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While Scrat (Chris Wedge) inadvertently causes the break up of Pangaea, Manny (Ray Romano) and Ellie (Queen Latifah) are forced to deal with the trials and tribulations of their teenage daughter Peaches (Keke Palmer), who has trouble fitting in with her peers. Ellie tries to support her daughter, but Manny becomes exceedingly over-protective. Peaches' friend Louis (Josh Gad), a molehog, tries to warn her as she tries to approach a mammoth named Ethan (Drake) whom she has a crush on. Meanwhile Sid's (John Leguizamo) family returns, but only long enough to drop off the elderly Granny (Wanda Sykes) before abandoning them both. Manny catches Peaches sneaking off to meet Ethan, and they argue and fall out. Shortly after, a continental break-up separates Manny from the herd. Trapped on a moving chunk of ice with Sid and Diego (Denis Leary), Manny has no choice but to ride out the current. Meanwhile a giant land shift encroaches on Ellie, Peaches, and those remaining on land, causing them to make their way toward the land bridge.
Meanwhile Scrat, in a side adventure, finds an acorn that has a treasure map on it that directs him towards an island. After violent weather pushes them further away from land Manny, Sid, and Diego find Granny who was sleeping inside a hollow tree stump, on the ice raft. Shortly after they are captured by a band of pirates sailing on a floating iceberg led by a Gigantopithecus, Captain Gutt (Peter Dinklage), who attempts to press gang them into his crew. When they refuse Gutt forces them to walk the plank but Manny, Sid, Diego, Granny and Scrat (who was also taken captive) manages to escape and inadvertently cause the ship to sink. Gutt's first mate, a female sabertooth named Shira (Jennifer Lopez) joins them out of no will of her own.
After finding Switchback Cove, they learn that it has a current that would take them back home, but after washing ashore, Shira, still loyal to Gutt, warns him and the rest of the pirate crew, who are also on the island, of their presence. Gutt, wanting revenge on Manny for sinking his ship, plans an attack. In response, Manny coordinates a plan to steal Gutt's new ship to return home, and enlists the aid of the hyrax inhabitants of the island with the help of Sid. Shira decides to leave Gutt, after confronting her feelings about Diego, but she stays behind to ensure Gutt doesn't catch the herd who was attempting to escape on his new ice ship. In a fit of rage Gutt constructs a new ship by breaking two icebergs apart, and sails after the herd, determined to get revenge.
Meanwhile, Scrat, using one of the hyrax's leaf "planes", flies off the island, only to be swallowed by a shark. Peaches finally begins to fit in with the mammoths her age, but accidentally pushes away Louis, by saying they are not friends to fit in with the rest of the group. When she realizes the other teens' careless disregard for danger, she turns her back on them, warning that their extinction will come sooner than they think.
While sailing back on the current, Diego, Sid, and Granny encounter monstrous sirens preying on their deepest fantasies (Sid's siren is voiced by Ester Dean). Fortunately, Manny realizes just in time that none of it is real and saves them all at the last second. Scrat encounters a siren as well, taking on the shape of Scratte (Scrat's old flame from the third film), that Scrat ignores. The siren then turns into an acorn and Scrat immediately runs up and attempts to bury the "acorn", and kills the siren, but is attacked by other sirens but he manages to escape unharmed.
Manny, Sid, Diego, and Granny returns home only to find the land bridge destroyed in the earthquakes and that Gutt and his crew made it there first. The pirates captures Ellie and Peaches forcing Manny to face Gutt. Louis stands up for Peaches and a battle ensues between the herd and the pirates. Shira frees Ellie and helps fend off her former comrades, while Granny's "imaginary" pet whale, Precious, appears and turns the tables on the rest of the lot. Gutt then attempts to kill Ellie but Peaches, using her possum-like skills, manages to save her mother. Manny defeats Gutt in a duel on an ice floe and reunites with his family and friends. In the meanwhile Gutt encounters a siren taking on the shape of a female of his species (also voiced by Ester Dean) and is eaten. With their home destroyed, the entire party and inhabitants then sail to an island that they found to settle down, while Louis becomes a hero among the teenage mammoths, Shira agrees to stay with Diego and becomes a member of the herd.
At the end Scrat discovers the island on the map, known as Scratlantis (a mock up of Atlantis), but his uncontrollable urge to hunt acorns in the acorn-rich city inadvertently causes the entire island to sink when he unplugs an acorn drain holder; Scrat is then ejected into what is now known as Death Valley, California.
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