is mostly grasses and sedges, but they will opportunistically consume bark, leaves, buds, fruits, and roots. Compared to ruminants, zebras have a simpler and less efficient digestive system. Nevertheless, they can subsist on lower-quality vegetation. Zebras may spend 60–80% of their time feeding, depending on the availability of vegetation. The plains zebra is a pioneer grazer, mowing down the upper, less nutritious grass canopy and preparing the way for more specialised grazers, which depend on shorter and more nutritious grasses below. Zebras are preyed on mainly by lions. Leopards, cheetahs, spotted hyenas, brown hyenas and wild dogs pose less of a threat to adults. Nile crocodiles also prey on zebras when they near water. Biting and kicking are a zebra's defense tactics. When threatened by lions, zebras flee, and when caught they are rarely effective in fighting off the big cats. In one study, the maximum speed of a zebra was found to be while a lion was measured at . Zebras do not escape lions by speed alone but by sideways turning, especially when the cat is close behind. With smaller predators like hyenas and dogs, zebras may act more aggressively, especially in defense of their young. Social structure Zebra species have two basic social structures. Plains and mountain zebras live in stable, closed family groups or harems consisting of one stallion, several mares, and their offspring. These groups have their own home ranges, which overlap, and they tend to be nomadic. Stallions form and expand their harems by recruiting young mares from their natal (birth) harems. The stability of the group remains even when the family stallion is displaced. Plains zebras groups gather into large herds and may create temporarily stable subgroups within a herd, allowing individuals to interact with those outside their group. Females in harems can spend more time feeding, and gain protection both for them and their young. The females have a linear dominance hierarchy with the high-ranking females being the ones that have lived in the group longest. While traveling, the most dominant females and their offspring lead the group, followed by the next most dominant. The family stallion trails behind. Young of both sexes leave their natal groups as they mature; females are usually herded by outside males to become part of their harems. In the more arid-living Grévy's zebras, adults have more fluid associations and adult males establish large territories, marked by