to those of North America. In eastern Beringia from 35,000 YBP the northern Arctic areas experienced temperatures 1.5 °C (2.7 °F) warmer than today, but the southern sub-Arctic regions were 2 °C (3.5 °F) cooler. In 22,000YBP, during the Last Glacial Maximum, the average summer temperature was 3–5 °C (5.4–9 °F) cooler than today, with variations of 2.9 °C (5.2 °F) cooler on the Seward Peninsula to 7.5 °C (13.5 °F) cooler in the Yukon. Beringia received more moisture and intermittent maritime cloud cover from the north Pacific Ocean than the rest of the Mammoth steppe, including the dry environments on either side of it. Moisture occurred along a north–south gradient with the south receiving the most cloud cover and moisture due to the airflow from the North Pacific. This moisture supported a shrub-tundra habitat that provided an ecological refugium for plants and animals. In this Beringian refugium, eastern Beringia's vegetation included isolated pockets of larch and spruce forests with birch and alder trees. This environment supported large herbivores that were prey for Beringian wolves and their competitors. Steppe bison (Bison priscus), Yukon horse (Equus lambei), woolly mammoth (Mammuthus primigenius), and Wild yak (Bos mutus) consumed grasses, sedges, and herbaceous plants. Caribou (Rangifer tarandus) and woodland muskox (Symbos cavifrons) consumed tundra plants, including lichen, fungi, and mosses. Prey Isotope analysis can be used to allow researchers to make inferences about the diet of the species being studied. Two isotope analyses of bone collagen extracted from the remains of Late Pleistocene wolves found in Beringia and Belgium indicate that wolves from both areas preyed mainly on Pleistocene megafauna, which became rare at the beginning of the Holocene 12,000 years ago. The Beringian wolf preyed most often on horse and steppe bison. In the period leading up to the Last Glacial Maximum (50,000YBP–23,000YBP), they also ate woodland muskox, and after this time they also ate mammoth. The analysis supports the conclusion that these wolves were capable of killing and dismembering large prey. In another stable isotope analysis, half of the Beringian wolves were found to be muskox and caribou specialists, and the other half were either horse and bison specialists or generalists. Two wolves from the full-glacial period (23,000–18,000 YBP) were found to be mammoth specialists, but it is not clear if this was due to scavenging or predation. The analysis of other carnivore fossils from the Fairbanks region of Alaska found