3. Modification in the colour of Salamandra maculosa induced by change in the colour of the soil on which the animals were kept.
1. I will take first the case of Alytes,[12] because it is the most definite example, and because it is the case which most readily admits of repetition and verification.
The habits of Alytes obstetricans are well known. The animals copulate on land. As the strings of eggs leave the female they are entangled by the hind legs of the male, and being adhesive they stick to him and undergo their development attached to his back and legs. The number of eggs varies from 18 to 86, a number much smaller than is usual in toads and frogs which lay their eggs in water. The eggs are large and full of yolk.
There are two breeding seasons, one about April and the other about September, and a winter hibernation. Not only animals brought in from outside, but their offspring reared in domestication maintain these normal habits in confinement, if the temperature does not exceed 17° C. (pp. 499 and 534).
If, however, the temperature be artificially raised and kept at 25-30° C., the males do not attach the eggs to themselves when spawning occurs on land but let them lie. The adhesion of the eggs is said to be hindered by the comparatively rapid drying of their surfaces.
More usually in the high temperatures the animals take to the water and copulate there. The eggs are ejected into the water, and as their gelatinous coverings immediately swell up, they do not stick to the males.
The offspring thus derived from the parents subjected to heat for one breeding-period only, whether they were laid in water or on land, did not show departures from the normal type.
Kammerer states next, however, that in subsequent breeding-periods the same parents frequently take to the water to breed, though they have become quite accustomed to the heated chamber; and furthermore that if such animals, having thus lost their instinct to brood their young, be transferred to ordinary temperatures they do not readily reassume their normal habits, but for several breeding seasons—at least four—will take to the water. These parents lay from 90 to 115 eggs, which are small and contain little yolk, and the larvae, on hatching, breathe with their embryonic gills until they are absorbed instead of being broken off as normally.
The offspring thus abnormally developed when they mature are said never to brood their eggs. If they are derived from the earlier spawnings of their parents, before, that is to say, the parents had been submitted to the changed conditions long enough to transmit their effects, they lay on land; but if they are derived from the later spawnings, they lay in the water. These changes of habit are manifested without the continued application of the abnormal experimental conditions, and, as I understand the account, in normal conditions of temperature.
If the abnormal experimental conditions are continued, the toads always lay in water, and their eggs become progressively smaller and more numerous. The larvae in the fourth generation acquire three pairs of gills instead of one pair, and are in other respects also different from the normal form.