Two years after the events of the previous film, Larry Daley (Ben Stiller) is now CEO of Daley Devices, a direct response television company that sells inventions inspired by his experiences as a night security guard at the American Museum of Natural History. While wealthy and successful he has not had the time to see his museum exhibit friends in several months.
When Larry returns to the museum he learns that it is closed for renovations and upgrades. Interactive holograms will replace many of the exhibits, which are moving to the Federal Archives at the Smithsonian Institution in Washington DC. While Theodore Roosevelt (Robin Williams), Rexy the Tyrannosaurus skeleton, the Easter Island moai (Brad Garrett), and Akhmenrah will remain at the museum, Jedediah (Owen Wilson), Dexter the Capuchin Monkey, and others will no longer come to life since the Golden Tablet of Akhmenrah that animated the exhibits each night will also remain.
After the exhibits leave Larry receives a call from Jedediah. Dexter stole Akhmenrah's tablet and brought it to the Federal Archives, and the evil Pharaoh Kahmunrah (Hank Azaria), Akhmenrah's older brother, is attacking Jedediah and the other exhibits. Larry travels to Washington and visits the National Air and Space Museum, the National Gallery of Art, and the Smithsonian Institution Building, searching for the Federal Archives with the help of his son Nick (Jake Cherry).
Larry locates the exhibits, frozen in their shipping container in the middle of a battle with Kahmunrah and his troops. As Larry obtains the tablet the sun sets and the exhibits, and others at the Smithsonian, come alive. Kahmunrah tells Larry that bringing exhibits to life is just one of the tablet's powers, and that he intends to use it to conquer the world by raising an army from the underworld.
Larry tries to dupe Kahmunrah into giving back the tablet, by telling him about the cube of rubik although is threatened by Kahmunrah. Larry escapes from the Egyptian troops with the tablet, with the help of General George Armstrong Custer (Bill Hader). He meets the beautiful and adventurous Amelia Earhart (Amy Adams), who is attracted to Larry and accompanies him through the National Gallery. Larry and Amelia trap Kahmunrah's soldiers in a photograph of V–J day in Times Square, but Napoleon Bonaparte (Alain Chabat) captures them. Napoleon takes Larry to Kahmunrah—who has also allied with historical leaders Ivan the Terrible (Christopher Guest) and Al Capone (Jon Bernthal), and has rejected fellow villains Darth Vader and Oscar the Grouch—and Jedediah is captured trying to rescue Larry.
Kahmunrah attempts to open the Gate of the Underworld by pressing the symbols on the tablet, which resembles numbers on a keypad, but the combination has changed. The pharaoh forces Larry to obtain the new combination before sunrise by trapping Jedediah in a filling hourglass. Larry and Amelia flee from Russian troops and visit the statue of Abraham Lincoln at the Lincoln Memorial. At the National Air and Space Museum, they consult a group of Albert Einstein bobbleheads who tell them that the new combination is the value of pi (3.14159265). When they are found by Napoleon, Ivan, and Capone, Larry and Amelia escape in the Wright Flyer and return to the Smithsonian Institution Building. The two separate, with Amelia searching for help while Larry delays Kahmunrah.
Napoleon, Ivan, and Capone obtain the combination from one of the bobbleheads. Kahmunrah opens the Gate of the Underworld and summons an army of Horus warriors. The Lincoln statue suddenly arrives, forcing the warriors to retreat to the underworld, and Amelia frees the New York exhibits from their container and recruits other Smithsonian exhibits. As the exhibits battle Larry banishes Kahmunrah to the underworld. Amelia flies Larry and the New York exhibits back to the Natural History Museum. Knowing that she will turn to dust at sunrise if she does not return to the Smithsonian, Amelia kisses Larry and leaves.
Larry sells his company and donates money to the museum to restore the exhibits. Roosevelt and Akhmenrah as tour guides, Attila as a storyteller, and other exhibits come to life in public as "animatronics" at the museum, which now stays open after sunset. Larry is rehired as the night guard, and meets a young woman who looks like Amelia.
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