Revisiting the exclusion principle in epidemiology at its ultimate limit

TL;DR

This paper challenges the traditional epidemiological exclusion principle by demonstrating that even with a significant transmissibility advantage, two pathogen strains can stably coexist, contradicting the expectation of eventual dominance.

Contribution

It provides a rigorous analysis showing that the exclusion principle does not always hold when one strain has a large transmissibility advantage, especially under broad conditions.

Findings

Coexistence of strains with vastly different transmissibility is possible.

The exclusion principle is not unconditional and can be invalidated.

Stable endemic equilibrium with multiple strains can occur despite competitive advantages.

Abstract

The competitive exclusion principle in epidemiology implies that when competing strains of a pathogen provide complete protection for each other, the strain with the largest reproduction number outcompetes the other strains and drives them to extinction. The introduction of various trade-off mechanisms may facilitate the coexistence of competing strains, especially when their respective basic reproduction numbers are close so that the competition between the strains is weak. Yet, one may expect that a substantial competitive advantage of one of the strains will eventually outbalance trade-off mechanisms driving less competitive strains to extinction. The literature, however, lacks a rigorous validation of this statement. In this work, we challenge the validity of the exclusion principle at an ultimate limit in which one strain has a vast competitive advantage over the other strains. We show that when one strain is significantly more transmissible than the others, and under broad conditions, an epidemic system with two strains has a stable endemic equilibrium in which both strains coexist with comparable prevalence. Thus, the competitive exclusion principle does not unconditionally hold beyond the established case of complete immunity.