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Nature of the Checks to Increase - Chapter III - Origin of Species

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That the mere physiological importance of an organ does not determine its classificatory value, is almost proved by the fact, that in allied groups, in which the same organ, as we have every reason to suppose, has nearly the same physiological value, its classificatory value is widely different. No naturalist can have worked at any group without being struck with this fact; and it has been fully acknowledged in the writings of almost every author. It will suffice to quote the highest authority, Robert Brown, who, in speaking of certain organs in the Proteaceae, says their generic importance, "like that of all their parts, not only in this, but, as I apprehend in every natural family, is very unequal, and in some cases seems to be entirely lost." Again, in another work he says, the genera of the Connaraceae "differ in having one or more ovaria, in the existence or absence of albumen, in the imbricate or valvular aestivation. Any one of these characters singly is frequently of more than generic importance, though here even, when all taken together, they appear insufficient to separate Cnestis from Connarus." To give an example among insects: in one great division of the Hymenoptera, the antennae, as Westwood has remarked, are most constant in structure; in another division they differ much, and the differences are of quite subordinate value in classification; yet no one will say that the antennae in these two divisions of the same order are of unequal physiological importance. Any number of instances could be given of the varying importance for classification of the same important organ within the same group of beings.

Again, no one will say that rudimentary or atrophied organs are of high physiological or vital importance; yet, undoubtedly, organs in this condition are often of much value in classification. No one will dispute that the rudimentary teeth in the upper jaws of young ruminants, and certain rudimentary bones of the leg, are highly serviceable in exhibiting the close affinity between Ruminants and Pachyderms. Robert Brown has strongly insisted on the fact that the position of the rudimentary florets is of the highest importance in the classification of the Grasses.

Numerous instances could be given of characters derived from parts which must be considered of very trifling physiological importance, but which are universally admitted as highly serviceable in the definition of whole groups. For instance, whether or not there is an open passage from the nostrils to the mouth, the only character, according to Owen, which absolutely distinguishes fishes and reptiles--the inflection of the angle of the lower jaw in Marsupials--the manner in which the wings of insects are folded--mere colour in certain Algae--mere pubescence on parts of the flower in grasses--the nature of the dermal covering, as hair or feathers, in the Vertebrata. If the Ornithorhynchus had been covered with feathers instead of hair, this external and trifling character would have been considered by naturalists as an important aid in determining the degree of affinity of this strange creature to birds.

The importance, for classification, of trifling characters, mainly depends on their being correlated with many other characters of more or less importance. The value indeed of an aggregate of characters is very evident in natural history. Hence, as has often been remarked, a species may depart from its allies in several characters, both of high physiological importance, and of almost universal prevalence, and yet leave us in no doubt where it should be ranked. Hence, also, it has been found that a classification founded on any single character, however important that may be, has always failed; for no part of the organisation is invariably constant. The importance of an aggregate of characters, even when none are important, alone explains the aphorism enunciated by Linnaeus, namely, that the characters do not give the genus, but the genus gives the character; for this seems founded on the appreciation of many trifling points of resemblance, too slight to be defined. Certain plants, belonging to the Malpighiaceae, bear perfect and degraded flowers; in the latter, as A. de Jussieu has remarked, "The greater number of the characters proper to the species, to the genus, to the family, to the class, disappear, and thus laugh at our classification." When Aspicarpa produced in France, during several years, only these degraded flowers, departing so wonderfully in a number of the most important points of structure from the proper type of the order, yet M. Richard sagaciously saw, as Jussieu observes, that this genus should still be retained among the Malpighiaceae. This case well illustrates the spirit of our classifications.

Practically, when naturalists are at work, they do not trouble themselves about the physiological value of the characters which they use in defining a group or in allocating any particular species. If they find a character nearly uniform, and common to a great number of forms, and not common to others, they use it as one of high value; if common to some lesser number, they use it as of subordinate value. This principle has been broadly confessed by some naturalists to be the true one; and by none more clearly than by that excellent botanist, Aug. St. Hilaire. If several trifling characters are always found in combination, though no apparent bond of connexion can be discovered between them, especial value is set on them. As in most groups of animals, important organs, such as those for propelling the blood, or for aerating it, or those for propagating the race, are found nearly uniform, they are considered as highly serviceable in classification; but in some groups all these, the most important vital organs, are found to offer characters of quite subordinate value. Thus, as Fritz Muller has lately remarked, in the same group of crustaceans, Cypridina is furnished with a heart, while in two closely allied genera, namely Cypris and Cytherea, there is no such organ; one species of Cypridina has well-developed branchiae, while another species is destitute of them.

We can see why characters derived from the embryo should be of equal importance with those derived from the adult, for a natural classification of course includes all ages. But it is by no means obvious, on the ordinary view, why the structure of the embryo should be more important for this purpose than that of the adult, which alone plays its full part in the economy of nature. Yet it has been strongly urged by those great naturalists, Milne Edwards and Agassiz, that embryological characters are the most important of all; and this doctrine has very generally been admitted as true. Nevertheless, their importance has sometimes been exaggerated, owing to the adaptive characters of larvae not having been excluded; in order to show this, Fritz Muller arranged, by the aid of such characters alone, the great class of crustaceans, and the arrangement did not prove a natural one. But there can be no doubt that embryonic, excluding larval characters, are of the highest value for classification, not only with animals but with plants. Thus the main divisions of flowering plants are founded on differences in the embryo--on the number and position of the cotyledons, and on the mode of development of the plumule and radicle. We shall immediately see why these characters possess so high a value in classification, namely, from the natural system being genealogical in its arrangement.

Our classifications are often plainly influenced by chains of affinities. Nothing can be easier than to define a number of characters common to all birds; but with crustaceans, any such definition has hitherto been found impossible. There are crustaceans at the opposite ends of the series, which have hardly a character in common; yet the species at both ends, from being plainly allied to others, and these to others, and so onwards, can be recognised as unequivocally belonging to this, and to no other class of the Articulata.

Geographical distribution has often been used, though perhaps not quite logically, in classification, more especially in very large groups of closely allied forms. Temminck insists on the utility or even necessity of this practice in certain groups of birds; and it has been followed by several entomologists and botanists.

Finally, with respect to the comparative value of the various groups of species, such as orders, suborders, families, subfamilies, and genera, they seem to be, at least at present, almost arbitrary. Several of the best botanists, such as Mr. Bentham and others, have strongly insisted on their arbitrary value. Instances could be given among plants and insects, of a group first ranked by practised naturalists as only a genus, and then raised to the rank of a subfamily or family; and this has been done, not because further research has detected important structural differences, at first overlooked, but because numerous allied species, with slightly different grades of difference, have been subsequently discovered.

All the foregoing rules and aids and difficulties in classification may be explained, if I do not greatly deceive myself, on the view that the natural system is founded on descent with modification--that the characters which naturalists consider as showing true affinity between any two or more species, are those which have been inherited from a common parent, all true classification being genealogical--that community of descent is the hidden bond which naturalists have been unconsciously seeking, and not some unknown plan of creation, or the enunciation of general propositions, and the mere putting together and separating objects more or less alike.

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