The purpose of these lectures is to discuss some of the familiar phenomena of biology in the light of modern discoveries. In the last decade of the nineteenth century many of us perceived that if any serious advance was to be made with the group of problems generally spoken of as the Theory of Evolution, methods of investigation must be devised and applied of a kind more direct and more penetrating than those which after the general acceptance of the Darwinian views had been deemed adequate. Such methods obviously were to be found in a critical and exhaustive study of the facts of variation and heredity, upon which all conceptions of evolution are based. To construct a true synthetic theory of Evolution it was necessary that variation and heredity instead of being merely postulated as axioms should be minutely examined as phenomena. Such a study Darwin himself had indeed tentatively begun, but work of a more thorough and comprehensive quality was required. In the conventional view which the orthodoxy of the day prescribed, the terms variation and heredity stood for processes so vague and indefinite that no analytical investigation of them could be contemplated. So soon, however, as systematic inquiry into the natural facts was begun it was at once found that the accepted ideas of variation were unfounded. Variation was seen very frequently to be a definite and specific phenomenon, affecting different forms of life in different ways, but in all its diversity showing manifold and often obvious indications of regularity. This observation was not in its essence novel. Several examples of definite variation had been well known to Darwin and others, but many, especially Darwin himself in his later years, had nevertheless been disposed to depreciate the significance of such facts. They consequently then lapsed into general disparagement. Upon more careful inquiry the abundance of such phenomena proved to be far greater than was currently supposed, and a discussion of their nature brought into prominence a consideration of greater weight, namely that the differences by which these definite or discontinuous variations are constituted again and again approximate to and are comparable with the class of differences by which species are distinguished from each other.

The interest of such observations could no longer be denied. The more they were examined the more apparent it became that by means of the facts of variation a new light was obtained on the physiological composition and capabilities of living things. Genetics thus cease to be merely a method of investigating theories of evolution or of the origin of species but provide a novel and hitherto untried instrument by which the nature of the living organism may be explored. Just as in the study of non-living matter science began by regarding the external properties of weight, opacity, colour, hardness, mode of occurrence, etc., noting only such evidences of chemical attributes and powers as chance spontaneously revealed; and much later proceeded to the discovery that these casual manifestations of chemical properties, rightly interpreted, afford a key to the intrinsic nature of the diversity of matter, so in biology, having examined those features of living things which ordinary observations can perceive, we come at last to realize that when studied for their own sake the properties of living organisms in respect of heredity and variation are indications of their inner nature and provide evidences of that nature which can be obtained from no other source.

While such ideas were gradually forming in our minds, came the rediscovery of Mendel's work. Investigations which before had only been imagined as desirable now became easy to pursue, and questions as to the genetic inter-relations and compositions of varieties can now be definitely answered. Without prejudice to what the future may disclose whether by way of limitation or extension of Mendelian method, it can be declared with confidence and certainty that we have now the means of beginning an analysis of living organisms, and distinguishing many of the units or factors which essentially determine and cause the development of their several attributes.

Briefly put, the essence of Mendelism lies in the discovery of the existence of unit characters or factors. For an account of the Mendelian method, how it is applied and what it has already accomplished, reference must be made to other works.[1] With this part of the subject I shall assume a sufficient acquaintance. In these lectures I have rather set myself the task of considering how certain problems appear when viewed from the standpoint to which the application of these methods has led us. It is indeed somewhat premature to discuss such questions. The work of Mendelian analysis is progressing with great rapidity and anything I can say may very soon be superseded as out of date. Nevertheless a discussion of this kind may be of at least temporary service in directing inquiry to the points of special interest.

The Problem of Species and Variety

Nowhere does our new knowledge of heredity and variation apply more directly than to the problem what is a species and what is a variety? I cannot assert that we are already in a position to answer this important question, but as will presently appear, our mode of attack and the answers we expect to receive are not those that were contemplated by our predecessors. If we glance at the history of the scientific conception of Species we find many signs that it was not till comparatively recent times that the definiteness of species became a strict canon of the scientific faith and that attempts were made to give precise limits to that conception. When the diversity of living things began to be accurately studied in the sixteenth and seventeenth centuries names were applied in the loosest fashion, and in giving a name to an animal or a plant the naturalists of those times had no ulterior intention. Names were bestowed on those creatures about which the writer proposed to speak. When Gesner or Aldrovandi refer to all the kinds of horses, unicorns, dogs, mermaids, etc., which they had seen or read of, giving to each a descriptive name, they do not mean to "elevate" each named kind to "specific rank"; and if anyone had asked them what they meant by a species, it is practically certain that they would have had not the slightest idea what the question might imply, or any suspicion that it raised a fundamental problem of nature.

Spontaneous generation being a matter of daily observation, then unquestioned, and supernatural events of all kinds being commonly reported by many witnesses, transmutation of species had no inherent improbability. Matthioli,[2] for instance, did not expect to be charged with heresy when he declared Stirpium mutatio to be of ordinary occurrence. After giving instances of induced modifications he wrote, "Tantum enim in plantis naturae germanitas potest, ut non solum saepe praedictos praestet effectus, sed etiam ut alteram in alteram stirpem facile vertat, ut cassiam in cinnamomum, sisymbrium in mentham, triticum in lolium, hordeum in avenam, et ocymum in serpyllum."

I do not know who first emphasized the need for a clear understanding of the sense in which the term species is to be applied. In the second half of the seventeenth century Ray shows some degree of concern on this matter. In the introduction to the Historia Plantarum, 1686, he discusses some of the difficulties and lays down the principle that varieties which can be produced from the seed of the same plant are to be regarded as belonging to one species, being, I believe, the first to suggest this definition. That new species can come into existence he denies as inconsistent with Genesis 2, in which it is declared that God finished the work of Creation in six days. Nevertheless he does not wholly discredit the possibility of a "transmutation" of species, such that one species may as an exceptional occurrence give rise by seed to another and nearly allied species. Of such a phenomenon he gives illustrations the authenticity of which he says he is, against his will, compelled to admit. He adds that some might doubt whether in the cases quoted the two forms concerned are really distinct species, but the passage is none the less of value for it shews that the conception of species as being distinct unchangeable entities was not to Ray the dogma sacrosanct and unquestionable which it afterwards became.[3]

In the beginning of the eighteenth century Marchant,[4] having observed the sudden appearance of a lacinated variety of Mercurialis, makes the suggestion that species in general may have arisen by similar mutations. Indeed from various passages it is manifest that to the authors of the seventeenth and early eighteenth centuries species appeared simply as groups more or less definite, the boundaries of which it was unnecessary to determine with great exactitude. Such views were in accord with the general scientific conception of the time. The mutability of species is for example sometimes likened (see for instance Sharrock, loc. cit.) to the metamorphoses of insects, and it is to be remembered that the search for the Philosopher's Stone by which the transmutation of metals was to be effected had only recently fallen into discredit as a pursuit.

The notion indeed of a peculiar, fixed meaning to be attached to species as distinct from variety is I think but rarely to be found categorically expressed in prae-Linnaean writings.

But with the appearance of the Systema Naturae a great change supervened. Linnaeus was before all a man of order. Foreseeing the immense practical gain to science that must come from a codification of nomenclature, he invented such a system.

It is not in question that Linnaeus did great things for us and made Natural History a manageable and accessible collection of facts instead of a disorderly heap; but orderliness of mind has another side, and inventors and interpreters of systems soon attribute to them a force and a precision which in fact they have not.

The systematist is primarily a giver of names, as Ray with his broader views perceived. Linnaeus too in the exordium to the Systema Naturae naively remarks, that he is setting out to continue the work which Adam began in the Golden Age, to give names to the living creatures. Naming however involves very delicate processes of mind and of logic. Carried out by the light of meagre and imperfect knowledge it entails all the mischievous consequences of premature definition, and promotes facile illusions of finality. So was it with the Linnaean system. An interesting piece of biological history might be written respecting the growth and gradual hardening of the conception of Species. To readers of Linnaeus's own writings it is well known that his views cannot be summarized in a few words. Expressed as they were at various times during a long life and in various connexions, they present those divers inconsistencies which commonly reflect a mind retaining the power of development. Nothing certainly could be clearer than the often quoted declaration of the Philosophia Botanica, "Species tot numeramus quot diversae formae in principio sunt creatae," with the associated passage "Varietates sunt plantae ejusdem speciei mutatae a caussa quacunque occasionali." Those sayings however do not stand alone. In several places, notably in the famous dissertation on the peloric Linaria he explicitly contemplates the possibility that new species may arise by crossing, declaring nevertheless that he thinks such an event to be improbable. In that essay he refers to Marchant's observation on a laciniate Mercurialis, but though he states clearly that that plant should only be regarded as a variety of the normal, he does not express any opinion that the contemporary genesis of new species must be an impossibility. In the later dissertation on Hybrid Plants he returns to the same topic. Again though he states the belief that species cannot be generated by cross-breedings, he treats the subject not as heretical absurdity but as one deserving respectful consideration.

The significance of the aphorisms that precede the lectures on the Natural Orders is not easy to apprehend. These are expressed with the utmost formality, and we cannot doubt that in them we have Linnaeus's own words, though for the record we are dependent on the transcripts of his pupils.

The text of the first five is as follows:

1. Creator T. O. in primordio vestiit Vegetabile Medullare principiis constitutivis diversi Corticalis unde tot difformia individua, quot Ordines Naturales prognata.

2. Classicas has (1) plantas Omnipotens miscuit inter se, unde tot Genera ordinum, quot inde plantae.

3. Genericas has (2) miscuit Natura, unde tot Species congeneres quot hodie existunt.

4. Species has miscuit Casus, unde totidem quot passim occurrunt, Varietates.

5. Suadent haec (1-4) Creatoris leges a simplicibus ad Composita.

Naturae leges generationis in hybridis.

Hominis leges ex observatis a posteriori.

I am not clear as to the parts assigned in the first sentence respectively to the "Medulla" and the "Cortex," beyond that Linnaeus conceived that multiformity was first brought about by diversity in the "Cortex." The passage is rendered still more obscure if read in connection with the essay on "Generatio Ambigena," where he expresses the conviction that the Medulla is contributed by the mother, and the Cortex by the father, both in plants and animals.[5]

But however that may be, he regards this original diversity as resulting in the constitution of the Natural Orders, each represented by one individual.

In the second aphorism the Omnipotent is represented as creating the genera by intermixing the individual plantae classicae, or prototypes of the Natural Orders.

The third statement is the most remarkable, for in it he declares that Species were formed by the act of Nature, who by inter-mixing the genera produced Species congeneres, namely species inside each genus, to the number which now exist. Lastly, Chance or Accident, intermixing the species, produced as many varieties as there are about us.

Linnaeus thus evidently regarded the intermixing of an originally limited number of types as the sufficient cause of all subsequent diversity, and it is clear that he draws an antithesis between Creator, Natura, and Casus, assigning to each a special part in the operations. The acts resulting in the formation of genera are obviously regarded as completed within the days of the Creation, but the words do not definitely show that the parts played by Nature and Chance were so limited.

Recently also E. L. Greene[6] has called attention to some curious utterances buried in the Species Plantarum, in which Linnaeus refers to intermediate and transitional species, using language that even suggests evolutionary proclivities of a modern kind, and it is not easy to interpret them otherwise.

Whatever Linnaeus himself believed to be the truth, the effect of his writings was to induce a conviction that the species of animals and plants were immutably fixed. Linnaeus had reduced the whole mass of names to order and the old fantastical transformations with the growth of knowledge had lapsed into discredit; the fixity of species was taken for granted, but not till the overt proclamation of evolutionary doctrine by Lamarck do we find the strenuous and passionate assertions of immutability characteristic of the first half of the nineteenth century.

It is not to be supposed that the champions of fixity were unacquainted with varietal differences and with the problem thus created, but in their view these difficulties were apparent merely, and by sufficiently careful observation they supposed that the critical and permanent distinctions of the true species could be discovered, and the impermanent variations detected and set aside.

This at all events was the opinion formed by the great body of naturalists at the end of the eighteenth and beginning of the nineteenth centuries, and to all intents and purposes in spite of the growth of evolutionary ideas, it remains the guiding principle of systematists to the present day. There are 'good species' and 'bad species' and the systematists of Europe and America spend most of their time in making and debating them.

In some of its aspects the problem of course confronted earlier naturalists. Parkinson for instance (1640) in introducing his treatment of Hieracium wrote, "To set forth the whole family of the Hawkeweedes in due forme and order is such a world of worke that I am in much doubt of mine own abilitie, it having lyen heavie on his shoudiers that hath already waded through them ... for such a multitude of varieties in forme pertaining to one herbe is not to be found againe in rerum natura as I thinke," and the same idea, that the difficulty lay rather in man's imperfect powers of discrimination than in the nature of the materials to be discriminated, is reflected in many treatises early and late.

It was however with the great ouburst of scientific activity which followed Linnaeus that the difficulty became acute. Simultaneously vast masses of new material were being collected from all parts of the world into the museums, and the products of the older countries were re-examined with a fresh zeal and on a scale of quantity previously unattempted. But the problem how to name the forms and where to draw lines, how much should be included under one name and where a new name was required, all this was felt, rather as a cataloguer's difficulty than as a physiological problem. And so we still hear on the one hand of the confusion caused by excessive "splitting" and subdivisions, and on the other of the uncritical "lumpers" who associate together under one name forms which another collector or observer would like to see distinguished.

In spite of Darwin's hopes, the acceptance of his views has led to no real improvement—scarcely indeed to any change at all in either the practice or aims of systematists. In a famous passage in the Origin he confidently declares that when his interpretation is generally adopted "Systematists will be able to pursue their labours as at present; but they will not be incessantly haunted by the shadowy doubt whether this or that form be a true species. This, I feel sure, and I speak after experience, will be no slight relief. The endless disputes whether or not some fifty species of British brambles are good species will cease." Those disputes nevertheless proceed almost exactly as before. It is true that biologists in general do not, as formerly, participate in these discussions because they have abandoned systematics altogether; but those who are engaged in the actual work of naming and cataloguing animals and plants usually debate the old questions in the old way. There is still the same divergence of opinion and of practice, some inclining to make much of small differences, others to neglect them.

Not only does the work of the systematists as a whole proceed as if Darwin had never written but their attitude towards these problems is but little changed. In support of this statement I may refer to several British Museum Catalogues, much of the Biologia Centrali-Americana, Ridgway's Birds of North America, the Fauna Hawaiensis, indeed to almost any of the most important systematic publications of England, America, or any other country. These works are compiled by the most proficient systematists of all countries in the several groups, but with rare exceptions they show little misgiving as to the fundamental reality of specific differences. That the systematists consider the species-unit as of primary importance is shown by the fact that the whole business of collection and distribution of specimens is arranged with regard to it.

Almost always the collections are arranged in such a way that the phenomena of variation are masked. Forms intermediate between two species are, if possible, sorted into separate boxes under a third specific name. If a species is liable to be constantly associated with a mutational form, the mutants are picked out, regardless of the circumstances of their origin, from the samples among which they were captured, and put apart under a special name. Only by a minute study of the original labels of the specimens and by redistributing them according to locality and dates, can their natural relations be traced. The published accounts of these collections often take no notice of variations, others make them the subject of casual reference. Very few indeed treat them as of much importance. From such indications it is surely evident that the systematists attach to the conception of species a significance altogether different from that which Darwin contemplated.

I am well aware that some very eminent systematists regard the whole problem as solved. They hold as Darwin did that specific diversity has no physiological foundation or causation apart from fitness, and that species are impermanent groups, the delimitations of which are ultimately determined by environmental exigency or "fitness." The specific diversity of living things is thus regarded as being something quite different in nature from the specific diversity of inorganic substances. In practice those who share these opinions are, as might be anticipated, to be found among the 'lumpers' rather than among the 'splitters.' In their work, certainly, the Darwinian theory is actually followed as a guiding principle; unanalysed inter-gradations of all kinds are accepted as impugning the integrity of species; the underlying physiological problem is forgotten, and while the product is almost valueless as a contribution to biological research, I can scarcely suppose that it aids greatly in the advances of other branches of our science.

But why is it that, with these exceptions, the consequences of the admittedly general acceptance of a theory of evolution are so little reflected in the systematic treatment of living things? Surely the reason is that though the systematist may be convinced of the general truth of the evolution theory at large, he is still of opinion that species are really distinct things. For him there are still 'good' species and 'bad' species and his experience tells him that the distinction between the two is not simply a question of degree or a matter of opinion.

To some it may seem that this is mere perversity, a refusal to see obvious truth, a manifestation of the spirit of the collector rather than of the naturalist. But while recognising that from a magnification of the conception of species the systematists are occasionally led into absurdity I do not think the grounds for their belief have in recent times been examined with the consideration they deserve. The phenomenon of specific diversity is manifested to a similar degree by living things belonging to all the great groups, from the highest to the lowest, Vertebrates, Invertebrates, Protozoa, Vascular Plants, Algae, and Bacteria, all present diversities of such a kind that among them the existence of specific differences can on the whole be recognised with a similar degree of success and with very similar limitations. In all these groups there are many species quite definite and unmistakable, and others practically indefinite. The universal presence of specificity, as we may call it, similarly limited and characterised, is one of its most remarkable features. Not only is this specificity thus universally present among the different forms of life, but it manifests itself in respect of the most diverse characteristics which living things display. Species may thus be distinguished by peculiarities of form, of number, of geometrical arrangement, of chemical constitution and properties, of sexual differentiation, of development, and of many other properties. In any one or in several of these features together, species may be found distinguished from other species. It is also to be observed that the definiteness of these distinctions has no essential dependence on the nature of the characteristic which manifests them. It is for example sometimes said that colour-distinctions are of small systematic importance, but every systematist is familiar with examples (like that of the wild species of Gallus) in which colours though complex, show very little variation. On the other hand features of structure, sexual differentiation, and other attributes which by our standards are estimated as essential, may be declared to show much variation or little, not according to any principle which can be detected, but simply as the attention happens to be applied to one species or group of species, or to another. In many groups of animals and plants observers have hit upon characters which were for a time thought to be finally diagnostic of species. The Lepidoptera and Diptera for instance, have been re-classified according to their neuration. Through a considerable range of forms determinations may be easily made on these characters, but as is now well known, neuration is no more immune from variation than any other feature of organisation, and in some species great variability is the rule. Again it was once believed by some that the genitalia of the Lepidoptera provided a basis of final determination—with a similar sequel. In some groups, for example the Lycaenidae, or the Hesperidae, there are forms almost or quite indistinguishable on external examination, but a glance at the genitalia suffices to distinguish numerous species, while on the contrary among Pieridae a great range of species show scarcely any difference in these respects: and again in occasional species the genitalia show very considerable variations.

The proposition that animals and plants are on the whole divisible into definite and recognisable species is an approximation to the truth. Such a statement is readily defensible, whereas to assert the contrary would be palpably absurd. For example, a very competent authority lately wrote: "In the whole Lepidopterous fauna of England there is no species of really uncertain limits."[7] Others may be disposed to make certain reservations, but such exceptions would be so few as scarcely to impair the validity of the general statement. The declaration might be extended to other orders and other lands.

We know, of course, that the phenomenon of specific diversity is complicated by local differentiation: that, in general, forms which cannot disperse themselves freely exhibit a multitude of local races, and that of these some are obviously adaptative, and that a few even owe their peculiarity to direct environmental effects. Every systematist also is perfectly aware that in dealing with collections from little explored countries the occurrence of polymorphism or even of sporadic variation may make the practical business of distinguishing the species difficult and perhaps for the time impossible; still, conceding that a great part of the diversity is due to geographical differentiation, and that some is sporadic variation, our experience of our own floras and faunas encourages the belief that if we were thoroughly familiar with these exotic productions it would usually be possible to assign their specific limitations with an approach to certainty.

For apart from any question of the justice of these wider inferences, if we examine the phenomenon of specificity as it appears in those examples which are nearest to hand, surely we find signs in plenty that specific distinction is no mere consequence of Natural Selection. The strength of this proposition has lain mainly in the appeal to ignorance. Steadily with the growth of knowledge has its cogency diminished, and such a belief could only have been formulated at a time when the facts of variation were unknown.

In Darwin's time no serious attempt had been made to examine the manifestations of variability. A vast assemblage of miscellaneous facts could formerly be adduced as seemingly comparable illustrations of the phenomenon "Variation." Time has shown this mass of evidence to be capable of analysis. When first promulgated it produced the impression that variability was a phenomenon generally distributed amongst living things in such a way that the specific divisions must be arbitrary. When this variability is sorted out, and is seen to be in part a result of hybridisation, in part a consequence of the persistence of hybrids by parthenogenetic reproduction, a polymorphism due to the continued presence of individuals representing various combinations of Mendelian allelomorphs, partly also the transient effect of alteration in external circumstances, we see how cautious we must be in drawing inferences as to the indefiniteness of specific limits from a bare knowledge that intermediates exist. Conversely, from the accident of collocation or from a misleading resemblance in features we deem essential, forms genetically distinct are often confounded together, and thus the divergence of such forms in their other features, which we declare to be non-essential, passes as an example of variation. Lastly, and this is perhaps the most fertile of all the sources of confusion, the impression of the indefiniteness of species is created by the existence of numerous local forms, isolated geographically from each other, forms whose differences may be referable to any one of the categories I have enumerated.

The advance has been from many sides. Something has come from the work of systematists, something from cultural experiments, something from the direct study of variation as it appears in nature, but progress is especially due to experimental investigation of heredity. From all these lines of inquiry we get the same answer; that what the naturalists of fifty years ago regarded as variation is not one phenomenon but many, and that what they would have adduced as evidence against the definiteness of species may not in fact be capable of this construction at all.

If we may once more introduce a physical analogy, the distinctions with which the systematic naturalist is concerned in the study of living things are as multifarious as those by which chemists were confronted in the early days of their science. Diversities due to mechanical mixtures, to allotropy, to differences of temperature and pressure, or to degree of hydration, had all to be severally distinguished before the essential diversity due to variety of chemical constitution stood out clearly, and I surmise that not till a stricter analysis of the diversities of animals and plants has been made on a comprehensive scale, shall we be in a position to declare with any confidence whether there is or is not a natural and physiological distinction between species and variety.

As I have said above, it is in the cases nearest to hand that the problem may be most effectively studied. Comparison between forms from dissimilar situations contributes something; but it is by a close examination of the behaviour, especially the genetic behaviour, of familiar species when living in the presence of their nearest allies that the most direct light on the problem is to be obtained. I cannot understand the attitude of those who, contemplating such facts as this examination elicits, can complacently declare that specific difference is a mere question of degree. With the spread of evolutionary ideas to speak much of the fixity of species has become unfashionable, and yet how striking and inscrutable are the manifestations of that fixity!

Consider the group of species composing the agrestis section of the genus Veronica, namely Tournefortii, agrestis, and polita.

These three grow side by side in my garden, as they do in suitable situations over a vast area of the temperate regions. I have for years noticed them with some care and become familiar with their distinctions and resemblances. Never is there any real doubt as to the identity of any plant. The species show some variability, but I have never seen one which assumed any of the distinguishing features of the others. A glance at the fruits decides at once to which species a plant belongs. I find it impossible to believe that the fixity of these distinctions is directly dependent on their value as aids in the struggle for existence. The mode of existence of the three forms in so far as we can tell is closely similar. By whatever standard we reckon systematic affinity I suppose we shall agree that these species come very near indeed to each other. Bentham even takes the view that polita is a mere variety of agrestis.

Now in such cases as this it has been argued that the specific features of the several types have been separately developed in as many distinct localities, and that their present association is due to subsequent redistribution. Of these Veronicas indeed we know that one, Tournefortii (= Buxbaumii) is as a matter of fact a recent introduction from the east.[8] But this course of argument leads to still further difficulties. For if it is true that the peculiarities of the several species have been perfected and preserved on account of their survival-value to their possessors, it follows that there must be many ways of attaining the same result. But since sufficient adaptation may be ensured in so many ways, the disappearance of the common parent of these forms is difficult to understand. Obviously it must have been a plant very similar in general construction to its modern representatives. Like them it must have been an annual weed, with an organisation conformable to that mode of life. Why then, after having been duly perfected for that existence should it have been entirely superseded in favour of a number of other distinct contrivances for doing the same thing, and—if a gradual transition be predicated—not only by them, but by each intermediate stage between them and the original progenitor? Surely the obvious inference from such facts is that the burden cast upon the theory of gradual selection is far greater than it can bear; that adaptation is not in practice a very close fit, and that the distinctions between these several species of Veronica have not arisen on account of their survival-value but rather because none of their diversities was so damaging as to lead to the extermination of its possessor. When we see these various Veronicas each rigidly reproducing its parental type, all comfortably surviving in competition with each other, are we not forced to the conclusion that tolerance has as much to do with the diversity of species as the stringency of Selection? Certainly these species owe their continued existence to the fact that they are each good enough to live, but how shall we refer the distinctions between them directly or indirectly to the determination of Natural Selection?

The control of Selection is loose while the conformity to specific distinction is often very strict and precise, and no less so even when several closely related species co-exist in the same area and in the same circumstances.

The theory of Selection fails at exactly the point where it was devised to help: Specific distinction.

Let us examine a somewhat different set of facts in the case of another pair of nearly allied species Lychnis diurna and vespertina. The two plants have much in common. Both are dioecious perennials, with somewhat similar flowers, the one crimson, the other white. Each however has its peculiarities which are discernible in almost any part of its structure, whether flower, leaf, fruit or seed, distinctions which would enable a person thoroughly familiar with the plants to determine at once from which species even a small piece had been taken. There is so much resemblance however as readily to support the surmise that the two were mere varieties of one species. Bentham, following Linnaeus, in fact actually makes this suggestion, with what propriety we will afterwards consider. Now this case is typical of many. The two forms have a wide distribution, occurring sometimes separately, sometimes in juxtaposition. L. diurna is a plant of hedgerows and sheltered situations. L. vespertina is common in fields and open spaces, where diurna is hardly ever found; but not rarely vespertina occurs in association with diurna in the places which that plant frequents. In this case I do not doubt that we have to do with organisms of somewhat different aptitudes. That L. vespertina has powers which diurna has not is shown very clearly by the fact that diurna is sometimes entirely absent from areas where vespertina can abound.[9] But in order to understand the true genetic relations of the two plants to each other it is necessary to observe their behaviour when they meet as they not unfrequently do. If the Lychnis population of such a locality be examined it will be found to consist of many undoubted and unmodified diurna, a number—sometimes few, sometimes many—of similarly unmodified vespertina, and an uncertain but usually rather small proportion of plants obviously hybrids between the two. How is it possible to reconcile these facts with the view that specific distinction has no natural basis apart from environmental exigency?

Darwinian orthodoxy suggests that by a gradual process of Natural Selection either one of these two types was evolved from the other, or both from a third type. I cannot imagine that anyone familiar with the facts would propose the first hypothesis in the case of Lychnis, nor can I conceive of any process, whether gradual or sudden, by which diurna could have come out of vespertina, or vespertina out of diurna. Both however may no doubt have been derived from some original third type. It is conceivable that Lychnis macrocarpa of Boissier, a native of Southern Spain and Morocco, may be this original form. This species is said to combine a white flower (like that of L. vespertina), with capsule-teeth rolled back (like those of diurna).[10] But whatever the common progenitor may have been, if we are to believe that these two species have been evolved from it by a gradual process of Natural Selection based on adaptation, enormous assumptions must be made regarding the special fitness of these two forms and the special unfitness of the common parent, and these assumptions must be specially invoked and repeated for each several feature of structure or habits distinguishing the three forms.

Why, if the common parent was strong enough to live to give rise to these two species, is it either altogether lost now, or at least absent from the whole of Northern Europe? Its two putative descendants, though so distinct from each other, are, as we have seen, able often to occupy the same ground. If they were gradually derived from a common progenitor—necessarily very like themselves—can we believe that this original form should always, in all the diversities of soil and situation which they inhabit, be unable to exist? Some one may fancy that the hybrids which are found in the situations occupied by both forms are this original parental species. But nothing can be more certain than that these plants are simply heterozygous combinations made by the union of gametes bearing the characters of diurna and vespertina.[11] For they may be reproduced exactly in F₁ or in later generations of that cross when it is artificially made; when bred from their families exhibit palpable phenomena of segregation more or less complex; and usually, if perhaps not always, they are partially sterile.[12] In a locality on the Norfolk coast that I know well, there is a strip of rough ground chiefly sand-bank, which runs along the shore. This ground is full of vespertina. Not a hundred yards inland is a lane containing diurna, and among the vespertina on the sand-bank are always some of the hybrid form, doubtless the result of fertilisation from the neighbouring diurna population. Seed saved from these hybrids gave vespertina and hybrids again, having obviously been fertilised by other vespertina or by other hybrids, and I have no doubt that such hybrid plants if fertilised by diurna would have shown some diurna offspring. The absence of diurna in such localities may fairly be construed as an indication that diurna is there at a real disadvantage in the competition for life.

But if, admitting this, we proceed to consider how the special aptitude of vespertina is constituted, or what it is that puts diurna at a disadvantage, we find ourselves quite unable to show the slightest connexion between the success of one or the failure of the other on the one hand, and the specific characteristics which distinguish the two forms on the other. The orthodox Selectionist would, as usual, appeal to ignorance. We ask what can vespertina gain by its white flowers, its more lanceolate leaves, its grey seeds, its almost erect capsule-teeth, its longer fruits, which diurna loses by reason of its red flowers, more ovate leaves, dark seeds, capsule-teeth rolled back, and shorter fruits? We are told that each of these things may affect the viability of their possessors. We cannot assert that this is untrue, but we should like to have evidence that it is true. The same problem confronts us in thousands upon thousands of examples, and as time goes on we begin to feel that speculative appeals to ignorance, though dialectically admissible, provide an insufficient basis for a proposition which, if granted, is to become the foundation of a vast scheme of positive construction.

One thing must be abundantly clear to all, that to treat two forms so profoundly different as one, because intermediates of unknown nature can be shown to exist between them, is a mere shirking of the difficulties, and this course indeed creates artificial obstacles in the way of those who are seeking to discover the origin of organic diversity.

In the enthusiasm with which evolutionary ideas were received the specificity of living things was almost forgotten. The exactitude with which the members of a species so often conform in the diagnostic, specific features passed out of account; and the scientific world by dwelling with a constant emphasis on the fact of variability, persuaded itself readily that species had after all been a mere figment of the human mind. Without presuming to declare what future research only can reveal, I anticipate that, when variation has been properly examined and the several kinds of variability have been successfully distinguished according to their respective natures, the result will render the natural definiteness of species increasingly apparent. Formerly in such a case as that of the two Lychnis species, the series of "intermediates" was taken to be a palpable proof that vespertina "graded" to diurna. It is this fact, doubtless, upon which Bentham would have relied in suggesting that both may be one species.[13] Genetic tests, though as yet imperfectly applied, make it almost certain that these inter-grading forms are not in any true sense variations from either species in the direction of the other, but combinations of elements derived from both.

The points in which very closely allied species are distinguished from each other may be found in the most diverse features of their organisation. Sometimes specific difference is to be seen in a character which we can believe to be important in the struggle, but at least as often it is some little detail that we cannot but regard as trivial which suffices to differentiate the two species. Even when the diagnostic point is of such a nature that we can imagine it to make a serious difference in the economy we are absolutely at a loss to suggest why this feature should be a necessity to species A and unnecessary to species B its nearest ally. The house sparrow (Passer domesticus) is in general structure very like the tree sparrow (P. montanus). They differ in small points of colour. For instance montanus has a black patch on the cheek which is absent in domesticus. The presence in the one species and the absence in the other are equally definite, and in both cases we are equally unable to suggest any consideration of utility in relation to these features. The two species are distinguished also by a characteristic that may well be supposed to be of great significance. In domesticus the two sexes are strongly differentiated, the cock being more ornate than the hen. On the other hand the two sexes in montanus are alike, and, if we take a standard from domesticus, we may fairly say that in montanus the hen has the colouration of the male. It is not unreasonable to suppose that such a distinction may betoken some great difference in physiological economy, but the economical significance of this perhaps important distinction is just as unaccountable as that of the seemingly trivial but equally diagnostic colour-point.

I have spoken of the fixed characteristics of the two species. If we turn to a very different feature, their respective liability to albinistic variation, we find ourselves in precisely similar difficulty. Passer domesticus is a species in which individuals more or less pied occur with especial frequency, but in P. montanus such variation is extremely rare if it occurs at all. The writer of the section on Birds in the Royal Natural History (III., 1894-5, p. 393) calls attention to this fact and remarks that in that species he knows no such instance.

The two species therefore, apart from any differences that we can suppose to be related to their respective habits, are characterised by small fixed distinctions in colour-markings, by a striking difference in secondary sexual characters, and by a difference in variability. In all these respects we can form no surmise as to any economic reason why the one species should be differentiated in the one way and the other in the other way, and I believe it is mere self-deception which suggests the hope that with fuller knowledge reasons of this nature would be discovered.

The two common British wasps, Vespa vulgaris and Vespa germanica, are another pair of species closely allied although sharply distinguished, which suggest similar reflexions. Both usually make subterranean nests but of somewhat different materials. V. vulgaris uses rotten wood from which the nest derives a characteristic yellow colour, while V. germanica scrapes off the weathered surfaces of palings and other exposed timber, material which is converted into the grey walls of the nest. The stalk by which the nest is suspended (usually to a root) in the case of germanica passes freely through a hole in the external envelope, but vulgaris unites this external wall solidly to the stalk. In bodily appearance and structure the two species are so much alike that they have often been confounded even by naturalists, and to the untrained observer they are quite indistinguishable. There are nevertheless small points of difference which almost though not quite always suffice to distinguish the two forms. For example the yellow part of the sinus of the eyes is emarginate in vulgaris but not emarginate in germanica. V. vulgaris often has black spots on the tibiae while in germanica the tibiae are usually plain yellow. In both species there is a horizontal yellow stripe on the thorax, but whereas in vulgaris this is a plain narrow stripe, it is in germanica enlarged downwards in the middle. These and other apparently trivial details of colouration, though not absolutely constant, are yet so nearly constant that irregularities in these respects are quite exceptional. Lastly the genitalia of the males, though not very different, present small structural points of distinction which are enough to distinguish the two species at a glance.[14]

In considering the meaning of the distinctions between these two wasps we meet the old problem illustrated by the Sparrows. The two species have somewhat different habits of life and we should readily expect to find differences of bodily organisation corresponding with the differences of habits. But is that what we do find? Surely not. To suppose that there is a correspondence between the little points of colour and structure which we see and the respective modes of life of the two species is perfectly gratuitous. We have no inkling of the nature of such a correspondence, how it can be constituted, or in what it may consist.

Is it not time to abandon these fanciful expectations which are never realised? Everywhere both among animals and plants does the problem of specific difference reiterate itself in the same form. In view of such facts as I have related and might indefinitely multiply, the fixity of specific characters cannot readily be held to be a measure of their economic importance to their possessors. The incidence of specific fixity is arbitrary and capricious, sometimes lighting on a feature or a property which can be supposed to matter much, but as often is it attached to the most trifling of superficial peculiarities.

The incidence of variability is no less paradoxical, and without investigation of the particular case no one can say what will be found to show much or little variability. The very characteristic which in one species may exhibit extreme variability may in an allied species show extreme constancy. Illustrations will occur to any naturalist, but nowhere is this truth more strikingly presented than in the British Noctuid Moths. Many are so variable that, in the common phrase, "scarcely two can be found alike," while others show comparatively slight variation. It need scarcely be remarked that, in the instances I have in mind, the evidence of great variability is in no way due to the abundance with which the particular species occurs, for common species may show constancy, and less abundant species may show great variability. The polymorphism seems to be now at least a general property of the variable species, as the fixity is a property of the fixed species. In illustration I may refer to the following examples.

Dianthoecia capsincola is a common and widely distributed moth which feeds on Lychnis. It shows little variation. Dianthoecia carpophaga is another species which feeds chiefly on Silene. Its habits are very similar to those of capsincola. Like that species it has a wide geographical range and is abundant in its localities, but in contrast to the fixity of capsincola, carpophaga exhibits a complex series of varieties. Agrotis suffusa (= ypsilon) is a moth widely spread through the southern half of England. It is very constant in colour and markings. Agrotis segetum and tritici are excessively variable both in ground colour and markings, being found in an immense profusion of dissimilar forms throughout their distribution. Of these and several other species of Agrotis there are many named varieties, some of which have by various writers been regarded as specifically distinct. Of the genus Noctua many species (e. g. festiva) show a similar polymorphism, but N. triangulum, though showing some variation in certain respects, is usually very constant to its type, and the same is true of N. umbrosa.

In several species of Taeniocampa, especially instabilis, the multiplicity of forms is extreme, while cruda (= pulverulenta) is a comparatively constant species. The genus Plusia contains a number of constant species, but in Plusia interrogationis we meet the fact that the central silvery mark undergoes endless variation. "Truly no two are alike," says Mr. Tutt, "and to look down a long series of interrogationis is something like looking at a series of Chinese characters." In contrast to this we have the fact that in Plusia gamma the very similar silvery mark is by no means variable.

I have taken this series of cases from the Noctuid moths, but it would be as easy to illustrate the same proposition from the Geometridae or the Micro-Lepidoptera.[15] I have a longseries of Peronea cristana, for example, which was given to me by Mr. W. H. B. Fletcher, of Bognor. All were beaten out of the same hedge, and their polymorphism is such that no one unaccustomed to such examples could suppose that they belonged to a single species. Another common form, P. schalleriana, which lives in similar circumstances, exhibits comparatively slight variability.

It should be expressly noted that the variation of which I am speaking is a genuine polymorphism. Several of the species enumerated exhibit also geographical variation, possessing definite and often strikingly distinct races peculiar to certain localities; but apart from the existence of such local differentiation, stands out the fact upon which I would lay stress, that some species are excessively variable while others are by comparison constant, in circumstances that we may fairly regard as comparable.

This fact is difficult to reconcile with the conventional view that specific type is directly determined by Natural Selection and that the precision with which a species conforms to its pattern is an indication of the closeness of that control. Anyone familiar with the characteristics of Moths will agree that the Noctuids, Geometrids and Tortricids are creatures whose existence depends in some degree on the success with which they can escape detection by their enemies in the imaginal state. We are therefore not surprised to find that some species of these orders exhibit definite geographical variation in conformity with the character of the ground, which may reasonably be supposed to aid in their protection. If this were all, there would be nothing to cause surprise. We might even be disposed to allow that variability might contribute to the perpetuation of animals so situated, on the principle that among a variety of surroundings some would probably be in harmony with the objects on which they rest. But we cannot admit the plausibility of an argument which demands on the one hand that the extreme precision with which species A adheres in the minutest details of its colour and pattern to a certain type shall be ascribed to the protective fitness of those details, and on the other hand that the abundant variability of species B shall be ascribed to the same determination. If it is absolutely necessary for A to conform to one type how comes it that B may range through some twenty distinct forms, any two of which differ more from each other than the regular species of many other genera? The only reply I can conceive is a suggestion that there may be some circumstance which differentiates the various classes of cases, that the exigencies of the fixed species may be different from those of the variable. Those who make such appeals to ignorance do not always perhaps realise whither this course of reasoning may lead. If admissible here the same argument would lead us to suggest that because albino moles have for an indefinite period occurred on a certain land near Bath there may be something in the soil or in the conditions of life near Bath which requires a proportion of albinos in its mole population. Or again, because the butterfly Thais rumina in one locality, Digne in the south of France, has a percentage of individuals of the variety Honoratii (with certain normally yellow spots on the hind wing coloured bright red) and nowhere else throughout its distribution, that therefore we may suggest that there is some difference in the condition of life at Digne which makes the continuance of Honoratii there possible and beneficial.

A polymorphism offering a parallel to that of the variable moths is afforded by the breeding plumage of the Ruff, the male of Machetes pugnax. The variety of plumage which these cocks exhibit is such that the statement that no two can be found alike is only a venial exaggeration. Newton remarks[16] "that all this wonderful 'show' is the consequence of the polygamous habit of the Ruff can scarcely be doubtful"; but even if it be conceded that the great external differentiation of the cocks may be a result of sexual selection, the problem of their polymorphism remains unsolved, for, as we are well aware, polygamy is not usually associated with polymorphism of the male. The Black Cock (Tetrao tetrix), for example, is as polygamous as the Ruff, but in that and countless other cases, both sexes are constant to one type of plumage.

When we thus compare the polymorphism of one species with the fixity of another, and attempt to determine the causes which have led to these extraordinary contrasts, two distinct lines of argument are open to us. We may ascribe the difference either to causes external to the organisms, primarily, that is to say, to a difference in the exigencies of Adaptation under Natural Selection; or on the other hand we may conceive the difference as due to innate distinctions in the chemical and physiological constitutions of the fixed and the variable respectively. There is truth undoubtedly in both conceptions. If the mole were physiologically incapable of producing an albino that variety would not have come into being, and if the albino were totally incapable of getting its living it would not be able to hold its own. Were Plotheia frontalis constructed on a chemical plan which admitted of no variation, the countless varieties would not have been produced; and if one of its varieties had an overwhelming success out of all proportion to that of the rest, then the species would soon become monomorphic again. We cannot declare that Natural Selection has no part in the determination of fixity or variability; nevertheless looking at the whole mass of fact which a study of the incidence of variation provides, I incline to the view that the variability of polymorphic forms should be regarded rather as a thing tolerated than as an element contributing directly to their chances of life; and on the other hand that the fixity of the monomorphic forms should be looked upon not so much as a proof that Natural Selection controls them with a greater stringency, but rather as evidence of a natural and intrinsic stability of chemical constitution.

Compare the condition of a variable form like the male Ruff (or in a less degree the Red Grouse in both its sexes) with that of the common Pheasant which is comparatively constant. In the Pheasant no doubt variations do occur as in other wild birds, but apart from the effects of mongrelisation the species is unquestionably uniform. Could it seriously be proposed that we should regard the constancy of the pheasant's plumage in this country as depending on the special fitness of that type of colouration? Even if the pheasant be not an alien in Western Europe, it has certainly been protected for centuries, and for a considerable period has existed in a state of semi-domestication. Such conditions should give good opportunity for polymorphism to be produced. In some coverts various aberrations do of course occur and persist, yet there is nothing indicative of a general relaxation of the fixity of the specific type, and the pheasant remains substantially a fixed species.[17] The common pheasant (Phasianus colchicus) even shows little of that disposition to form local races which appears in the species of Further India. Are we not then on safer ground in regarding the fixity of our species as a property inherent in its own nature and constitution? Just as in ages of domestication no rose has ever given off a blue variety so has the pheasant never broken out into the polymorphism of the Ruff.

As soon as it is realised how largely the phenomena of variation and stability must be an index of the internal constitution of organisms, and not mere consequences of their relations to the outer world, such phenomena acquire a new and more profound significance.