of subspecies [48] may be cultivated in the same garden without any real danger of their intercrossing. They remain as pure as under perfect isolation.

It is very interesting to observe the aspect of such types, when growing near each other. Hundreds of rosettes exhibit one type, and are undoubtedly similar. The alternative group is distinguishable at first sight, though the differentiating marks are often so slight as to be traceable with difficulty. Two elementary species occur in Holland, one with narrow leaves in the western provinces and one with broader foliage in the northern parts. I have cultivated them side by side, and was as much struck with the uniformity within each group, as with the contrast between the two sets.

Nearly all organs show differences. The most marked are those of the leaves, which may be small or large, linear or elliptic or oblong and even rhomboidal in shape, more or less hairy with simple or with stellate branched hairs, and finally of a pure green or of a glaucous color. The petals are as a rule obcordate, but this type may be combined with others having more or less broad emarginations at the summit, and with differences in breadth which vary from almost linear types to others which touch along their margins. The pods are short and broad, or long and narrow, or varying in sundry other [49] ways. All in all there are constant differences which are so great that it has been possible to distinguish and to describe large numbers of types.

Many of them have been tested as to their constancy from seed. Jordan made numerous cultures, some of which lasted ten or twelve years; Thuret has verified the assertion concerning their constancy by cultures extending over seven years in some instances; Villars and de Bary made numerous trials of shorter duration. All agree as to the main points. The local races are uniform and come true from seed; the variability of the species is not of a fluctuating, but of a polymorphous nature. A given elementary species keeps within its limits and cannot vary beyond them, but the whole group gives the impression of variability by its wide range of distinct, but nearly allied forms.

The geographic distribution of these elementary species of the whitlow-grass is quite distinct from that of the violets. Here predominant species are limited to restricted localities. Most of them occupy one or more departments of France, and in Holland two of them are spread over several provinces. An important number are native in the centre of Europe, and from the vicinity of Lyons, Jordan succeeded in establishing about fifty elementary [50] species in his garden. In this region they are crowded together and not rarely two or even more quite distinct forms are observed to grow side by side on the same spot. Farther away from this center they are more widely dispersed, each holding its own in its habitat. In all, Jordan has distinguished about two hundred species of Draba verna from Europe and western Asia. Subsequent authors have added new types to the already existing number from time to time.

The constancy of these elementary species is directly proven by the experiments quoted above, and moreover it may be deduced from the uniformity of each type within its own domain. These are so large that most of the localities are practically isolated from one another, and must have been so for centuries. If the types were slowly changing such localities would often, though of course not always, exhibit slighter differences, and on the geographic limits of neighboring species intermediates would be found. Such however, are not on record. Hence the elementary species must be regarded as old and constant types.

The question naturally arises how these groups of nearly allied forms may originally have been produced. Granting a common origin for all of them, the changes may have been [51] simultaneous or successive. According to the geographic distribution, the place of common origin must probably be sought in the southern part of central Europe, perhaps even in the vicinity of Lyons. Here we may assume that the old Draba verna has produced a host or a swarm of new types. Thence they must have spread over Europe, but whether in doing so they have remained constant, or whether some or many of them have repeatedly undergone specific mutations, is of course unknown.

The main fact is, that such a small species as Draba verna is not at all a uniform type, but comprises over two hundred well distinguished and constant forms.

It is readily granted that violets and whitlowgrasses are extreme instances of systematic variability. Such great numbers of elementary species are not often included in single species of the system. But the numbers are of secondary importance, and the fact that systematic species consist, as a rule, of more than one independent and constant subspecies, retains its almost universal validity.

In some cases the systematic species are manifest groups, sharply differentiated from one another. In other instances the groups of elementary forms as they are shown by direct observation, have been adjudged by many authors [52] to be too large to constitute species. Hence the polymorphous genera, concerning the systematic subdivisions of which hardly two authors agree. Brambles and roses are widely known instances, but oaks, elms, apples, and pears, Mentha, Prunus, Vitis, Lactuca, Cucumis, Cucurbita and numerous others are in the same condition.

In some instances the existence of elementary species is so obvious, that they have been described by taxonomists as systematic varieties or even as good species. The primroses afford a widely known example. Linnaeus called them Primula veris, and recognized three types as pertaining to this species, but Jacquin and others have elevated these subspecies to the full rank of species. They now bear the names of Primula elatior with larger, P. officinalis with smaller flowers, and P. acaulis. In the last named the common flower-stalk is lacking and the flowers of the umbel seem to be borne in the arils of the basal leaves.

In other genera such nearly allied species are more or less universally recognized. Galium Mollugo has been divided into G. elatum with a long and weak stem, and G. erectum with shorter and erect stems; Cochlearia danica, anglica and officinalis are so nearly allied as to be hardly distinguishable. Sagina apetala and patula, [53] Spergula media and salina and many other pairs of allied species have differentiating characters of the same value as those of the elementary species of Draba verna. Filago, Plantago, Carex, Ficaria and a long series of other genera afford proofs of the same close relation between smaller and larger groups of species. The European frost-weeds or Helianthemum include a group of species which are so closely allied, that ordinary botanical descriptions are not adequate to give any idea of their differentiating features. It is almost impossible to determine them by means of the common analytical keys. They have to be gathered from their various native localities and cultivated side by side in the garden to bring out their differences. Among the species of France, according to Jordan, Helianthemum polifolium, H. apenninum, H. pilosum and H. pulverulentum are of this character.

A species of cinquefoil, Potentilla Tormentilla, which is distinguished by its quaternate flowers, occurs in Holland in two distinct types, which have proved constant in my cultural experiments. One of them has, broad petals, meeting together at the edges, and constituting rounded saucer without breaks. The other has narrow petals, which are strikingly separated from one another and show the sepals between them. [54] In the same manner bluebells vary in the size and shape of the corolla, which may be wide or narrow, bell-shaped or conical, with the tips turned downwards, sidewards or backwards.

As a rule all of the more striking elementary types have been described by local botanists under distinct specific names, while they are thrown together into the larger systematic species by other authors, who study the distribution of plants over larger portions of the world. Everything depends on the point of view taken. Large floras require large species. But the study of local floras yields the best results if the many forms of the region are distinguished and described as completely as possible. And the easiest way is to give to each of them a specific name. If two or more elementary species are united in the same district, they are often treated in this way, but if each region had its own type of some given species, commonly the part is taken for the whole, and the sundry forms are described under the same name, without further distinctions.

Of course these questions are all of a practical and conventional nature, but involve the different methods in which different authors deal with the same general fact. The fact is that systematic species are compound groups, exactly like the genera and that their real units [55] can only be recognized by comparative experimental studies.

Though the evidence already given might be esteemed to be sufficient for our purpose, I should like to introduce a few more examples; two of them pertain to American plants.

The Ipecac spurge or Euphorbia Ipecacuanha occurs from Connecticut to Florida, mainly near the coast, preferring dry and sandy soil. It is often found by the roadsides. According to Britton and Brown's "Illustrated Flora" it is glabrous or pubescent, with several or many stems, ascending or nearly erect; with green or red leaves, which are wonderfully variable in outline, from linear to orbicular, mostly opposite, the upper sometimes whorled, the lower often alternate. The glands of the involucres are elliptic or oblong, and even the seeds vary in shape.

Such a wide range of variability evidently points to the existence of some minor types. Dr. John Harshberger has made a study of those which occur in the vicinity of Whitings in New Jersey. His types agree with the description given above. Others were gathered by him at Brown's Mills in the pinelands, New Jersey, where they grew in almost pure sand in the bright sunlight. He observed still other differentiating characters. The amount of seed [56] produced and the time of flowering were variable to a remarkable degree.

Dr. Harshberger had the kindness to send me some dried specimens of the most interesting of these types. They show that the peculiarities are individual, and that each specimen has its own characters. It is very probable that a comparative experimental study will prove the existence of a large number of elementary species, differing in many points; they will probably also show differences in the amount of the active chemical substances, especially of emetine, which is usually recorded as present in about 1%, but which will undoubtedly be found in larger quantities in some, and in smaller quantities in other elementary species. In this way the close and careful distinction of the really existing units might perhaps prove of practical importance.

MacFarlane has studied the beach-plum or Prunus maritima, which is abundant along the coast regions of the Eastern States from Virginia to New Brunswick. It often covers areas from two to two hundred acres in extent, sometimes to the exclusion of other plants. It is most prolific on soft drifting sand near the sea or along the shore, where it may at times be washed with ocean-spray. The fruit usually become ripe about the middle of August, and show extreme [57] variations in size, shape, color, taste, consistency and maturation period, indicating the existence of separate races or elementary species, with widely differing qualities. The earlier varieties begin to ripen from August 10 to 20, and a continuous supply can be had till September 10, while a few good varieties continue to ripen till September 20. But even late in October some other types are still found maturing their fruits.

Exact studies were made of fruit and stone variations, and their characteristics as to color, weight, size, shape and consistency were fully described. Similar variations have been observed, as is well known, in the cultivated plums. Fine blue-black fruits were seen on some shrubs and purplish or yellow fruits on others. Some exhibit a firmer texture and others a more watery pulp. Even the stones show differences which are suggestive of distinct races.

Recently Mr. Luther Burbank of Santa Rosa, California, has made use of the beach-plum to produce useful new varieties. He observed that it is a very hardy species, and never fails to bear, growing under the most trying conditions of dry and sandy, or of rocky and even of heavy soil. The fruits of the wild shrubs are utterly worthless for anything but preserving. [58] But by means of crossing with other species and especially with the Japanese plums, the hardy qualities of the beach-plum have been united with the size, flavor and other valuable qualities of the fruit, and a group of new plums have been produced with bright colors, ovoid and globular forms which are never flattened and have no suture. The experiments were not finished, when I visited Mr. Burbank in July, 1904, and still more startling improvements were said to have been secured.

I may perhaps be allowed to avail myself of this opportunity to point out a practical side of the study of elementary species. This always appears whenever wild plants are subjected to cultivation, either in order to reproduce them as pure strains, or to cross them with other already cultivated species. The latter practice is as a rule made use of whenever a wild species is found to be in possession of some quality which is considered as desirable for the cultivated forms. In the case of the beach-plum it is the hardiness and the great abundance of fruits of the wild species which might profitably be combined with the recognized qualities of the ordinary plums. Now it is manifest, that in order to make crosses, distinct individual plants are to be chosen, and that the variability of the wild species may be of very great importance. [59] Among the range of elementary species those should be used which not only possess the desired advantages in the highest degree, but which promise the best results in other respects or their earliest attainment. The fuller our knowledge of the elementary species constituting the systematic groups, the easier and the more reliable will be the choice for the breeder. Many Californian wild flowers with bright colors seem to consist of large numbers of constant elementary forms, as for instance, the lilies, godetias, eschscholtias and others. They have been brought into cultivation many times, but the minutest distinction of their elementary forms is required to attain the highest success.

In concluding, I will point out a very interesting difficulty, which in some cases impedes the clear understanding of elementary species. It is the lack of self-fertilization. It occurs in widely distant families, but has a special interest for us in two genera, which are generally known as very polymorphous groups.

One of them is the hawkweed or Hieracium, and the other is the dandelion or Taraxacum officinale. Hawkweeds are known as a genus in which the delimitation of the species is almost impossible, Thousands of forms may be cultivated side by side in botanical gardens, exhibiting [60] slight but undoubted differentiating features, and reproduce themselves truly by seed. Descriptions were formerly difficult and so complicated that the ablest writers on this genus, Fries and Nageli are said not to have been able to recognize the separate species by the descriptions given by each other. Are these types to be considered as elementary species, or only as individual differences? The decision of course, would depend upon their behavior in cultures. Such tests have been made by various experimenters. In the dandelion the bracts of the involucre give the best characters. The inner ones may be linear or linear-lanceolate, with or without appendages below the tip; the outer ones may be similar and only shorter, or noticeably larger, erect, spreading or even reflexed, and the color of the involucre may be a pure green or glaucous; the leaves may be nearly entire or pinnatifid, or sinuate-dentate, or very deeply runcinate-pinnatifid, or even pinnately divided, the whole plant being more or less glabrous.

Raunkiaer, who has studied experimentally a dozen types from Denmark, found them constant, but observed that some of them have no pollen at all, while in others the pollen, though present, is impotent. It does not germinate on the stigma, cannot produce the ordinary tube, [61] and hence has no fertilizing power. But the young ovaries do not need such fertilization. They are sufficient unto themselves. One may cut off all the flowers of a head before the opening of the anthers, and leave the ovaries untouched, and the head will ripen its seeds quite as well. The same thing occurs in the hawkweeds. Here, therefore, we have no fertilization and the extensive widening of the variability, which generally accompanies this process is, of course, wanting. Only partial or vegetative variability is present. Unfertilized eggs when developing into embryos are equivalent to buds, separated from the parent-plant and planted for themselves. They repeat both the specific and the individual characters of the parent. In the case of the hawkweed and the dandelion there is at present no means of distinguishing between these two contrasting causes of variability. But like the garden varieties which are always propagated in the vegetative way, their constancy and uniformity are only apparent and afford no real indication of hereditary qualities.

In addition to these and other exceptional cases, seed-cultures are henceforth to be considered as the sole means of recognizing the really existing systematic units of nature. All other groups, including systematic species and [62] genera, are equally artificial or conventional. In other words we may state "that current misconceptions as to the extreme range of fluctuating variability of many native species have generally arisen from a failure to recognize the composite nature of the forms in question," as has been demonstrated by MacDougal in the case of the common evening-primrose, Oenothera biennis. "It is evident that to study the behavior of the characters of plants we must have them in their simplest combinations; to investigate the origin and movements of species we must deal with them singly and uncomplicated."

[63]

LECTURE III

ELEMENTARY SPECIES OF CULTIVATED PLANTS

Recalling the results of the last lecture, we see that the species of the systematists are not in reality units, though in the ordinary course of floristic studies they may, as a rule, seem to be so. In some cases representatives of the same species from different countries or regions, when compared with one another do not exactly agree. Many species of ferns afford instances of this rule, and Lindley and other great systematists have frequently been puzzled by the wide range of differences between the individuals of a single species.

In other cases the differing forms are observed to grow near each other, sometimes in neighboring provinces, sometimes in the same locality, growing and flowering in mixtures of two or three or even more elementary types. The violets exhibit widespread ancient types, from which the local species may be taken to have arisen. The common ancestors of the Whitlow-grasses are probably not to be found [64] among existing forms, but numerous types are crowded together in the southern part of central Europe and more thinly scattered elsewhere, even as far as western Asia. There can be little doubt that their common origin is to be sought in the center of their geographic distribution.

Numerous other cases exhibit smaller numbers of elementary units within a systematic species; in fact purely uniform species seem to be relatively rare. But with small numbers there are of course no indications to be expected concerning their common origin or the starting point of their distribution.

It is manifest that these experiences with wild species must find a parallel among cultivated plants. Of course cultivated plants were originally wild and must have come under the general law. Hence we may conclude that when first observed and taken up by man, they must already have consisted of sundry elementary subspecies. And we may confidently assert that some must have been rich and others poor in such types.

Granting this state of things as the only probable one, we can easily imagine what must have been the consequences. If a wild species had been taken into cultivation only once, the cultivated form would have been a single elementary [65] type. But it is not very likely that such partiality would occur often. The conception that different tribes at different times and in distant countries would have used the wild plants of their native regions seems far more natural than that all should have obtained plants for cultivation from the same source or locality. If this theory may be relied upon, the origin of many of the more widely cultivated agricultural plants must have been multiple, and the number of the original elementary species of the cultivated types must have been so much the larger, the more widely distributed and variable the plants under consideration were before the first period of cultivation.

Further it would seem only natural to explain the wide variability of many of our larger agricultural and horticultural stocks by such an incipient multiformity of the species themselves. Through commercial intercourse the various types might have become mixed so as to make it quite impossible to point out the native localities for each of them.

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