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Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo, and that she occasionally laid an egg in another bird's nest. If the old bird profited by this occasional habit through being enabled to migrate earlier or through any other cause; or if the young were made more vigorous by advantage being taken of the mistaken .. instinct of another species than when reared by their own mother, .. encumbered as she could hardly fail to be by having eggs and young of different ages at the same time; then the old birds or the fostered young would gain an advantage. And analogy would lead us to believe, that the young thus reared would be apt to follow by inheritance the occasional and aberrant habit of their mother, and in their turn would be apt to lay their eggs in other birds' nests, and thus be more successful in rearing their young. By a continued process of this nature, I believe that the strange instinct of our cuckoo ... has been generated. It has, also, recently been ascertained on sufficient evidence, by Adolf Müller, that the cuckoo occasionally lays her eggs on the bare ground, sits on them, and feeds her young. This rare event is probably a case of reversion to the long-lost, aboriginal instinct of nidification.
It has been objected by some authors that I have not noticed other related instincts and points of structure in the cuckoo, which are falsely spoken of as necessarily co-ordinated. But in all cases, speculation on any instinct or character known in only a single species, is useless, for we have no facts to guide us. Until quite recently the instincts of the European and of the non-parasitic American cuckoo alone were known; now, owing to Mr. E. Ramsay's observations, we know something about three Australian species, which lay their eggs in other birds' nests. The chief points referred to are three: firstly, that the cuckoo, with rare exceptions, lays only one egg in a nest, so that the large and voracious young cuckoo receives ample food. Secondly, that the egg is so remarkably small, that it does not exceed in size that of the skylark,—a bird not more than one-fourth of the size of the cuckoo; that this is a real case of adaptation we may infer from the fact of the non-parasitic American cuckoo laying eggs of full size proportionally with her body. Thirdly and lastly, that the young cuckoo, soon after birth, has the instinct, the strength, and a properly shaped back for ejecting its foster-brothers, which then perish from cold and hunger. This, it has been boldly maintained, is beneficently designed, in order that the young cuckoo may get sufficient food, and that its foster-brothers may perish, before, as it is supposed, they have acquired much feeling!
Turning now to the Australian species; though these birds generally lay only one egg in a nest, it is not rare to find two and even three eggs of the same species of cuckoo in the same nest. In the Bronze cuckoo the eggs vary greatly in size, from eight to ten lines in length. Now if it had been of any advantage to this species to have laid eggs even smaller than those now laid by her, so as to have deceived certain foster-parents, or, as is more probable, to have been hatched within some shorter period (for it is asserted that there is a relation between the size of eggs and the period of incubation), then there is no difficulty in believing that a race or species might have been formed which would have laid smaller and smaller eggs; for these would have been more safely hatched and reared. Mr. Ramsay remarks that two of the Australian cuckoos, when they lay their eggs in an open or not domed nest, manifest a decided preference for nests containing eggs similar to their own. The European species certainly manifests some tendency towards a similar instinct, but not rarely departs from it, as is shown by her laying her dull and pale-coloured eggs in the nest of the Hedge-warbler with its bright greenish-blue eggs: had she invariably displayed the above instinct, it would assuredly have been added to those which it is assumed must all have been acquired together. The eggs of the Australian Bronze cuckoo vary, according to Mr. Ramsay, in an extraordinary manner in colour; so that in this respect, as well as in size, natural selection assuredly might have secured and fixed any advantageous variation.
In the case of the European cuckoo, the offspring of the foster-parents are commonly ejected from the nest within three days after the cuckoo is hatched; and as the latter at this age is in a most helpless condition, Mr. Gould was formerly inclined to believe that the act of ejection was performed by the foster-parents themselves. But he has now received a trustworthy account of a young cuckoo which was actually seen, whilst still blind and not able even to hold up its own head, in the act of ejecting its foster-brothers. One of these was replaced in the nest by the observer, and was again thrown out. Now, if it had been of great importance to the young cuckoo to have received as much food as possible during the first few days after birth, I can see no especial difficulty, if it possessed sufficient strength, in its gradually acquiring, during successive generations, the habit (first, perhaps, through mere unintentional restlessness) and the structure best fitted for ejecting its foster-brothers; for those young cuckoos which had such habits and structure would have been the best fed and most securely reared. The first step towards the acquisition of the proper instinct might have been mere unintentional restlessness on the part of the young bird, when somewhat advanced in age and strength; the habit having been afterwards improved, and transmitted to an earlier age. I can see no more difficulty in this, than in young birds acquiring the instinct and the temporary hard tips to their beaks for breaking through their own shells;—or than in the young snake having in its upper jaw, as Owen has remarked, a transitory sharp tooth for cutting through the tough egg-shell. For if each part is liable to variation at any age, and the variations tend to be inherited at a corresponding age,—propositions which cannot, as we shall hereafter see, rightfully be disputed,—then the instincts and structure of the young can be slowly modified as well as those of the adult, and both cases must stand or fall together with the whole theory of natural selection.
Some species of Molothrus, a widely distinct genus of American birds, allied to our starlings, have parasitic habits like those of the cuckoo; and the species present an interesting gradation in the perfection of their instincts. The sexes of Molothrus badius are stated by an excellent observer, Mr. Hudson, sometimes to live promiscuously together in flocks, and sometimes to pair. They either build a nest of their own, or seize on one belonging to some other bird, occasionally throwing out the nestlings of the stranger. They either lay their eggs in the nest thus appropriated, or oddly enough build one for themselves on the top of it. They usually sit on their own eggs and rear their own young; but Mr. Hudson says it is probable that they are occasionally parasitic, for he has seen the young of this species following old birds of a distinct kind and clamouring to be fed by them. The parasitic habits of another species of Molothrus, the M. bonariensis, are much more highly developed than those of the last, but are still far from perfect. This bird, as far as it is known, invariably lays its eggs in the nests of strangers; but it is remarkable that several together sometimes commence to build an irregular untidy nest of their own, placed in singularly ill-adapted situations, as on the leaves of a large thistle. They never, however, as far as Mr. Hudson has ascertained, complete a nest for themselves. They often lay so many eggs— from fifteen to twenty— in the same foster-nest, that few or none can possibly be hatched. They have, moreover, the extraordinary habit of pecking holes in the eggs, whether of their own species or of their foster-parents, which they find in the appropriated nests. They drop also many eggs on the bare ground, which are thus wasted. A third species, the M. pecoris of North America, has acquired instincts as perfect as those of the cuckoo, for it never lays more than one egg in a foster-nest, so that the young bird is securely reared. Mr. Hudson is a strong disbeliever in evolution, but he appears to have been so much struck by the imperfect instincts of the Molothrus bonariensis that he quotes my words, and asks, "Must we consider these habits, not as especially endowed or created instincts, but as small consequences of one general law, namely, transition? "
Various birds, as has already been remarked, occasionally lay their eggs in the nests of other birds. This habit is not very uncommon with the Gallinaceæ, and throws some light on the singular instinct of the ... ostrich. In this family several hen-birds unite and lay first a few eggs in one nest and then in another; and these are hatched by the males. This instinct may probably be accounted for by the fact
Nevertheless, I could give several instances of various birds which have been known occasionally to lay their eggs in other birds' nests. Now let us suppose that the ancient progenitor of our European cuckoo had the habits of the American cuckoo; but that occasionally she laid an egg in another bird's nest. If the old bird profited by this occasional habit, or if the young were made more vigorous by advantage having been taken of the mistaken maternal instinct of another bird, than by their own mother's care, encumbered as she can hardly fail to be by having eggs and young of different ages at the same time, and by having to migrate at a very early period; then the old birds or the fostered young would gain an advantage. And analogy would lead me to believe, that the young thus reared would be apt to follow by inheritance the occasional and aberrant habit of their mother, and in their turn would be apt to lay their eggs in other birds' nests, and thus be successful in rearing their young. By a continued process of this nature, I believe that the strange instinct of our cuckoo could be, and has been, generated. I may add that, according to Dr. Gray and to some other observers, the European cuckoo has not utterly lost all maternal love and care for her own offspring.
The occasional habit of birds laying their eggs in other birds' nests, either of the same or of a distinct species, is not very uncommon with the Gallinaceæ; and this perhaps explains the origin of a singular instinct in the allied group of ostriches. For several hen ostriches, at least in the case of the American species, unite and lay first a few eggs in one nest and then in another; and these are hatched by the males. This instinct may probably be accounted for by the fact