Milankovitch cycles and their effects on species in ecological and evolutionary time

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Abstract
The Quaternary ice ages were paced by astronomical cycles with periodicities of 20–100 k.y. (Milankovitch cycles). These cycles have been present throughout earth history. The Quaternary fossil record, marine and terrestrial, near to and remote from centers of glaciation, shows that communities of plants and animals are temporary, lasting only a few thousand years at the most. Response of populations to the climatic changes of Quaternary Milankovitch cycles can be taken as typical of the way populations have behaved throughout earth history. Milankovitch cycles thus force an instability of climate and other aspects of the biotic and abiotic environment on time scales much less than typical species durations (1–30 m.y.). Any microevolutionary change that accumulates on a time scale of thousands of years is likely to be lost as communities are reorganized following climatic changes. A four-tier hierarchy of time scales for evolutionary processes can be constructed as follows: ecological time (thousands of years), Milankovitch cycles (20–100 k.y.), geological time (millions of years), mass extinctions (approximately 26 m.y.). “Ecological time” and “geological time” are defined temporally as the intervals between events of the second and fourth tiers, respectively. Gould's (1985) “paradox of the first tier” can be resolved, at least in part, through the undoing of Darwinian natural selection at the first tier by Milankovitch cycles at the second tier.