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Art. I.— On the Origination and Distribution of Species:Introductory Essay to the Flora of Tasmania; by Dr. JOSEPH D. Hooker*

§ 1. Preliminary Remarks.

The Island of Tasmania does not contain a vegetation peculiar to itself, nor constitute an independent botanical region. Ita plants are, with comparatively few exceptions, natives of extratropical Australia; and I have consequently found it necessary to study the vegetation of a great part of that vast continent, in order to determine satisfactorily the nature, distribution, and

* To the Editors of the American Journal of Science, &c.:—The sheets of this Introductory Essay, having been obligingly communicated to me in advance of the publication of the concluding part of the Flora of Tasmania, to which it belongs, I asked and have received the distinguished author's permission to reprint them, or a considerable portion of them, in jour Journal, and now offer them for that purpose. This is in order that we may have before us, at the earliest date, an essay which cannot fail to attract the immediate and profound attention of scientific men; but which, if confined to the pages of the Flora of Tasmania, would be seen by very few American readers. To those who have intelligently observed the course of scientific investigation, and the tendency of speculation, it has for some time been manifest that a re statement of the Lamarkian hypothesis is at hand. We have this, in an improved and truly scientific form, in the theories which, recently propounded by Mr. Darwin, followed by Mr. Wallace, are here so ably and altogether independently maintained. When these views are fully laid before them, the naturalists of this country will be able to take part in the interesting discussion which they will not fail to call forth.

To save room, a few paragraphs are omitted which do not directly bear upon the subject in hand. A. O.

SECOND SERIES, Vox. XXIX, No. 85 JAN., 1860.

affinities of the Tasmanian Flora. From the study of certain extratropical genera and species in their relation to those of Tasmania, I have been led to the far more comprehensive undertaking of arranging and classifying all the Australian plants accessible to me. This I commenced in the hope of being able thereby to extend our knowledge of the affinities of its Flora, and, if possible, to throw light on a very abstruse subject, viz. the origin of its vegetation, and the sources or causes of its peculiarity. This again has induced me to proceed with the inquiry into the oxigin and distribution of existing species; and, as I have already treated of these subjects in the Introduction to the New Zealand Flora, I now embrace the opportunity afforded me by a similar Introduction to the Tasmanian Flora, of revising the opinions I then entertained, and of again investigating the whole subject of the creation of species by variation, with the aid of the experience derived from my subsequent studies of the Floras of India and Australia in relation to one another and to those of neighboring countries, and of the recently published hypotheses of Mr. Darwin and Mr. Wallace. * * *

In the Introductory Essay to the New Zealand Flora, I advanced certain general propositions as to the origin of species, which I refrained from endorsing as articles of my own creed: amongst others was the still prevalent doctrine that these are, in the ordinary acceptation of the term, created as such, and are immutable. In the present Essay I shall advance the opposite hypothesis, that species are derivative and mutable; and this chiefly because, whatever opinions a naturalist may have adopted with regard to the origin and variation of species, every candid mind must admit that the facts and arguments upon which he has grounded his convictions require revision since the recent publication by the Linnean Society of the ingenious and original reasonings and theories of Mr. Darwin and Mr. Wallace.

Further, there must be many who, like myself, having hitherto refrained from expressing any positive opinion, now, after a careful consideration of these naturalists' theories, find the aspect of the question materially changed, and themselves freer to adopt such a theory as may best harmonize with the facts adduced by their own experience.

The Natural History of Australia seemed to me to be especially suited to test such a theory, on account of the comparative uniformity of its physical features being accompanied with a great variety in its Flora; of the differences in the vegetation of its several parts; and of the peculiarity both of its Fauna and Flora, as compared with those of other countries. I accordingly prepared a classified catalogue of all the Australian species in the Herbarium, with their ranges in longitude, latitude, and elevation, as far as I could ascertain them, and added what further information I could obtain from books. At the same time I made a careful study of the affinities and distribution of all the Tasmanian species, and of all those Australian ones which I believed to be found in other countries. I also determined as accurately as I could the genera of the remainder, and especially of those belonging to genera which are found in other countries, and I distinguished the species from one another in those genera which had not been previously arranged. In this manner I have brought together evidence of nearly 8000 flowering plants having been collected or observed in Australia, of which I have seen and catalogued upwards of 7000. About two-thirds of these are ascertained specifically with tolerable accuracy, and the remainder are distinguished from one another, and referred to genera with less certainty, being either undescribed, or described under several names, whilst some are members of such variable groups that I was left in doubt how to dispose of them.

To many who occupy themselves with smaller and better worked botanical districts, such results as may be deduced from the skeleton Flora I have compiled for Australia may seem too crude and imperfect to form data from which to determine its relations. But it is not from a consideration of specific details that such problems as those of the relations of Floras and the origin and distribution of organic forms will ever be solved, though we must eventually look to these details for proofs of the solutions we propose. The limits of the majority of species are so undefinable that few naturalists are agreed upon them;* to a great extent they are matters of opinion, even amongst those persons who believe that species are original and immutable creations; and as our knowledge of the forms and allies of each increases, so do these differences of opinions; the progress of systematic science being, in short, obviously unfavorable tothe view that most species are limitable by descriptions or characters, unless large allowances are made for variation. On the other hand, when dealing with genera, or other combinations of species, all that is required is that these be classified in natural groups; and that such groups are true exponents of affinitiessettled by Nature is abundantly capable of demonstration. It is to an investigation of the extent, relations, and proportions of these natural combinations of species, then, that we must look for the means of obtaining and expressing the features of a Flora; and if in this instance the exotic species are well ascertained, it matters little whether or not the endemic arc in all cases accurately distinguished from one another. Further, in a Flora so large as that of Australia, if the species are limited and

* The most conspicuous evidence of this lies in the fact, that the number of known species of flowering plants is by some assumed to be under 80,000, and by others over 150,000.

estimated by one mind and eye, the errors made under each genus will so far counteract one another, that the mean results for the genera and orders will scarcely be affected. As it is, the method adopted has absorbed many weeks of labor during the last five years, and a much greater degree of accuracy could only have been obtained by a disproportionately greater outlay of time, whilst it would not have materially affected the general results.

With regard to my own views on the subjects of the variability of existing species and the fallacy of supposing we can ascertain anything through these alone of their ancestry or of originally created types, they are, in so far as they are liable to influence my estimate of the value of the facts collected for the analysis of the Australian Flora, unaltered from those which I maintained in*the 'Flora of New Zealand.' On such theoretical questions, however, as the origin and ultimate permanence of species, they have been greatly influenced by the views and arguments of Mr. Darwin and Mr. Wallace above alluded to, which incline me to regard more favorably the hypothesis that it is to variation that we must look as the means which Nature has adopted for peopling the globe with those diverse existing forms which, when they tend to transmit their characters unchanged through many generations, are called species. Nevertheless I must repeat, what I have fully stated elsewhere, that these hypotheses should not influence our treatment of species, either as subjects of descriptive science, or as the means of investigating the phenomena of the succession of organic forms in time, or their dispersion and replacement in area, though they should lead us to more philosophical conceptions on these subjects, and stimulate us to seek for such combinations of their characters as may enable us to classify them better, and to trace their origin back to an epoch anterior to that of their present appearance and condition. In doing this, however, the believer in species being lineally related forms must employ the same methods of investigation and follow the same principles that guide the believer in their being actual creations, for the latter assumes that Nature has created species with mutual relations analogous to those which exist between the lineally-descended members of a family, and this is indeed the leading idea in all natural systems. On the other hand, there are so many checks to indiscriminate variation, so many inviolable laws that regulate the production of varieties, the time required to produce wide variations from any given specific type is so great, and the number of species and varieties known to propagate for indefinite periods a succession of absolutely identical members is so large, that all naturalists are agreed that for descriptive purposes species must be treated as if they were at their origin distinct, and are destined so to remain. Hence the descriptive naturalist who believes all species to be derivative and mutable, only differs in practice from him who asserts the contrary, in expecting that the posterity of the organism he describes as species may, at some indefinitely distant period of time, require redescription.

I need hardly remark that the classificatory branch of Botany is the only one from which this subject can be approached; for a good system must be founded on a due appreciation of all the attributes of individual plants,—upon a balance of their morphological, physiological, and anatomical relations at all periods of their growth. Species are conventionally assumed to represent, with a great amount of uniformity, the lowest degree of such relationship; and the facts that individuals are more easily grouped into species limited by characters, than into varieties, or than species are into limitable genera or groups of higher value, and that the relationships of species are transmitted hereditarily in a very eminent degree, are the strongest appearances in favor of species being original creations, and genera, etc., arbitrarily limited groups of these.

The difference between varieties and species and genera in respect to definable limitation is however one of degree only, and if increased materials and observation confirm the doctrine which I have for many years labored to establish, that far more species are variable, and far fewer limitable, than has been supposed, that hypothesis will be proportionally strengthened which assumes species to be arbitrarily limited groups of varieties. With the view of ascertaining how far my own experience in classification will bear out such a conclusion, I shall now endeavor to review, without reference to my previous conclusions, the impressions which I have derived from the retrospect of twenty years' study of plants. During that time I have classified many large and small Floras, arctic, temperate and tropical, insular and continental: embracing areas so extensive ana varied as to justify, to my apprehension, the assumption that the results derived from these would also be applicable to the whole vegetable kingdom. I shall arrange these results successively under three heads; viz., facts derived from a study of classification; secondly, from distribution; thirdly, from fossils; after which I shall examine the theories with which these facts should harmonize.

§ 2. On the General Phenomena of Variation in the Vegetable

Kingdom.

1. All vegetable forms are more or less prone to vary as to their sensible properties, or (as it has been happily expressed in regard to all organisms), "they are in a state of unstable equilibrium."* No organ is exactly symmetrical, no two are exact

* Essays: Scientific, Political, and Speculative; by Herbert Spencer: p. 280.

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