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Alfred Russel Wallace : Alfred Wallace : A. R. Wallace :
Russel Wallace : Alfred Russell Wallace (sic)

The Comparative Richness of Faunas and
Floras Tested Numerically (S280: 1877)

Editor Charles H. Smith's Note: A letter to the Editor printed in the 6 December 1877 number of Nature. Original pagination indicated within double brackets. To link directly to this page, connect with: http://people.wku.edu/charles.smith/wallace/S280.htm

    [[p. 100]] In his letter in Nature, vol. xvii. p. 9, Prof. Newton1 has strongly brought out the absurdity of comparing districts of very different areas by the proportionate number of species to area in each. On this principle he shows that to be equally rich with the small island of Rodriguez, Madagascar ought to possess four times as many species of birds as exist throughout the whole world. It does not, however, by any means follow that the method thus exposed may not be of value in comparing regions of approximately equal area, as is the case with several of the primary regions, to determine the comparative richness of which Mr. Sclater2 first applied it. I have not Mr. Sclater's paper at hand, but it is my impression that he made no attempt to show--"that the proper mode of comparing the wealth or poverty of one fauna with another was to state the proportion which the number of species composing it bears to the area over which they range"--as Prof. Newton implies that he did, but that he merely adopted this method as the only one readily available for the comparison of his regions. Although I took the opportunity of making some corrections in the figures, I never committed myself to the principle; and I very soon afterwards found that it was not to be trusted. As, however, several later writers have made use of it without remark, it will be interesting to consider where the exact point of the fallacy lies, and with what modifications the method can be trusted to give useful and consistent results.

    If we compare two islands of almost exactly equal areas, such as Ceylon and Tasmania, and find that the one has twice or three times as many species of mammals or birds as the other, it will be generally admitted that we express the fact correctly when we say that, as regards such a group of animals, the one is twice or thrice as rich as the other; and the same may be said of two countries or two continents of identical areas. For on the supposition that there is a general correspondence between the numbers of rare and common, of local and of wide-spread species in the two areas compared (and this seems probable), then the average number of distinct species to be met with on one spot, or to be seen during a journey of equal length, will be proportionate to the total number of species in the two areas. But now let us divide one of the two continents or islands which we are comparing into two or more parts. We know, as a matter of fact, that one-half the area will always contain much more than half the total number of species, while one-tenth of the area will contain immensely more than one-tenth of the species. To take an example: the county of Sussex is about one-eightieth part the area of the British Isles, yet it actually contains full two-thirds of the total number of flowering plants, both being estimated by the same flora (Babington's "Manual," fifth edition, British Isles 1,536 species, Sussex 1,059 species). If we now compare either Britain or Sussex with an equal area on the continent of Europe or North America, we may obtain an instructive estimate of the comparative richness of their respective floras; but if we compare unequal areas, and then endeavour to equalise them by getting the proportions of species to area, we shall obtain erroneous results, which will become literally absurd when the areas compared are very unequal.

    The problem remains, how to compare unequal areas of which we possess the zoological or botanical statistics. We can only do so by equalising them, and this may not be so difficult as at first sight appears. For example, let us take the Palæarctic and North American regions, in which the species of birds are nearly equal in number, but the areas are as about seven to three. The number of the Palæarctic species have, however, been proportionately increased of late years, and if we take the western half of the Palæarctic region so as to include North Africa and Persia we shall have an area about equal to the Nearctic region, and a number of species perhaps one-sixth or one-eighth less, which will thus represent the comparative richness of these two areas. The eastern half of the region, including Japan and North China, is probably as rich as the western; while the intermediate portion is poorer in species. Combining these three portions, and taking the average, we should perhaps find the Palæarctic region about four-fifths or five-sixths as rich as the Nearctic, instead of less than one-half, as shown by the method of proportionate areas.

    Whenever we know how many peculiar species any district contains, we can deduct its area from the total area of the region to be compared, and this number of peculiar species, from the fauna of the region; and by this means we may reduce two unequal regions to comparative equality. Again, all detached portions or islands should be omitted in estimating the comparative richness of regions, because they affect these regions very unequally. By adding Britain to Europe you increase the area without adding to the fauna, and thus make the region seem poorer; while by adding Madagascar to Africa, or New Zealand [[p. 101]] to Australia, you add to the fauna in a greater proportion than you increase the area, and thus make the region seem richer. For a fair comparison continents should be compared with continents, and islands with islands, and these should in every case be brought to an approximate equality of area by lopping off outlying portions with their peculiar species. We shall then get results which will be instructive, and which will afford us a true estimate of the comparative richness of different countries in the several classes of animals and plants.

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Editor's Notes

1Alfred Newton (1829-1907), English ornithologist
2Philip Lutley Sclater (1829-1913), English ornithologist

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