This paper describes the last of a series of field trials designed to discover the factors preventing unlimited increase in the numbers of the short-tailed vole (Microtus agrestis). The present study may be regarded as a pilot test of the hypothesis that 'a decline in numbers is a necessary consequence of the selective action of hostility against the genotypes present during the phase of rapid increase' (Chitty & Phipps 1961). Here we describe findings which, for the first time in the course of these investigations, did not compel us to modify our initial point of view. Unfortunately they did not enable us to establish it either, since they merely confirmed that further progress was impossible without critical studies of behaviour and genetics. One such study on behaviour has since been performed by Sadleir (1965), using the deermouse Peromyscus maniculatus. The eliminative procedures used in earlier studies are described in the first paper in the series (Chitty 1952, especially pp. 535-7), and in later papers also; but because some people still believe in the ideas thus discarded we shall summarize the events that led up to the present study-references and reviews are given by Chitty (1960), Newson & Chitty (1962), Barnett (1964) and Krebs (1964). An assumption that is essential to the whole argument is that these studies are concerned with a single phenomenon, i.e. that the population changes are not mere changes in numbers but are associated with diagnostic events. These other events include variations in mean adult body weight, differences in survival rates between juveniles and adults, and between males and females, together with such less constant features as a tendency for fluctuations to recur every 4 years and to be synchronous with those in neighbouring populations. Given a recognizable class of events and enough comparable instances we can eliminate as not sufficient all factors which are sometimes present when the phenomenon is absent, and as not necessary all factors which are absent when the phenomenon is present. (One does not, of course, reach the absurd conclusion that factors so eliminated play no part in producing numerical differences of other kinds.) Prior to 1959 the more obvious explanations had been rejected for reasons explained by Elton (1942) and subsequent workers. On the positive side it was known that: (a) populations were qualitatively different at different stages of abundance; (b) declines continued for one or more generations after peak densities had been passed; (c) individuals from rapidly declining populations survived well in captivity; and (d) some but not all of these individuals reached high body weights in captivity, whereas none did in the field. The present hypothesis is consistent with these findings and, though not known to be true, has the merit of being highly testable, since falsifying instances can be looked for in other species (Chitty 1964, 1965). As shown in the following account it also offers a possible explanation for otherwise puzzling changes in survival. The nearest alternative explanation is that of Christian & Davis (1964), who believe that there are 'endocrine mechanisms which have the prolonged effects necessary to * Present address: Department of Zoology, University of British Columbia, Vancouver 8, B.C., Canada.
Read full abstract