These differences are illustrated in the accompanying coloured plate, which has been most kindly prepared for me under the instructions of Dr. F. M. Chapman of the American Museum of Natural History. Before going further it is worth considering the nature of these differences a little more closely. All but the last are large differences which no one would overlook even in a hasty glance at the birds. If the only distinction lay in the colour of the quills we might feel fairly sure that auratus was a recessive form of cafer, and so probably it is in this respect. Similarly the black malar stripe of auratus is in all probability recessive to the red malar stripe of cafer and I imagine the pigments concerned are comparable with those in the Gouldian Finch (Poephila gouldiae) of Australia. Both sexes in that species may have the head black, red, or, less often, yellow, and though it is not any longer in question that birds may breed in either plumage, I believe that the young are always black-headed and I imagine that those which become red-headed possess a dominant factor absent from the permanently black-headed birds.[2] Yellow as a recessive form of a red is certainly very common, but red and black as variants of the same pigment are less usual. In the Gouldian Finch we seem to have a case where a pigment can assume all three forms. It would be interesting to know whether the red of the malar stripes in Colaptes is a pigment of the same nature as the red of the quills. Both in Colaptes and in Poephila gouldiae I have seen specimens intermediate between the black and the red, and the appearance of the part affected was exactly alike in the two cases, red feathers coming up among the black ones, and many feathers containing both red and black pigments mixed together.
The development of the scarlet nuchal crescent in auratus and the absence of this conspicuous mark in cafer constitute from the physiological point of view the most remarkable pair of differences. When the red crescent is not formed, the feathers which would bear it are exactly like the rest, and no special pigment is visible in them which one can regard as ready to be modified into red. If the crescent is due to a factor it must therefore be supposed that this factor has the power of modifying the pigment of the neck in one special place alone. Dr. W. D. Miller called my attention to the fact that a similar variation occurs in another American woodpecker, the Sapsucker, Sphyropicus varius.[3]
I do not suggest that such variations are without parallel: indeed in P. gouldiae the factor which turns the black of the head into scarlet affects one special region of the black only, being sharply distinct from the unmodified black of the throat. These regions of the head are however often the seat of special colours in birds.[4] So also may be instanced the variety of the Common Guillemot (Uria troile) which has a white line round the eyes and at the sides of the head where the normal has no such mark; but this line is formed in a very special place, the groove joining the eye to the ear, whereas the feathers of the nuchal crescent are not ostensibly distinguished from those adjacent.[5]
The transposition of the brown and the grey on the back and front of the neck also constitutes a very remarkable difference. If either grey or brown depends on a factor then it must be supposed that auratus has one of these factors and cafer the other.
From these several considerations it is quite clear that if auratus and cafer are modifications of the same type produced by presence or absence of factors, several independent elements must be concerned, and to unravel their inter-relations would be most difficult even if it were possible to breed the types under observation, which is of course quite beyond present possibilities.
The distribution of the two is as follows. On the east side of the Continent C. auratus, relatively pure, occupies the whole of Canada and the States from the North to Galveston. Westward it extends across the whole continent in the more northern region to Alaska, but in its pure form it only reaches down the Pacific coast to about the northern border of British Columbia. Its southern and western limit is thus roughly a line drawn from north of Vancouver, southeast to North Dakota and then south to Galveston. C. cafer in the comparatively pure form inhabits Mexico, Arizona, California (except Lower California and the opposite coast), central and western Nevada, Utah, Oregon, and is bounded on the east by a line drawn from the Pacific south of Washington, south and eastward through Colorado to the mouth of the Rio Grande or the Gulf of Mexico. Between the two lines thus roughly defined is a band of country about 1,200-1,300 miles long and 300-400 miles wide, which contains some normal birds of each type, but chiefly birds exhibiting the characters of both, mixed together in various and irregular ways. Even in the areas occupied by the pure forms occasional birds are recorded with more or less indication of characteristics of the other form, but within the area in which the two forms are conterminous, the mixed birds are in the majority. The condition of these birds of mixed character is described by Allen as follows:
"As has been long known—indeed, as shown by Baird in 1858—the 'intermediates' or 'hybrids' present ever-varying combinations of the characters of the two birds, from individuals of C. auratus presenting only the slightest traces of the characters of C. cafer, or, conversely—individuals of C. cafer presenting only the slightest traces of the characters of C. auratus—to birds in which the characters of the two are about equally blended. Thus we may have C. auratus with merely a few red feathers in the black malar stripe, or with the quills merely slightly flushed with orange, or C. cafer with either merely a few black feathers in the red malar stripe, or a few red feathers at the sides of the nape, or an incipient, barely traceable scarlet nuchal crescent. Where the blending of the characters is more strongly marked, the quills may be orange-yellow or orange-red, or of any shade between yellow and red, with the other features of the two birds about equally blended. But such examples are exceptional, an unsymmetrical blending being the rule, the two sides of the same bird being often unlike. The quills of the tail, for example, may be part red and part yellow, the number of yellow or red feathers varying in different individuals, and very often in the opposite sides of the tail in the same bird. The same irregularity occurs also, but apparently less frequently, in the quills of the wings. In such cases the quills may be mostly yellow with a few red or orange quills intermixed, or red with a similar mixture of yellow. A bird may have the general colouration of true cafer combined with a well-developed nuchal crescent, or nearly pure auratus with the red malar stripes of a cafer. Sometimes the body plumage is that of C. auratus with the head nearly as in pure cafer, or exactly the reverse may occur. Or we may have the general plumage as in cafer with the throat and crown as in auratus, and the malar stripe either red or black, or mixed red and black, and so on in almost endless variations, it being rare to find, even in birds of the same nest, two individuals alike in all their features of colouration. Usually the first trace of cafer seen in auratus manifests itself as a mixture of red in the black malar stripe, either as a few red feathers, or as a tipping of the black feathers with red, or with merely the basal portion of the feathers red. Sometimes, however, there is a mixture of orange or reddish quills, while the malar stripe remains normal. In C. cafer the traces of auratus are usually shown by a tendency to an incipient nuchal crescent, represented often by merely a few red-tipped feathers on the sides of the nape; at other times by a slight mixture of black in the red malar stripe."
Such a state of things accords very imperfectly with expectations under any received theory of Evolution. As in some of the instances discussed in the first chapter we have here two fairly definite forms, nearly allied, which on any evolutionary hypothesis must have been evolved either the one from the other, or both from a third form at a time not very remote from the present, as time must be measured in evolution. Yet though intermediates exist in some quantity, no one can for a moment suggest that they are that definite intermediate from which auratus and cafer descend in common. One cannot imagine that the immediate ancestor of these birds was a mosaic, made up of asymmetrical patches of each sort: but that is what many of the intermediates are. It is not much easier to suppose the ancestor to have been a nondescript, with a compromise between the developed characters of each, with quills buff, malar stripes neither black nor red, with a trace of nuchal crescent, and so on. Such Frankenstein-monsters have played, a considerable part in the imaginations of evolutionary philosophers, but if it were true that there was once a population of these monsters capable of successful existence, surely they should now be found as a population occupying the neutral zone between the two modern forms. Yet, though much remains to be done in clearing up the facts, one thing is certain, namely that the neutral zone has not a definite and normally intermediate population, but on the contrary it is peopled by fragments of the two definite types and miscellaneous mongrels between them.
On the other hand, one cannot readily suppose that either form was the parent of the other. The process must have involved both addition and loss of factors, for whatever hypothesis be adopted, such changes must be supposed to have occurred. A careful statistical tabulation of the way in which the characters are distributed in the population of the mixed zone would be of great value, and till that has been done there is little that can be said with certainty as to the genetics of these characters. In the collection of Dr. Bishop of New Haven I was very kindly allowed to examine a sample, all taken at random, near together, in Saskatchewan. There were females 4 adult, 2 young; males 4 adult and 5 young. This number, though of course insufficient, is enough to give some guide as to the degree of definiteness which the characters generally show in their variations. Of the 15 birds, 8 had simply yellow quills; 2 had red; 1 was almost red but had one yellow tail-quill; 3 were intermediate and 1 was buff. As regards the malar patch, which can only be determined properly in the adult males, 1 was red, 1 was approximately red, 2 intermediate. As to nuchal crescent 4 females had none, 2 females very slight; 7 males had it, 1 had only a slight crescent, and 1 had none. In point of quills therefore 10 were definite out of 15; in point of crescent, 11 were definite out of 15; and in point of malar patch 1 only was definite out of 4. The last is a feature directly dependent on age and so counts for less, but as regards the other two features there is some indication that the factors show definiteness in their behaviour. It must be remembered that we have no knowledge what the heterozygous form may be, and in the case of red and yellow it is probably a reddish buff. The patch-works are no doubt to be compared with other well-known pied forms, and in these we must suppose the active factor broken up, which it probably can be very easily. The asymmetry, which Allen notices as so marked a feature, in the distribution of the red and yellow quills of the tail especially, recalls that of the black markings in the pied Canaries. As is well known to students of variations some pigment-factors in some animals are apparently uncontrolled by symmetry, while in other specific cases symmetry is the rule. On the other hand the blackness or redness of the malar patches is, I think, as a rule nearly symmetrical. It should be mentioned that two of Dr. Bishop's young birds belonged to the same nest, one a female with red quills, the other a male with yellow. Both are without crescent.
As to the question whether certain combinations of characters occur with special frequency, the evidence is insufficient to give a definite answer. Among all the birds I have seen in America or in England I have not yet found one having the malar patches black without any nuchal crescent. Of Dr. Bishop's 8 adults not one, however, showed the combination of the three chief features normal for auratus or for cafer.