The paradox of Vito Volterra's predator-prey model

This article is dedicated to the late Giorgio Israel. R{\'e}sum{\'e}. The aim of this article is to propose on the one hand a brief history of modeling starting from the works of Fibonacci, Robert Malthus, Pierre Francis Verhulst and then Vito Volterra and, on the other hand, to present the main hypotheses of the very famous but very little known predator-prey model elaborated in the 1920s by Volterra in order to solve a problem posed by his son-in-law, Umberto D'Ancona. It is thus shown that, contrary to a widely-held notion, Volterra's model is realistic and his seminal work laid the groundwork for modern population dynamics and mathematical ecology, including seasonality, migration, pollution and more. 1. A short history of modeling 1.1. The Malthusian model. If the rst scientic view of population growth seems to be that of Leonardo Fibonacci [2], also called Leonardo of Pisa, whose famous sequence of numbers was presented in his Liber abaci (1202) as a solution to a population growth problem, the modern foundations of population dynamics clearly date from Thomas Robert Malthus [20]. Considering an ideal population consisting of a single homogeneous animal species, that is, neglecting the variations in age, size and any periodicity for birth or mortality, and which lives alone in an invariable environment or coexists with other species without any direct or indirect inuence, he founded in 1798, with his celebrated claim Population, when unchecked, increases in a geometrical ratio, the paradigm of exponential growth. This consists in assuming that the increase of the number N (t) of individuals of this population, during a short interval of time, is proportional to N (t). This translates to the following dierential equation : (1) dN (t) dt = $\epsilon$N (t) where $\epsilon$ is a constant factor of proportionality that represents the growth coe-cient or growth rate. By integrating (1) we obtain the law of exponential growth or law of Malthusian growth (see Fig. 1). This law, which does not take into account the limits imposed by the environment on growth and which is in disagreement with the actual facts, had a profound inuence on Charles Darwin's work on natural selection. Indeed, Darwin [1] founded the idea of survival of the ttest on the 1. According to Frontier and Pichod-Viale [3] the correct terminology should be population kinetics, since the interaction between species cannot be represented by forces. 2. A population is dened as the set of individuals of the same species living on the same territory and able to reproduce among themselves.