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The story continues from the events of the second film, where the Penguins left Alex the Lion, Marty the Zebra, Melman the Giraffe, and Gloria the Hippopotamus, alongside lemurs King Julien, Maurice and Mort in Africa. The group wave goodbye as they leave, but Skipper suddenly shouts that they are never coming back, and the aircraft "warps" away. Alex turns around, and finds that all his friends and he himself have become extremely old while waiting for the penguins to come back. He wakes up suddenly, realizing the whole event was just a dream, but is still miserable to be stuck in Africa.
Marty, Gloria and Melman have built him a mud model of New York City in honor of his birthday, but it only makes the lion more homesick. He decides that the group should go to Monte Carlo, get the Penguins, and have them fly them back to New York City. Everyone agrees.In Monte Carlo, the Penguins and two chimpanzees, Mason and Phil (disguising as the King of Versailles) keep winning in gambling. Their plan is to come go back for the group after this final win, but the Four's attempt to reach them blunders and starts chaos in the Hotel de Paris. The security call Monaco Animal Control Captain Chantel DuBois in order to capture the animals, but she wants them (specifically Alex) dead as trophies. During a high-speed chase between the relentless DuBois and the animals in a truck driven by the Penguins to reach their aircraft, DuBois's four associates all end up injured, but DuBois keeps after the group. When they finally get to the plane, Dubois manages to get a wire hoop around Melman's neck and holds onto it even as the aircraft takes off. Alex manages to cut the wire, and DuBois falls into a hotel's rooftop pool.
Above the skies of France, the plane's engines fail again and it crashes into a suburban train yard with the authorities closing in. Their only chance of escape is passage on the train of Circus Zaragoza, but the Russian tiger Vitaly will not let them on. However, when the Four desperately claim they are circus animals themselves, Stefano the sea lion and Gia the jaguar who Alex is smitten with upon first sight of her let them come aboard.
When asked about their act, Alex says that he climbs onto his rock, roars, and jumps off. The circus animals seem unimpressed, and the group quickly "enhances" it -- Alex jumps off his rock towards a pool of aquatic cobras, then pulls up at the last second by igniting a jetpack, and then he releases balloons into the air for all the children of the world. Stefano thinks this must be "Trapeze Americano", and Gia seems to perk up when she hears this.
The animals soon learn from Stefano that they are performing two legs in Rome and London, where they plan to impress a promoter to get them on their first American tour. When Vitaly begins to threaten Alex, since stowaways are not allowed on the train, the penguins drop in through the ceiling and state their intentions to purchase the circus. The next day the chimps (again posing as the King of Versailles) offer the circus manager a bag of jewels for the circus deed. He and the human acts seem suspiciously delighted to sell the circus.
Meanwhile, Julien encounters Sonya the Bear, a huge non-speaking brown bear who rides a tricycle. He falls in love with her and they go on a city tour across Rome while the others are preparing for the performance at the Roman Coliseum. During their escapades (which include knocking down ancient ruins and stealing the ring of a monarch), Sonya breaks her tricycle. In an effort to comfort her, Julian trades the ring he stole for a motorcycle, and the pair zoom off just as DuBois arrives in Rome.
She spots Julian and Sonya and leaps onto a police motor scooter to follow; however, the scooter she is riding is chained to several other scooters and moves slower. She eventually looses control of the scooter and crashes into a fountain. Several police officers come to confront her, and she shoots each with a tranquilizing dart. However, when she comes to the last officer, she has run out of darts, and is arrested.
To Alex's horror, the circus is awful; Vitaly moves his plastic hoop instead of jumping through it, Stefano swallows some of his juggling balls and they hit people or other acts as he coughs them out, an elephant sits on a boy, and one of the dancing dogs is knocked into a man's hot dog. The angered audience demands refunds to the point chasing the circus to the departing train. Julian instructs Phil and Mason, who are not yet on the train, to simply throw him the bags of money. They try, but the money ends up scattering all over the station. Though this serves as a distraction and they escape the enraged people, the circus is left completely broke.
En route to London, Stefano reveals to Alex that Vitaly used to be their inspiration. Once a professional hoop jumper, he would perform physically impossible feats by dousing himself in olive oil to become slippery and leaping through increasingly smaller hoops. One day, however, he attempted to jump through a flaming pinkie ring. The highly flammable oil caused his fur to catch fire and destroyed his confidence; the other animals suffered by his example. An inspired Alex has the train make a stop at the Alps and convinces the performers to rework their previous act to become the opposite of the successful human-only Cirque du Soleil as an animal-only circus. Among the acts developed, Stefano and Marty train to be living cannon balls, Gloria and Melman develop a tightrope dance act, and Alex teaches Gia the beginner's level of "Trapeze Americano"; it is painfully improvised but they both enjoy it.
Meanwhile, DuBois was captured in Rome after causing problems with the local police officers, but escapes and learns about her quarry. Once accomplished, DuBois recruits her injured men and heads towards the Alps, forcing the circus (now going by the name Fur Power) to proceed to London despite their incomplete rehearsals. In London, the Penguins learns that the promoter has arrived and Alex learns from Stefano that Vitaly wants to quit the circus. Alex confronts Vitaly, and after a while, the tiger realizes that he loved his act because it was impossible, and makes the decision he will perform, no matter what the outcome may be. Per Alex's suggestion, Vitaly opens the act by using slippery conditioner instead of olive oil. He leaps through the flaming pinkie ring to the crowd's delight, and the show is both spectacular and successful, with a killer light show and fantastic acts; Alex and Gia perform trapeze, Stefano and Marty are repeatedly blasted out of cannons, Melman and Gloria dance across multiple tightropes, Sonya the Bear rides her motorcycle with Julien, the dancing dogs "dance" on rocket-powered skates Kowalski designed, the horses prance around wearing butterfly costumes, and the elephants blow colored smoke, resembling fire at the audience. Both the audience and the animals have the times of their lives.
However, after the impressed promoter arranges for an American tour, DuBois shows up. The Penguins manage to blast her and her companions out of the cannon, but Alex is forced to confess to the circus group about their zoo past and intentions to return there. The circus animals are so hurt, they turn their backs on them. The Penguins tearfully pretend to be just as surprised by this as the circus animals. Meanwhile, Julian and Sonya fight, and he decides to leave her so he can be King of New York; however, neither are happy about his decision.
Finally, both the zoo animals with King Julien and circus animals (along with the Penguins,Chimps,Maurice and Mort) arrive in New York City separately. While the circus finds that they are incomplete without the Four, that group finally return to the closed Central Park Zoo and realize that their time of freedom has changed them too much for them to return to captivity. Unfortunately, they are all tranquilized by DuBois. The Zoo Staff suddenly appear and thank the annoyed DuBois for (they think) returning Alex the Lion. A tranquilized Julian manages to stumble back to the circus and tells Sonya he wants to stay with her. When the circus animals learn what has happened to the Four, they make the decision to rescue them.
Alex and the others wake up in their old habitats, now with tall chain-link fences so they can't escape. A ceremony is being held in honor of DuBois rescuing and returning Alex the lion "and three other animals" to the zoo. However, Alex watches DuBois load her dart gun with a poisoned dart and conceal the weapon inside an "Alex #1" glove. She fires at Alex, but just before the dart hits him, Gia suddenly pulls him out of the way.
The circus animals built a flying circus, called the "Operation: Afro Circus Rescue", and held up by multiple clumps of balloons. The animals manage to stall DuBois long enough to rescue the Four, finally flinging her right through the brick wall of the reptile house. However, as they start to pull away, they realize Stefano is still on the ground.
DuBois comes out of the ruins of the reptile house, followed by several cobras who slip into the Penguin's old habitat. She grabs Stefano and slowly begins dragging him closer to her gun, with the intent of killing him. Alex and Gia trapeze down and manage to grab Stefano, but DuBois is still holding on. Alex grabs DuBois and then lets go of the trapeze, leaving Gia and Stefano to swing back to safety.
The struggling pair tumble downwards toward the Penguin habitat, where the now-aquatic cobras are waiting. At the last second, however, two of the dancing dogs, flying on their rocket skates, grab Alex and pull him up. The lion drops an enraged DuBois in his habitat, where she is tranquilized by Maurice and Mort. The dogs fly Alex back to the Afro Circus, and one clump of balloons comes loose. The children at the zoo are delighted by the balloons, and Stefano realizes that Trapeze Americano is real.
Deeply grateful for their new friends' help and now valuing their freedom, Alex and his company decide to stay with the circus and they all happily proceed on their American tour while DuBois and her men find themselves on a cargo box bound for Madagascar.
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