Conclusion
If we can comprehend and interpret on some such lines as these the form and mode of growth of the foraminiferal shell, we may also begin to understand two striking features of the group, namely, on the one hand the large number of diverse types or families which exist and the large number of species and varieties within each, and on the other the persistence of forms which in many cases seem to have undergone little change or none at all from the Cretaceous or even from earlier periods to the present day. In few other groups, perhaps only among the Radiolaria, do we seem to possess so nearly complete a picture of all possible transitions between form and form, and of the whole branching system of the evolutionary tree: as though little or nothing of it had ever perished, and the whole web of life, past and present, were as complete as ever. It leads one to imagine that these shells have grown according to laws so simple, so much in harmony with their material, with their environment, and with all the forces internal and external to which they are exposed, that none is better than another and none fitter or less fit to survive. It invites one also to contemplate the possibility of the lines of possible variation being here so narrow and determinate that identical forms may have come independently into being again and again.
While we can trace in the most complete and beautiful manner the passage of one form into another among these little shells, and ascribe them all at last (if we please) to a series which starts with the simple sphere of Orbulina or with the amoeboid body of Astrorhiza, the question stares us in the face whether this be an “evolution” which we have any right to correlate with historic time. The mathematician can trace one conic section into another, and “evolve” for example, through innumerable graded ellipses, the circle from the straight line: which tracing of continuous steps is a true “evolution,” though time has no part therein. It was after this fashion that Hegel, and for that matter Aristotle himself, was an evolutionist—to whom evolution was {608} a mental concept, involving order and continuity in thought, but not an actual sequence of events in time. Such a conception of evolution is not easy for the modern biologist to grasp, and harder still to appreciate. And so it is that even those who, like Dreyer and like Rhumbler, study the foraminiferal shell as a physical system, who recognise that its whole plan and mode of growth is closely akin to the phenomena exhibited by fluid drops under particular conditions, and who explain the conformation of the shell by help of the same physical principles and mathematical laws—yet all the while abate no jot or tittle of the ordinary postulates of modern biology, nor doubt the validity and universal applicability of the concepts of Darwinian evolution. For these writers the biogenetisches Grundgesetz remains impregnable. The Foraminifera remain for them a great family tree, whose actual pedigree is traceable to the remotest ages; in which historical evolution has coincided with progressive change; and in which structural fitness for a particular function (or functions) has exercised its selective action and ensured “the survival of the fittest.” By successive stages of historic evolution we are supposed to pass from the irregular Astrorhiza to a Rhabdammina with its more concentrated disc; to the forms of the same genus which consist of but a single tube with central chamber; to those where this chamber is more and more distinctly segmented; so to the typical many-chambered Nodosariae; and from these, by another definite advance and later evolution to the spiral Trochamminae. After this fashion, throughout the whole varied series of the Foraminifera, Dreyer and Rhumbler (following Neumayr) recognise so many successions of related forms, one passing into another, and standing towards it in a definite relationship of ancestry or descent. Each evolution of form, from simpler to more complex, is deemed to have been attended by an advantage to the organism, an enhancement of its chances of survival or perpetuation; hence the historically older forms are, on the whole, structurally the simpler; or conversely the simpler forms, such as the simple sphere, were the first to come into being in primeval seas; and finally, the gradual development and increasing {609} complication of the individual within its own lifetime is held to be at least a partial recapitulation of the unknown history of its race and dynasty.
We encounter many difficulties when we try to extend such concepts as these to the Foraminifera. We are led for instance to assert, as Rhumbler does, that the increasing complexity of the shell, and of the manner in which one chamber is fitted on another, makes for advantage; and the particular advantage on which Rhumbler rests his argument is strength. Increase of strength, die Festigkeitssteigerung, is according to him the guiding principle in foraminiferal evolution, and marks the historic stages of their development in geologic time. But in days gone by I used to see the beach of a little Connemara bay bestrewn with millions upon millions of foraminiferal shells, simple Lagenae, less simple Nodosariae, more complex Rotaliae: all drifted by wave and gentle current from their sea-cradle to their sandy grave: all lying bleached and dead: one more delicate than another, but all (or vast multitudes of them) perfect and unbroken. And so I am not inclined to believe that niceties of form affect the case very much: nor in general that foraminiferal life involves a struggle for existence wherein breakage is a constant danger to be averted, and increased strength an advantage to be ensured.
In the course of the same argument Rhumbler remarks that Foraminifera are absent from the coarse sands and gravels, as Williamson indeed had observed many years ago: so averting, or {610} at least escaping, the dangers of concussion. But this is after all a very simple matter of mechanical analysis. The coarseness or fineness of the sediment on the sea-bottom is a measure of the current: where the current is strong the larger stones are washed clean, where there is perfect stillness the finest mud settles down; and the light, fragile shells of the Foraminifera find their appropriate place, like every other graded sediment, in this spontaneous order of lixiviation.
The theorem of Organic Evolution is one thing; the problem of deciphering the lines of evolution, the order of phylogeny, the degrees of relationship and consanguinity, is quite another. Among the higher organisms we arrive at conclusions regarding these things by weighing much circumstantial evidence, by dealing with the resultant of many variations, and by considering the probability or improbability of many coincidences of cause and effect; but even then our conclusions are at best uncertain, our judgments are continually open to revision and subject to appeal, and all the proof and confirmation we can ever have is that which comes from the direct, but fragmentary evidence of palaeontology.
But in so far as forms can be shewn to depend on the play of physical forces, and the variations of form to be directly due to simple quantitative variations in these, just so far are we thrown back on our guard before the biological conception of consanguinity, and compelled to revise the vague canons which connect classification with phylogeny.
The physicist explains in terms of the properties of matter, and classifies according to a mathematical analysis, all the drops and forms of drops and associations of drops, all the kinds of froth and foam, which he may discover among inanimate things; and his task ends there. But when such forms, such conformations and configurations, occur among living things, then at once the biologist introduces his concepts of heredity, of historical evolution, of succession in time, of recapitulation of remote ancestry in individual growth, of common origin (unless contradicted by direct evidence) of similar forms remotely separated by geographic space or geologic time, of fitness for a function, of {611} adaptation to an environment, of higher and lower, of “better” and “worse.” This is the fundamental difference between the “explanations” of the physicist and those of the biologist.
In the order of physical and mathematical complexity there is no question of the sequence of historic time. The forces that bring about the sphere, the cylinder or the ellipsoid are the same yesterday and to-morrow. A snow-crystal is the same to-day as when the first snows fell. The physical forces which mould the forms of Orbulina, of Astrorhiza, of Lagena or of Nodosaria to-day were still the same, and for aught we have reason to believe the physical conditions under which they worked were not appreciably different, in that yesterday which we call the Cretaceous epoch; or, for aught we know, throughout all that duration of time which is marked, but not measured, by the geological record.
In a word, the minuteness of our organism brings its conformation as a whole within the range of the molecular forces; the laws of its growth and form appear to lie on simple lines; what Bergson calls the “ideal kinship” is plain and certain, but the “material affiliation” is problematic and obscure; and, in the end and upshot, it seems to me by no means certain that the biologist’s usual mode of reasoning is appropriate to the case, or that the concept of continuous historical evolution must necessarily, or may safely and legitimately, be employed.