Bloodstained from feeding, an adult female is greeted on her return to the den by two cubs that may or may not be her own. Spotted hyena clans are matriarchal, with females dominating groups of 80 or more animals. Clans often splinter into smaller cells, reassembling days or weeks later. To help ease the reintegration process, hyenas of all ages, ranks, and sexes engage in ceremonial one-on-one greetings. Hierarchical in all things, hyena etiquette usually requires the submissive animal to initiate the greeting.
Mile after mile the chase goes on. The gazelle tires, but not the spotted hyenas in pursuit. With a final surge of speed, the predators spring upon the flagging prey, drag it down, and disembowel it. Then a roar echoes across the East African savanna. The hyenas flee their feeding as a male lion takes over the kill. The thwarted hunters skulk nearby with empty bellies.
Hyenas have an undeserved reputation as thieves and scavengers that subsist on the leavings of the larger predator. "But it is far more frequent that the lion will steal a kill from the hyenas," says Kay Holekamp of Michigan State University. Biologists have known this for decades, she laments, yet hyenas are still viewed as "slobbering, mangy, stupid poachers" (not to mention goose-stepping fascists) in The Lion King, the movie that for many has defined the species.
Why do people grimace at the sight of them? With their patchy fur and odd proportions, maybe they flout our shallow standards for beauty in animals. "Our obsession with looks doesn't take into account how well their bodies and brains are adapted to an ecosystem," says Anup Shah, who, with his brother, Manoj, photographed hyenas in Kenya, their homeland, and Tanzania. Among Africans, hyenas arouse humor and horror—the former because the genitals of the females inexplicably mimic those of males, giving rise to the myth that hyenas are hermaphrodites, and the latter because of a link with death. The Masai leave corpses in the bush for hyenas to dispose of. Indeed, hyenas eat almost anything, which makes them valuable. "They are very important to the health of the ecosystem," says Marion East of the Institute for Zoo and Wildlife Research in Berlin. "As scavengers, they clean up a huge amount of dead matter. As hunters, they probably help maintain the genetic health of the great herds."
Cheetahs are usually too smart to tangle with hyenas. But someone forgot to tell these cubs, and a rare showdown ensues on the savanna. Soon after bringing down a gazelle, a mother cheetah and her three nearly grown cubs get an unwelcome dinner guest—a lone female hyena. The cats know hyenas will steal any food they can, so they often time their hunts to coincide with hyenas' midday naps, from about 10 a.m. to 4 p.m. Sometimes the strategy fails. With the wisdom of age, the mother cat immediately gives way to the more aggressive hyena. Her brash youngsters, however, mount a challenge. One cub makes a frontal charge before ﬂitting away. The second buzzes the slower hyena. "They must have been really hungry," says Michigan State's Kay Holekamp. "I've never seen a cheetah go after a hyena. It's a dangerous thing to do." Though the confrontation may have been unusual, the outcome was not. The hyena eats its fill in peace, while a young cheetah can only glare and hope for some leftovers.
Returning home with fast food—a chunk of speedy Thomson's gazelle—this young hyena has no intention of sharing. It could lose its lunch entirely, however, if a dominant hyena challenges it. That's why animals that rank low in a clan's hierarchy learn to eat fast and eat alone. This gazelle head isn't a prime cut, but the spotted hyena is adapted perhaps better than any other predator to making do with rougher fare. Vise-grip jaws and specialized teeth help slice through thick skin and stubborn tendons with ease. Hyenas will even crunch up bone, digesting the organic content and excreting calcium.
A growl and bared fangs signal that this voracious predator is fed up with the vultures vying for a wildebeest carcass it wants to devour single-handedly—an impossibility even for the hungriest hyena. Unsure of when they will next feed, hyenas will gorge themselves whenever they can. Although they often snap at vultures in the process, they rarely harm the rival scavengers. The relationship between the species is mutually beneficial: The birds often eat at hyena kills, and the sight of vultures alerts the hyenas to nearby dead meat. This hyena, disturbed by the swarm of scavengers, finally tore off a hunk of wildebeest meat and went looking for a more peaceful spot.
After a big meal—big meaning not a few steaks, but 30 to 40 pounds (14 to 18 kilograms) of meat, nearly a third of their weight and far more, proportionally, than a lion would eat—digestion raises hyena body temperature. On such occasions, few things are more pleasant than a cool, postprandial mud bath. But even in the mud, the hierarchy rules. A lower ranking animal lounging in the same spot quickly made way for this dominant female.
Snarling in the face of certain death, a hyena cornered by lions in a Masai Mara bog has nowhere to run. Male lions seem to relish harassing and killing their smaller competitors. After delivering the hyena a fatal bite and toying with its carcass, this male and another lion began to play. "They had all the signs of excitement—eyes shining, and they were rolling in the grass nuzzling each other," says photographer Anup Shah. "It was a successful assassination."
Shoulder to shoulder, a clan faces down a young male lion off camera that has come into the hyenas' territory in Masai Mara. Tails up, ears back, and emitting an eerie call of aggression known as a "low," they edge closer, but the lion stands its ground. Had it been a full-grown male, such a show of force—the same tactic hyenas use when confronting rival clans at the edges of their territory—would have been too dangerous even to attempt. Safe near a den in Tanzania's Serengeti, two members of a clan appear to be engaged in a painful fight, but they're just playing a friendly game, Anup Shah says. "It appeared one was saying, 'I trust you. You can do whatever you want with those teeth. I know you won't harm me.'