Qualitative Changes in Populations in Unstable Environments

Abstract Inactive moths of Malacosoma pluviale (Dyar) oviposit near their birthplaces, and most of their offspring also are inactive. More active moths can travel farther before they oviposit, and always have a higher proportion of vigorous individuals among their progeny. Such polymorphism allows the insect to cope with environmental diversity; e.g., inactive residents exploit favourable habitats, and active migrants colonize more severe habitats, or replenish the vigour of other populations. Because the most active moths usually export the most vigorous progeny, the population left behind becomes less vigorous during successive generations. This steadily decreasing vitality eliminates local populations that are not replenished by vigorous immigrants. Qualitative changes in Malacosoma populations follow this basic pattern, but the rate of deterioration is affected by the habitat. Departing migrants fly too high to be stopped by small trees in farmland, bur many are stopped near their source by tall trees in forests. Deterioration therefore is slower in forests. Forests also delay return migration to nearby farmlands, and thus allow some farmland populations to deteriorate unchecked. In a fluctuating climate, the size of the area tolerable for the species varies annually. When it begins to expand, the vigorous progeny of active moths can take immediate advantage of slight local improvements. Consequently, they are the first to exploit each marginal habitat that becomes tolerable. But while better climate persists, some less active descendants of these pioneers appear in all occupied habitats. When the regional climate deteriorates, the tolerable area contracts, and most marginal populations are totally destroyed. Moreover, even within the contracted tolerable area, the harsher climate becomes intolerable for any deteriorated stock. In the next generation, therefore, the only regional survivors are vigorous colonies deposited in the tolerable area by some of the few migrants that escaped the widespread destruction of the preceding generation in the margins. Their descendants recolonize depopulated sections of the refuge, and so preserve the species in the region while the climate remains severe.