In the past, Alex (Ben Stiller) the lion lived in Africa with his father Zuba (Bernie Mac), the pride's alpha lion, and mother Florrie (Sherri Shepherd), along the rest of the pride, including Makunga (Alec Baldwin), who always wanted to be the leader. One day Alex—known as Alakay to his parents—is captured by poachers after his father turns away for a moment because of Makunga. Zuba chases after his captured son, but fails in his attempt to rescue him. However, Alakay's crate falls into a nearby river and drifts off to sea. Alex is then discovered, found washed ashore in New York, were he is put in the zoo, named Alex, and later becomes an idol to many New Yorkers, also making friends with a young zebra, a hippo, and a giraffe.
Alex and the rest meet up with a giant group of animals including ones the same of them plus more. There Alex reunites with his father and mother. Marty meets an enormous pack of zebras that sound exactly like him, Gloria falls for a hippo named Moto Moto (will.i.am) which his name means "Hot, HOT" and makes Melman jealous, because he has deep feelings for her. The giraffe becomes the witch doctor for other giraffes so he can stop his species from dying in holes. Before Alex can join the pride he must complete a challenge, which is scheduled the next day. Makunga now has the chance to become the alpha lion by tricking the returning son.
The next day, Alex is told by Makunga that he should choose an opponent called Teetsi, and so he does and the entire pride gasps as the lion walks up to Alex, and Teetsi appears to be a gigantic beefed up lion. Alex mistakes the challenge as a dance off, and he is easily defeated. Zuba is forced to give up his role as the alpha lion and give it to Makunga and the lion is both confused and angry after Alex tells him he was a star unlike a king. After Zuba insults Alex saying that he isn't a "real" lion, he walks off offended and depressed. Marty becomes angry at Alex because he mistook him for another zebra, and Melman argues with Gloria after she says she is going out on a date with Moto Moto that night. Melman is too shy to admit his feelings to Gloria, but Julien attempts to teach Melman how, and so the giraffe takes off to find Gloria and Moto Moto. He is still to scared, and Gloria thinks it is just an apology and she accepts and then Melman tells Moto Moto to take great care of Gloria in case of the future he is sure she might have. Meanwhile the lost humans including Nana meet up with another group of people who were carjacked as well and they find a camp site and they worship Nana for her great leadership. Mort washes up to the shore (because he fell off the plane when it was about to crash) and he is chased by a shark.
The water hole is dried up the next morning, and all the animals panic because they will die of thirst. Makunga doesn't know what to do, but Julien suggests a sacrifice to the volcano so the water could return. Melman decides to volunteer as the sacrifice. Gloria is completely surprised by this, and tries to talk Melman out of it, but then she realizes he is doing this is impress her and sharing his complete feelings about her. Alex finds Marty and the zebra forgives him and they take off in the hunters preserve, which is were the jungle is located. The duo finds a dam blocking the river to the plains built by the humans. They are ambushed, Alex is captured, and Marty escapes to tell that his friend has been caught. Zuba gets the message and he chases after him. Melman declines the sacrifice to stay with Gloria, and the zebra arrives to tell them about Alex's capture. They then show up at the plane as it is completed and it flies off above the jungle.
The humans are spit roasting Alex above the fire alive, but Zuba arrives in a nick of time. Nana attempts to kill Zuba and Alex with a gun, but Alex shows the humans his famous dance moves to the people and soon enough they recognize the lion as Alex from back at the zoo. Zuba shortly joins in the performance. Nana is impressed, but then points the gun at Zuba and pulls the trigger, but the bullet is deflected by a big bucket lowered down by the plane carrying the rest of the group. The lions enter and they fly off, but before leaving the campsite, they destroy the dam and the water washes back to the pride, who are almost dying from thirst. At the volcano, Mort arrives still pursued by the shark, which falls into the volcano lava (Julien makes a joke about hoping the gods like seafood). The lemurs look down into the plains finding the water returning and making them think the sacrifice worked. Makunga is removed from the throne (after Alex tricks Nana into attacking him), and Zuba and Alex now the alpha lions.
Now that the water is back, the animals can live in peace without Makunga living there anymore. The penguins, chimps, and lemurs decide not to stay as they take off back to Madagascar. Now part of the pride, Alex, Marty, Gloria, and Melman dance off to their new home along with Zuba and Florrie.
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