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PHYLOGENY] old quarrel between evolution and epigenesis. The most striking general change has been against seeing in the facts of ontogeny any direct evidence as to phylogeny. The general proposition as to a parallelism between individual and ancestral development is no doubt indisputable, but extended knowledge of the very different ontogenetic histories of closely allied forms has led us to a much fuller conception of the mode in which stages in embryonic and larval history have been modified in relation to their surroundings, and to a consequent reluctance to attach detailed importance to the embryological argument for evolution.

The vast bulk of botanical and zoological work on living and extinct forms published during the last quarter of the 19th century increased almost beyond all expectation the evidence for the fact of evolution. The discovery of a single fossil creature in a geological stratum of a wrong period,

the detection of a single anatomical or physiological fact irreconcilable with origin by descent with modification, would have been destructive of the theory and would have made the reputation of the observer. But in the prodigious number of supporting discoveries that have been made no single negative factor has appeared, and the evolution from their predecessors of the forms of life existing now or at any other period must be taken as proved. It is necessary to notice, however, that although the general course of the stream of life is certain, there is not the same certainty as to the actual individual pedigrees of the existing forms. In the attempts to place existing creatures in approximately phylogenetic order, a striking change, due to a more logical consideration of the process of evolution, has become established and is already resolving many of the earlier difficulties and banishing from the more recent tables the numerous hypothetical intermediate forms so familiar in the older phylogenetic trees. The older method was to attempt the comparison between the highest member of a lower group and the lowest member of a higher group—to suppose, for example, that the gorilla and the chimpanzee, the highest members of the apes, were the existing representatives of the ancestors of man and to compare these forms with the lowest members of the human race. Such a comparison is necessarily illogical, as the existing apes are separated from the common ancestor by at least as large a number of generations as separate it from any of the forms of existing man. In the natural process of growth, the gap must necessarily be wider between the summits of the twigs than lower down, and, instead of imagining “missing links,” it is necessary to trace each separate branch as low down as possible, and to institute the comparisons between the lowest points that can be reached. The method is simply the logical result of the fact that every existing form of life stands at the summit of a long branch of the whole tree of life. A due consideration of it leads to the curious paradox that if any two animals be compared, the zoologically lower will be separated from the common ancestor by a larger number of generations, since, on the average, sexual maturity is reached more quickly by the lower form. Naturally very many other factors have to be considered, but this alone is a sufficient reason to restrain attempts to place existing forms in linear phylogenetic series. In embryology the method finds its expression in the limitation of comparisons to the corresponding stages of low and high forms and the exclusion of the comparisons between the adult stages of low forms and the embryonic stages of higher forms. Another expression of the same method, due to Cope, and specially valuable to the taxonomist, is that when the relationship between orders is being considered, characters of subordinal rank must be neglected. It must not be supposed that earlier writers all neglected this method, or still less that all writers now employ it, but merely that formerly it was frequently overlooked by the best writers, and now is neglected only by the worst. The result is, on the one hand, a clearing away of much fantastic phylogeny, on the other, an enormous reduction of the supposed gaps between groups.

There has been a renewed activity in the study of existing forms from the point of view of obtaining evidence as to the nature and origin of species. Comparative anatomists have been learning to refrain from basing the diagnosis of a species, or the description of the condition of an organ, on the evidence of a

single specimen. Naturalists who deal specially with museum collections have been compelled, it is true, for other reasons to attach an increasing importance to what is called the type specimen, but they find that this insistence on the individual, although invaluable from the point of view of recording species, is unsatisfactory from the point of view of scientific zoology; and propositions for the amelioration of this condition of affairs range from a refusal of Linnaean nomenclature in such cases, to the institution of a division between master species for such species as have been properly revised by the comparative morphologist, and provisional species for such species as have been provisionally registered by those working at collections. Those who work with living forms of which it is possible to obtain a large number of specimens, and those who make revisions of the provisional species of palaeontologists, are slowly coming to some such conception as that a species is the abstract central point around which a group of variations oscillate, and that the peripheral oscillations of one species may even overlap those of an allied species. It is plain that we have moved far from the connotation and denotation of the word species at the time when Darwin began to discuss the origin of species, and that the movement, on the one hand, tends to simplify the problem philosophically, and, on the other, to make it difficult for the amateur theorist.

The conception of evolution is being applied more rigidly to the comparative anatomy of organs and systems of organs. When a series of the modifications of an anatomical structure has been sufficiently examined, it is frequently possible to decide that one particular condition is primitive, ancestral or central, and that the other conditions have been derived from it. Such a condition has been termed, with regard to the group of animals or plants the organs of which are being studied, archecentric. The possession of the character in the archecentric condition in (say) two of the members of the group does not indicate that these two members are more nearly related to one another than they are to other members of the group; the archecentric condition is part of the common heritage of all the members of the group, and may be retained by any. On the other hand, when the ancestral condition is modified, it may be regarded as having moved outwards along some radius from the archecentric condition. Such modified conditions have been termed apocentric. It is obvious that the mere apocentricity of a character can be no guide to the affinities of its possessor. It is necessary to determine if the modification be a simple change that might have occurred in independent cases, in fact if it be a multiradial apocentricity, or if it involved intricate and precisely combined anatomical changes that we could not expect to occur twice independently; that is to say, if it be a uniradial apocentricity. Multiradial apocentricities lie at the root of many of the phenomena that have been grouped under the designation convergence. Especially in the case of manifest adaptations, organs possessed by creatures far apart genealogically may be moulded into conditions that are extremely alike. Sir E. Ray Lankester’s term, homoplasy, has passed into currency as designating such cases where different genetic material has been pressed by similar conditions into similar moulds. These may be called heterogeneous homoplasies, but it is necessary to recognize the existence of homogeneous homoplasies, here called multiradial apocentricities. A complex apocentric modification of a kind which we cannot imagine to have been repeated independently, and which is to be designated as uniradial, frequently forms a new centre around which new diverging modifications are produced. With reference to any particular group of forms such a new centre of modification may be termed a metacentre, and it is plain that the archecentre of the whole group is a metacentre of the larger group of which the group under consideration is a branch. Thus, for instance, the archecentric condition of any Avian structure is a metacentre of the Sauropsidan stem. A form of apocentricity extremely common and often perplexing may be termed pseudocentric; in such a condition there is an apparent simplicity that