In the prologue of Twilight, an unknown cat receives a prophecy from StarClan, stating that "it cannot be true". During the book, Squirrelflight struggles with Brambleclaw's hostility toward her, and at the same time she and Ashfur grow much closer, and she wonders if she loves him as she had loved Brambleclaw. Squirrelflight begins to mistrust Brambleclaw for spending time with his half-brother, Hawkfrost. Brambleclaw and Squirrelflight are farther apart than ever, and are constantly bickering. Squirrelflight doesn't know that Brambleclaw is jealous of Ashfur and Squirrelflight's growing relationship until later during the book. During this time, Daisy and her three kits join ThunderClan after Daisy witnesses the Twolegs taking Floss's kits away. Her arrival causes jealousy between Brightheart and her mate Cloudtail, who spends much of his time with Daisy.
Meanwhile, Leafpool and Crowfeather begin secretly meeting each other. Cinderpelt finally confronts Leafpool when she is with Crowfeather. The two medicine cats fight and Leafpool decides to run away with Crowfeather. Deciding her heart is with Crowfeather, she leaves.
Squirrelflight and Brambleclaw are even more hostile when Leafpool leaves until they go to WindClan after finding out that Crowfeather and Leafpool were meeting, thinking that Leafpool had gone to WindClan. Upon finding that Crowfeather had been missing too, Brambleclaw, Squirrelflight, Ashfoot and Tornear follow the missing cats into the moorland where their scents led, but ultimately lost the trail. Squirrelflight states that she could not think of another cat she would rather accompany with her than Brambleclaw, and "presses her cheek on Brambleclaw's muzzle." The two have settled their friendship back into place, but not enough to be in love again.
Badgers are angry with the Clans (especially ThunderClan and RiverClan) because they keep chasing them out. Together, the badgers invade ThunderClan camp and the cats of ThunderClan are thrown into chaos as it becomes increasingly more apparent that they cannot defeat them. A badger kills Sootfur. Leafpool and Crowfeather come across Midnight on their journey. Midnight knows of The trouble that the badgers are doing and is off to warn the Clans (it is clear that she doesn't know when they are attacking). Leafpool and Crowfeather decide to return to ThunderClan to help. Meanwhile, Sorreltail is having her kits, with Brackenfur guarding the nursery, Brightheart tells Leafpool to get Cinderpelt. After Cinderpelt disappers into the nursery, Leafpool returns to fight alongside her Clan. Midnight brings the cats of WindClan into the camp to help fight the badgers. When Leafpool can finally get to Cinderpelt, she is horrified to find her mentor dying, attacked by a badger who broke into the nursery. Cinderpelt reveals to Leafpool that StarClan had already prophesied her death. The badgers are eventually driven out with the help of WindClan. Near the end of the novel, Squirrelflight states that she pushed aside that fact that she had been more worried about Brambleclaw than Ashfur. In the last paragraph, Stormfur and Brook come to visit from the Tribe of Rushing Water.
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