The Variation of Animals and Plants under Domestication — Volume 2. Darwin Charles

or peloric, the change is generally looked at by botanists as a return to the primitive state. But Dr. Maxwell Masters (13/68. 'Natural Hist. Review' April 1863 page 258. See also his Lecture, Royal Institution, March 16, 1860. On same subject see Moquin-Tandon 'Elements de Teratologie' 1841 pages 184, 352. Dr. Peyritsch has collected a large number of very interesting cases 'Sitzb. d. k. Akad. d. Wissensch.' Wien b. 60 and especially b. 66 1872 page 125.), who has ably discussed this subject, remarks that when, for instance, all the sepals of a Tropaeolum become green and of the same shape, instead of being coloured with one prolonged into a spur, or when all the petals of a Linaria become simple and regular, such cases may be due merely to an arrest of development; for in these flowers all the organs during their earliest condition are symmetrical, and, if arrested at this stage of growth, they would not become irregular. If, moreover, the arrest were to take place at a still earlier period of development, the result would be a simple tuft of green leaves; and no one probably would call this a case of reversion. Dr. Masters designates the cases first alluded to as regular peloria; and others, in which all the corresponding parts assume a similar form of irregularity, as when all the petals in a Linaria become spurred, as irregular peloria. We have no right to attribute these latter cases to reversion, until it can be shown that the parent-form, for instance, of the genus Linaria had had all its petals spurred; for a chance of this nature might result from the spreading of an anomalous structure, in accordance with the law, to be discussed in a future chapter, of homologous parts tending to vary in the same manner. But as both forms of peloria frequently occur on the same individual plant of the Linaria (13/69. Verlot 'Des Varietes' 1865 page 89; Naudin 'Nouvelles Archives du Museum' tome 1 page 137.), they probably stand in some close relation to one another. On the doctrine that peloria is simply the result of an arrest of development, it is difficult to understand how an organ arrested at a very early period of growth should acquire its full functional perfection; — how a petal, supposed to be thus arrested, should acquire its brilliant colours, and serve as an envelope to the flower, or a stamen produce efficient pollen; yet this occurs with many peloric flowers. That pelorism is not due to mere chance variability, but either to an arrest of development or to reversion, we may infer from an observation made by Ch. Morren (13/70. In his discussion on some curious peloric Calceolarias quoted in 'Journal of Horticulture' February 24, 1863 page 152.) namely, that families which have irregular flowers often "return by these monstrous growths to their regular form; whilst we never see a regular flower realise the structure of an irregular one."

      Some flowers have almost certainly become more or less completely peloric through reversion, as the following interesting case shows. Corydalis tuberosa properly has one of its two nectaries colourless, destitute of nectar, only half the size of the other, and therefore, to a certain extent, in a rudimentary state; the pistil is curved towards the perfect nectary, and the hood, formed of the inner petals, slips off the pistil and stamen in one direction alone, so that, when a bee sucks the perfect nectary, the stigma and stamens are exposed and rubbed against the insect's body. In several closely allied genera, as in Dielytra, etc., there are two perfect nectaries, the pistil is straight, and the hood slips off on either side, according as the bee sucks either nectary. Now, I have examined several flowers of Corydalis tuberosa, in which both nectaries were equally developed and contained nectar; in this we see only the redevelopment of a partially aborted organ; but with this redevelopment the pistil becomes straight, and the hood slips off in either direction, so that these flowers have acquired the perfect structure, so well adapted for insect agency, of Dielytra and its allies. We cannot attribute these coadapted modifications to chance, or to correlated variability; we must attribute them to reversion to a primordial condition of the species.

      The peloric flowers of Pelargonium have their five petals in all respects alike, and there is no nectary so that they resemble the symmetrical flowers of the closely allied genus Geranium; but the alternate stamens are also sometimes destitute of anthers, the shortened filaments being left as rudiments, and in this respect they resemble the symmetrical flowers of the closely allied genus Erodium. Hence we may look at the peloric flowers of Pelargonium as having reverted to the state of some primordial form, the progenitor of the three closely related genera of Pelargonium, Geranium, and Erodium.

      In the peloric form of Antirrhinum majus, appropriately called the" Wonder," the tubular and elongated flowers differ wonderfully from those of the common snapdragon; the calyx and the mouth of the corolla consist of six equal lobes, and include six equal instead of four unequal stamens. One of the two additional stamens is manifestly formed by the development of a microscopically minute papilla, which may be found at the base of the upper lip of the flower of the common snapdragons in the nineteen plants examined by me. That this papilla is a rudiment of a stamen was well shown by its various degrees of development in crossed plants between the common and the peloric Antirrhinum. Again, a peloric Galeobdolon luteum, growing in my garden, had five equal petals, all striped like the ordinary lower lip, and included five equal instead of four unequal stamens; but Mr. R. Keeley, who sent me this plant, informs me that the flowers vary greatly, having from four to six lobes to the corolla, and from three to six stamens. (13/71. For other cases of six divisions in peloric flowers of the Labiatae and Scrophulariaceae see Moquin- Tandon 'Teratologie' page 192.) Now, as the members of the two great families to which the Antirrhinum and Galeobdolon belong are properly pentamerous, with some of the parts confluent and others suppressed, we ought not to look at the sixth stamen and the sixth lobe to the corolla in either case as due to reversion, any more than the additional petals in double flowers in these same two families. But the case is different with the fifth stamen in the peloric Antirrhinum, which is produced by the redevelopment of a rudiment always present, and which probably reveals to us the state of the flower, as far as the stamens are concerned, at some ancient epoch. It is also difficult to believe that the other four stamens and the petals, after an arrest of development at a very early embryonic age, would have come to full perfection in colour, structure, and function, unless these organs had at some former period normally passed through a similar course of growth. Hence it appears to me probable that the progenitor of the genus Antirrhinum must at some remote epoch have included five stamens and borne flowers in some degree resembling those now produced by the peloric form. The conclusion that peloria is not a mere monstrosity, irrespective of any former state of the species, is supported by the fact that this structure is often strongly inherited, as in the case of the peloric Antirrhinum and Gloxinia and sometimes in that of the peloric Corydalis solida. (13/72. Godron reprinted from the 'Memoires de l'Acad. de Stanislas' 1868.)

      Lastly I may add that many instances have been recorded of flowers, not generally considered as peloric, in which certain organs are abnormally augmented in number. As an increase of parts cannot be looked at as an arrest of development, nor as due to the redevelopment of rudiments, for no rudiments are present, and as these additional parts bring the plant into closer relationship with its natural allies, they ought probably to be viewed as reversions to a primordial condition.]

      These several facts show us in an interesting manner how intimately certain abnormal states are connected together; namely, arrests of development causing parts to become rudimentary or to be wholly suppressed, — the redevelopment of parts now in a more or less rudimentary condition, — the reappearance of organs of which not a vestige can be detected, — and to these may be added, in the case of animals, the presence during youth, and subsequent disappearance, of certain characters which occasionally are retained throughout life. Some naturalists look at all such abnormal structures as a return to the ideal state of the group to which the affected being belongs; but it is difficult to conceive what is meant to be conveyed by this expression. Other naturalists maintain, with greater probability and distinctness of view, that the common bond of connection between the several foregoing cases is an actual, though partial, return to the structure of the ancient progenitor of the group. If this view be correct, we must believe that a vast number of characters, capable of evolution, lie hidden in every organic being. But it would be a mistake to suppose that the number is equally great in all beings. We know, for instance, that plants of many orders occasionally become peloric; but many more cases have been observed in the Labiatae and Scrophulariaceae than in any other order; and in one genus of the Scrophulariaceae, namely Linaria, no less than thirteen species have been described in this condition (13/73. Moquin- Tandon 'Teratologie' page 186.) On this view of the nature of peloric flowers, and bearing in mind certain monstrosities in the animal kingdom, we must