GENETIC VARIATION IN THE HORSESHOE CRAB (LIMULUS POLYPHEMUS), A PHYLOGENETIC "RELIC".

To many early evolutionists, particularly those of the mutationist school, the existence of evolutionarily stable forms ("living fossils" or phylogenetic "relics") posed a difficult problem, and, even among contemporary biologists, the feeling that "a long-standing stability of organization seems antithetical to the concept of evolution" may persist (Volpe, 1967:140). In the early part of this century, it was popularly believed that stable or slowly evolving (bradytelic) lines lacked the genetic variation necessary for more rapid evolution as a result of having abnormally low mutation rates; as Mayr (1963) puts it, a lack of variation in time and space was interpreted as a reflection of genotypic stability. Largely because of the influence of Simpson (1944), Stebbins (1944), Schmalhausen (1949), and Dobzhansky (1951), the mutationists' interpretation of bradytely has now been all but abandoned by evolutionists in favor of the view that the principal factors controlling rates of evolution are to be found in the organism-environment relationship. Additionally, Mayr (1963 :304) has suggested that phenotypic "uniformity in space and time is due to a highly perfected 'buffering system,' in other words, to genetic homeostasis." The ecological hypothesis is well expressed in Simpson's (1944:141) conclusions that "the final and probably the most fundamental factor in the relationship is that bradytelic groups are so well adapted to a particular, continuously available environment that almost any mutation occurring in them must be disadvantageous" and that "bradytely results from the equilibrium of large breeding populations of animals specifically adapted to a continuously available environment that is relatively invariable . . . ." Yet, as Mayr (1963) notes, there is no evidence to refute (or to support) the hypothesis that bradytely results from loss of mutability. The recent development of electrophoretic techniques for demonstrating allelic variation at loci controlling enzymes and other proteins in natural populations (Hubby and Lewontin, 1966; Lewontin and Hubby, 1966) permits a test of the mutationists' thesis. The purpose of this study is to compare degrees of protein polymorphism and underlying genic heterozygosity in a classic phylogenetic "relic," the horseshoe or king crab (Limulus polyphemus), and in several representatives of horotelic lines (those evolving at standard rates) for which data are already available. Evidence of a level of genetic variation in Limulus similar to that in horotelic forms will support the modern ecological and homeostatic interpretations of bradytely. But if the mutationists' thesis is valid, Limulus should show an unusually low degree of polymorphism at its genetic loci. Limulus polyphemus is one of five living species of the Subclass Xiphosura, a group of marine arthropods usually placed with the Subclass Eurypterida (extinct giant water scorpions) in the Class Merostomata of the Subphylum Chelicerata. Following a period of adaptive radiation in the early and middle Paleozoic, the Merostomata became much less prominent after the Permian. Several Mesozoic forms are known in the Family Mesolimulidae, but by the Tertiary the fossil record is all but absent. There are presently three living genera, all belonging to the Family Limulidae, only one of which, Tachypleus, is known from the fossil record (Stormer,