As a cub, Alex the lion was called Alekey and was the son of Zuba (Bernie Mac), the alpha lion. Though Zuba tries to teach Alekey to be a hunter, the cub is more interested in dancing. When another lion named Makunga (Alec Baldwin) challenges Zuba to a fight for the position of alpha lion, Alekey is captured by poachers. Though Zuba tries to rescue his son, Alekey is forced into a crate. The crate falls into the ocean where it drifts to New York. There, Alekey is renamed Alex and sent to the Central Park Zoo where he grew up, meeting Gloria, Marty, and Melman there.
In the present time, Alex (Ben Stiller), Gloria (Jada Pinkett Smith), Melman (David Schwimmer), Marty (Chris Rock), the chimpanzees, King Julien, Maurice, and the penguins have fixed a crashed plane in the hopes of using it to fly back to New York (Mort manages to tag along as well, by clinging to the airplane wing). Though the plane does fly, it ends up crash-landing in Africa because of lack of fuel.
In Africa, the animals are amazed to find more of their kind. Alex is reunited with Zuba and his mother Florrie (Sherri Shephard). Marty quickly fits into a herd of zebra who all look, sound, and talk exactly like him. Melman takes on the position of Witch Doctor amongst the animals after discovering that sick giraffes have no other option but to die otherwise. Gloria, interested in finding a boyfriend, attracts the attention of a hippo named Moto Moto (Will.i.am).
Meanwhile, the penguins set about repairing the plane. They carjack several jeeps from New Yorkers on vacation (under "Operation Tourist Trap"), leaving the tourists stranded and lost in the jungle. A tough old woman called "Nana" (who had previously attacked Alex in a subway station in Madagascar) takes charge, reminding them that they are New Yorkers and always survive.
Unfortunately, life in Africa is not as wonderful as it first seems. Makunga, still determined to take the position of alpha lion, reminds Zuba that Alex must complete a traditional lion coming-of-age challenge. Alex, thinking that the challenge is a dance contest (it is actually a fight) competes against the strongest lion and loses quickly. To avoid being forced to banish Alex for failing, Zuba abdicates and Makunga immediately takes the position and banishes Alex and his family. Marty meanwhile begins to feel upset that all of the zebra are exactly the same as him, leaving him with nothing unique. Even Alex is unable to tell the difference between Marty and a different zebra, leaving Marty hurt. Melman is happy in his position as Witch Doctor until another giraffe tells him that he has a brown spot on his shoulder. Melman is initially not bothered (as he has brown spots all over his body), until he learns that the previous Witch Doctor apparently died two days after the "spot" appeared. Melman is also upset about Gloria dating Moto Moto, as Melman has secretly loved her for a long time. Gloria goes on a date with Moto Moto and quickly realizes that he only loves her for her body.
The next day, the animals are in a panic when they discover that the watering hole has dried up. Determined to make up for his earlier failure, Alex decides to risk being shot by hunters and leave the Reserve to discover what happened. Along the way, he makes amends with Marty by being able to pick his friend out of the herd of zebras. The pair discover that the stranded New Yorkers, under the instruction of Nana, have dammed the river and built a primitive civilization. Alex is captured and Marty runs for help. Meanwhile, Zuba hears what Alex did and goes to rescue him.
King Julien suggests that the animals sacrifice one of themselves into the volcano to appease the water gods and regain their water supply. Believing that he will die soon, Melman volunteers to be the sacrifice. Gloria stops him just in time and Marty arrives to tell them of Alex's fate. The trio, the penguins, and a cartload of chimpanzees use the newly-fixed plane to come to the rescue. Alex meanwhile manages to rescue both himself and his father by dancing for the New Yorkers, who quickly recognize him from the zoo. Zuba joins in as well. The other animals rescue the lions with the plane and destroy the dam, freeing the water. At the volcano, Mort arrives (he had fallen from the plane into the ocean). He was pursued the entire way from the beach by a shark, which falls into the lava just before the water returns.
Alex manages to remove Makunga from power by tricking Nana into attacking him. Skipper (a penguin) marries a bobble-head hula doll from the plane and leaves on a honeymoon, leaving the other animals to remain in Africa.
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