“The fact that such metallic colors can be retained in dead specimens by putting a drop of glycerine under the elytra, leads us to conclude that those colors are based upon fat substances. The hypodermal colors are never glossy, as far as I know; the dermal colors frequently.
“As the wings, elytra, and hairs all possess a cuticula, dermal colors are frequently to be found, together with hypodermal ones, chiefly in metallic colors. In the same place both colors may be present, or one of them alone. So we find hypodermal colors in the elytra of Lampyridæ. In the elytra of the Cicindelidæ the main metallic color is dermal, the white lines or spots are hypodermal, by which arrangement the variability in size and shape of those spots is explained.
“There occur in a number of insects external colors, that is, colors upon the cuticula, which I consider to be in fact displaced hypodermal colors: the mealy pale blue or white upon the abdomen of some Odonata, the white on many Hemiptera, the pale gray on the elytra and on the thorax of the Goliath beetle, and the yellowish powder on Lixus. Some of these colors dissolve easily by ether or melt in heat, and some of them are a kind of wax. I believe that those colors are produced in the hypodermis, and are exuded through the pore-canals.” (Hagen.)
The white colors are simply for the most part due to the inclusion of air in scales. The white mother-of-pearl spots of Argynnis are produced by a system of fine transverse pore-canals filled with air; in Hydrometra the white ventral marks have the same origin. (Leydig.)
The further statements and criticisms of Hagen regarding the relation of color to mimicry, sexual selection, and the origin of patterns are of much weight and will be referred to under those heads. Indeed, these subjects cannot well be discussed without reference to the fundamental facts stated in the masterly papers of Leydig and of Hagen, and much of the theorizing of these latter days is ill-founded, because the colors of insects and animals are attributed to natural selection, when they seem really the result of the action of the primary factors of organic evolution, such as changes of light, heat, cold, and chemical processes dependent on the former.
As to the chemical nature of color, Hagen, after quoting the results of Krukenberg and others, thinks that the colors of insects are chemically produced by a combination of fats or fat-acids with other acids or alkalis under the influence of air, light, and heat. He concludes:—
1. That some colors of insects can be changed or obliterated by acids.
2. That two natural colors, madder-lake and indigo, can be produced artificially by the influence of acid on fat-bodies.
3. As protein bodies in insects are changed into fat-bodies, and may be changed by acids contained in insects into fat-acids, the formation of colors in the same manner seems probable.
4. That colors can be changed by different temperatures.