Chapter 4

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_, _Prunu_s, _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.