Climate Change, Humans, and the Extinction of the Woolly Mammoth

Abstract
Woolly mammoths inhabited Eurasia and North America from late Middle Pleistocene (300 ky BP [300,000 years before present]), surviving through different climatic cycles until they vanished in the Holocene (3.6 ky BP). The debate about why the Late Quaternary extinctions occurred has centred upon environmental and human-induced effects, or a combination of both. However, testing these two hypotheses—climatic and anthropogenic—has been hampered by the difficulty of generating quantitative estimates of the relationship between the contraction of the mammoth's geographical range and each of the two hypotheses. We combined climate envelope models and a population model with explicit treatment of woolly mammoth–human interactions to measure the extent to which a combination of climate changes and increased human pressures might have led to the extinction of the species in Eurasia. Climate conditions for woolly mammoths were measured across different time periods: 126 ky BP, 42 ky BP, 30 ky BP, 21 ky BP, and 6 ky BP. We show that suitable climate conditions for the mammoth reduced drastically between the Late Pleistocene and the Holocene, and 90% of its geographical range disappeared between 42 ky BP and 6 ky BP, with the remaining suitable areas in the mid-Holocene being mainly restricted to Arctic Siberia, which is where the latest records of woolly mammoths in continental Asia have been found. Results of the population models also show that the collapse of the climatic niche of the mammoth caused a significant drop in their population size, making woolly mammoths more vulnerable to the increasing hunting pressure from human populations. The coincidence of the disappearance of climatically suitable areas for woolly mammoths and the increase in anthropogenic impacts in the Holocene, the coup de grâce, likely set the place and time for the extinction of the woolly mammoth. What caused the woolly mammoth's extinction? Climate warming in the Holocene might have driven the extinction of this cold-adapted species, yet the species had survived previous warming periods, suggesting that the more-plausible cause was human expansion. Testing these competing hypotheses has been hampered by the difficulty in generating quantitative estimates of the relationship between the mammoth's contraction and the climatic and/or human-induced drivers of extinction. In this study, we combined paleo-climate simulations, climate envelope models (which describe the climate associated with the known distribution of a species—its envelope—and estimate that envelope's position under different climate change scenarios), and a population model that includes an explicit treatment of woolly mammoth–human interactions to measure the extent to which climate changes, increased human pressures, or a combination of both factors might have been responsible. Results show a dramatic decline in suitable climate conditions for the mammoth between the Late Pleistocene and the Holocene, with hospitable areas in the mid-Holocene being restricted mainly to Arctic Siberia, where the latest records of woolly mammoths in continental Asia have been found. The population model results also support the view that the collapse of the climatically suitable area caused a significant drop in mammoth population size, making the animals more vulnerable to increasing hunting pressure from expanding human populations. The coincidence of the collapse of climatically suitable areas and the increase in anthropogenic impacts in the Holocene are most likely to have been the “coup de grâce,” which set the place and time for the extinction of the woolly mammoth.