biological designation. The first to challenge the concept of race on empirical grounds were the anthropologists Franz Boas, who provided evidence of phenotypic plasticity due to environmental factors, and Ashley Montagu, who relied on evidence from genetics. E. O. Wilson then challenged the concept from the perspective of general animal systematics, and further rejected the claim that "races" were equivalent to "subspecies". Human genetic variation is predominantly within races, continuous, and complex in structure, which is inconsistent with the concept of genetic human races. According to the biological anthropologist Jonathan Marks, Subspecies The term race in biology is used with caution because it can be ambiguous. Generally, when it is used it is effectively a synonym of subspecies. (For animals, the only taxonomic unit below the species level is usually the subspecies; there are narrower infraspecific ranks in botany, and race does not correspond directly with any of them.) Traditionally, subspecies are seen as geographically isolated and genetically differentiated populations. Studies of human genetic variation show that human populations are not geographically isolated, and their genetic differences are far smaller than those among comparable subspecies. In 1978, Sewall Wright suggested that human populations that have long inhabited separated parts of the world should, in general, be considered different subspecies by the criterion that most individuals of such populations can be allocated correctly by inspection. Wright argued that, "It does not require a trained anthropologist to classify an array of Englishmen, West Africans, and Chinese with 100% accuracy by features, skin color, and type of hair despite so much variability within each of these groups that every individual can easily be distinguished from every other." While in practice subspecies are often defined by easily observable physical appearance, there is not necessarily any evolutionary significance to these observed differences, so this form of classification has become less acceptable to evolutionary biologists. Likewise this typological approach to race is generally regarded as discredited by biologists and anthropologists. Ancestrally differentiated populations (clades) In 2000, philosopher Robin Andreasen proposed that cladistics might be used to categorize human races biologically, and that races can be both biologically real and socially constructed. Andreasen cited tree diagrams of relative genetic distances among populations published by Luigi Cavalli-Sforza as the basis for a phylogenetic tree of human races (p. 661). Biological anthropologist Jonathan Marks (2008) responded by arguing that Andreasen had misinterpreted the genetic literature: "These trees are phenetic