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The story begins around 150 years ago, in an English town named Wall. Wall is built beside a wall that is a portal to a magical realm of Stormhold. After a young man named Dunstan Thorne (the older version of whom is played by Nathaniel Parker later in the film) crosses the wall in spite of a guard trying to prevent this, he meets a nameless woman, who we later find out is named Una (Kate Magowan). She tells him she is a princess captured and used as the slave of the witch Ditchwater Sal (Melanie Hill). She gives him a glass flower, which she claims will bring him incredible luck, for the price of a kiss. Dunstan makes love to her and returns through the portal to his hometown of Wall the next day. Nine months later, Una sends him a baby in a basket, their infant son, Tristan.

Eighteen years later, the King of Stormhold (Peter O'Toole) is dying, and his four surviving sons (Septimus, Primus, Tertius, and Secundus) fight over who will be the next king while their three brothers already dead (Quartus, Quintus, and Sextus) look on as ghosts. Secundus (Rupert Everett) appears late, and the king tricks him into being pushed off a balcony and down a cliff by Septimus (Mark Strong). Secundus then appears along with the other three dead brothers, his face smashed in from the impact of the ground. The King turns a ruby into a diamond and says that whichever son retrieves it and returns it to a ruby will be king. The ruby knocks a star out of the sky, which the grown Tristan Thorne (Charlie Cox) sees. Tristan promises to retrieve the star for the beautiful yet conventional Victoria (Sienna Miller) to prove his love for her. However, at the same time, three witch sisters; Lamia, Empusa and Mormo, see the falling star, and Lamia (Michelle Pfeiffer), leaves to capture it so, by devouring its heart, she and her sisters will be restored to beauty and power. Tristan tries to cross the wall, but the guard, in spite of his old age, manages to stop him.

Dunstan gives Tristan the glass flower and a package left with him by his mother, which contains a letter and a Babylon candle that allows one to travel wherever one wishes. Though intending to travel to his mother, Tristan accidentally thinks of Victoria and the fallen star and instead finds himself at the star's crash site in Stormhold and ends up bowling over a startled young woman named Yvaine (Claire Danes), who, it turns out, is the star. Tristan resolves to take her back to Victoria anyway and traps her with an enchanted chain and promising her the remains of the Babylon candle if she comes with him to Victoria. Meanwhile, Tertius is killed by a poisoned glass of wine by Septimus, although there was another glass meant for Primus which was instead taken by a bishop.

Yvaine manages to escape Tristan with the help of a Unicorn, but is lured into a trap by Lamia, who is posing as an innkeeper. The stars whisper to Tristan while he sleeps, telling him he must protect 'their sister' Yvaine. He is told a coach is coming, and that he must get on it by any means possible. The coach is driven by Primus. Tristan fails to jump on as it races by, but Primus allows him to get on. When they arrive at the Inn, Lamia kills Primus by slitting his throat (leaving Septimus as the only remaining son and putative heir), and both Yvaine and Tristan barely escape with their lives and the help of the unicorn who dies trying to protect them, when Tristan uses the stub of his Babylon candle to transport them "home"; but because they have differing ideas of home, they go halfway between both their homes, and are left in the sky standing and arguing on clouds.

Tristan and Yvaine are captured by the "notorious" Captain Shakespeare (Robert De Niro) on a flying pirate ship, but he reveals that he is secretly a flamboyant cross dresser who shuns killing and is very proud of his reputation. He used to dream of crossing the Wall to England in much the same way that Tristan dreams of traveling to far away places for adventure. He teaches them various things during their stay on his ship, fencing for Tristan, the piano to Yvaine, and the waltz for both of them. Along the way, they stop at Ferdy the Fence (Ricky Gervais) home where he buys some of their products, which contain lightning. Shakespeare drops off Yvaine and Tristan near Wall (and gives advice to Tristan that his true love may be the star). Shortly afterwards, Shakespeare's crew saves him from the last surviving prince of Stormhold (the ruthless Septimus) and reveal they always knew he was a "whoopsie". While Septimus and Lamia both converge on Yvaine, she and Tristan fall in love though neither would admit it. They end up in front of Sal, who still has Una under her control. Sal tricks Tristan into giving up his magic flower (which protects the wearer from magic) for a ride to the wall and turns him into a 'mouse' (actually a sugar glider) out of spite. On the way Yvaine admits her love to Tristan who doesn't seem to notice anything but his piece of cheese. Sal keeps her end of the bargain and changes him back to human form and in an inn Tristan asks if Yvaine meant what she said in the caravan. Shocked and confused, she is comforted by Tristan who reveals he feels the same way.

While Yvaine sleeps, Tristan leaves her to deliver a lock of Yvaine's hair to Victoria as proof that he found the fallen star and to tell Victoria that he no longer loves her. While there Victoria, seeing how Tristan's appearance has changed, tries to win his heart but Tristan refuses. Humphrey, who planned to marry Victoria, tries to duel Tristan but Tristan proves that he knows how to fence and Humphrey backs away. At the same time, Lamia, Yvaine, Una, and Ditchwater Sal meet at the border between Wall and Stormhold (Yvaine believes that Tristan has returned to marry Victoria and intends to enter Wall to find him. Una steals Sal's caravan to warn her that if she enters Wall, Yvaine will turn into a rock). Lamia kills Sal, freeing Una from her chain. Then Lamia abducts Una and Yvaine and takes them to her home where her sisters are waiting. Septimus and Tristan both pursue Lamia, and form an uneasy alliance, while Yvaine has been strapped down by Lamia and her sisters to have her heart cut out. In the ensuing battle, Empusa and Mormo are killed, but Lamia kills Septimus in turn, and reanimates his corpse to attack Tristan. Tristan crushes him with a chandelier. As Lamia is about to strike the final blow, she instead cuts Yvaines bonds while weeping about how eternal youth and beauty is pointless without her sisters. At first the couple tries to leave but Lamia reveals her happiness that she can keep Yvaine's heart for herself. Yvaine tells Tristan to hold her, which sets off a massive burst of starlight so intensely hot that Yvaine's light kills Lamia. She tells Tristan that she could only do that when she was truly happy, as she was now that she knows he loves her.

Una reveals that she was the only daughter of the King of Stormhold, which makes her son, Tristan, the throne's only surviving male heir. After retrieving the gem from the necklace Yvaine had been carrying, the red from the colourless ruby returns, proving that Tristan is of royal blood. In the end Tristan becomes the new king, and Yvaine his queen. At the coronation, Una, now united with her love Dunstan, presents Yvaine with a Babylon candle. After eighty years of ruling (once their children and grandchildren have grown up), Tristan and Yvaine light the Babylon candle and travel into the heavens, where they become twin stars.