by Greg Mayer

Darwin’s theory of evolution (and ours), unlike that of Lamarck, is variational, rather than transformational: the process of evolution is a change in frequency of different variants within a population, not a transformation of the individuals.  Darwin thus made the origin, nature, and inheritance of variation key problems for biology; indeed, for much of the 20th century, evolution and genetics were often taught as a single course at universities.

One of the most distinctive sorts of variation is polymorphism, in which two or more discontinuous forms are found in a single species (this is distinct from sexual or age related variation). Darwin himself pioneered the study of polymorphisms. Such discontinuous variation often has a simple genetic basis, with allelic variation at one genetic locus accounting for all (or most) of the variability.The color polymorphism in peppered moths (Biston betularia) is a well known and well studied case involving industrial melanism, in which light and dark forms are adapted to polluted and unpolluted environments, respectively. A well known case of polymorphism in vertebrates are the two color phases of Cuban sparrow hawk (Falco sparverius sparverioides). This case is not well studied, though, and we know nothing about the genetics, nor the adaptive significance (if any) of the polymorphism.

A polymorphism in vertebrates that many Americans and Canadians are familiar with are the melanistic and gray forms of the gray squirrel (Sciurus carolinensis). The most frequent color form is gray, but blackish or dark brownish individuals are widely distributed, and in places quite frequent. I have seen them in Illinois (Cook County), Wisconsin (Racine and Kenosha Cos.) and Michigan (Ingham Co.), and also on the campus of Princeton University. (I was told at Princeton that, during football season, black squirrels are captured, and orange stripes applied to them, so that they resemble diminutive arboreal tigers, the tiger being Princeton’s mascot.)

A much less common color morph is the leucistic or albinistic form, which is whitish, cream or yellowish. They are famously common in Olney, Illinois (due to an introduction of two albinistic individuals to an area previously lacking any gray squirrels at all), and also occur regularly in Stevens Point, Wisconsin, but I had never seen one before my recent trip to Washington, DC, where I saw one on the tree right across from the steps on the Mall entrance to the USNM.  (The picture was taken through a bus window.)

Vertebrate polymorphisms are often less well understood than those of invertebrates, because their generally greater size and longer generation times make experimental study more difficult. Melanism in squirrels, for example, has been related to thermoregulation and fire frequency, but no thoroughly compelling explanation has been found. One exception to this is coat color variation in mice of the genus Peromyscus, where coat color seems to be an adaptation for camouflage in varying environments.

In the 1930s, F.B. Sumner conducted classic field and lab studies on light colored mice living on sandy soils and dark mice on dark soils. Unlike the melanistic and albinistic squirrels, which are variant individuals within a populations, there is an element of geographic variation in the mice, which live in distinct, though adjacent, places. Sumner’s studies showed that there were several (not just one) genetic loci involved in coat color, and the color forms intergrade where their habitats meet and they interbreed. Hopi Hoekstra of the Museum of Comparative Zoology is currently conducting exciting studies of some of the same species studied by Sumner.

Although the mice occur in distinct modal forms (white vs. brown), the intergradation where they meet shows an underlying continuous variation. The frogs below show that although we can pick out distinctly different individuals, the range of pattern from plain to mottled to striped makes it difficult to recognize a small number of discrete color morphs, and the variation approaches a continuous dictribution. Such continuous variations were thought by Darwin, and most biologists today as well, to be important raw material for the evolutionary process.