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Kate (Hayden Panettiere) and Humphrey (Justin Long) are two wolves from the same pack in Canada's Jasper National Park but on opposite ends of the social order. Kate, daughter of the main Alpha male Winston (Danny Glover) and his mate Eve (Vicki Lewis), is herself an Alpha and takes her duty to the pack seriously. Humphrey, on the other hand, is an Omega wolf, the lowest of the pack, and spends his days having fun with his other Omega friends. Despite his low rank in the pack, Humphrey falls in love with Kate and is valued in the pack as a peacemaker by virtue of his affable nature.
During her youth, Kate leaves to begin Alpha school with her father, much to Humphrey's disappointment. She returns as a fully trained Alpha ready to procure food for the pack, but on her first hunt, two members of the other, Eastern, pack cross into Western pack territory due to the lack of caribou in their own land. The hunt is ruined and the packs are nearly at war with each other. The two packs have had a bitter rivalry ever since Tony (Dennis Hopper), the Alpha male of the Eastern pack, commanded his wolves to cross over into Western territory (against pack law) in order to hunt, to prevent their pack from starving. Winston and Tony meet up one night and arrange for Kate to marry Garth (Chris Carmack), Tony's son. Kate, having a sense of responsibility and duty, agrees for the good of the pack and meets Garth at the Howling Rock, where the wolves meet to find partners.
Garth is also an Alpha, and at first appears perfect in every way, until it is revealed that he has a terrible howl. Kate, taken aback, leaves the scene to "get water". Humphrey then notices Kate and surprises her. The two talk about Garth, and Humphrey grows jealous because of his feelings for Kate, even though it is against pack law for them to marry. Suddenly, they are shot with tranquilizer darts by some park rangers and taken away to Idaho. The two wolves wake up at Sawtooth National Recreation Area, where they meet a French goose named Marcel, (Larry Miller) who has an avid interest in golf, and his caddy, a duck named Paddy (Eric Price). Humphrey and Kate learn that they were relocated to repopulate the species. Kate is horrified by the idea, but Humphrey is more interested, as this would be his only chance to be with Kate. Marcel and Paddy agree to help the wolves return home to Jasper Park. Humphrey is disappointed, but Kate demands to leave because of her responsibility to the packs. Humphrey offers to help her and Kate agrees.
Trouble brews back at Jasper Park as the two packs discover that Kate has gone missing. Tony warns Winston that on the next full moon if Kate doesn't return to marry Garth, he and his pack will declare war for their territory. Lilly (Christina Ricci), an Omega wolf who is Kate�s younger sister, has a crush on Garth and suggests that she show him around.
Meanwhile, Kate and Humphrey attempt to get home in the back of a camper, which goes well until they stop at a gas station. Humphrey leaves the camper to go to the bathroom, but is mistaken for a rabid wolf after he eats a cream-filled cupcake. Seeing one of the men about to shoot Humphrey, Kate attacks the man with the gun, and the two make their escape into the forest. The camper they had stowed away on leaves. Still seeking a way back to Jasper Park, Kate comes to a ravine and attempts to cross it. She slips down the cliff, but Humphrey saves her life by swinging from a vine and letting her grab his tail with her teeth. Kate is very grateful and seems to like Humphrey a lot more. After that the two find a den for the night.
The next morning, Marcel and Paddy find the wolves and direct them to a train over a snowy mountain that will take them to Jasper. Climbing the mountain, Humphrey meets a grizzly bear cub who is unfamiliar with wolves. While they are playing, the cub gets hurt when a snowball is thrown at him. This angers a trio of adult bears (most likely the cub's family) who think that Humphrey was trying to harm the cub. When Kate realizes that Humphrey is in danger, she rushes in to help him, causing them to slide down the other side of the mountain. Humphrey saves Kate using a log sled like the ones he learned to play on as a young Omega, and as the train passes by the two wolves manage to get into a boxcar just in time.
During this time, Lilly and Garth are bonding. At the midnight howl, Lilly is not deterred by Garth's less than perfect howl and compassionately teaches him. At the same time, Humphrey and Kate begin to howl together on the train. Even though both couples are not allowed to sing together, it is clear that they are in love. By this time, the full moon has come and the packs declare war against one another.
Shortly after, as the train passes by Jasper, Humphrey attempts to tell Kate how he feels about her, but the sight of the packs fighting causes her to jump off the train before he can finish. Kate stops the fight by announcing that she will marry Garth for the good of the packs, much to Humphrey, Garth, and Lilly's dismay.
The next day, a heartbroken Humphrey decides to leave Jasper and travel on his own, and shares his goodbyes with Kate, Marcel, and Paddy. Kate feels heartbroken after realizing how she feels about Humphrey. During the ceremony, Kate decides not to marry Garth, declaring her love for Humphrey, after which Garth declares his own love for Lilly. Winston is confused and Tony is enraged that his son is in love with an Omega, which goes against pack customs. His pack starts to fight the other pack out of anger, but the conflict is cut short when a massive stampede of caribou interrupts the scene. Winston and Tony get caught in the stampede and Kate and Humphrey (who decided to return) rush in to save them using a log as a sled. Kate is hit by a caribou, and Humphrey shields her with his body while Winston and Tony growl at the passing caribou in order to protect Humphrey from getting hit. After the stampede passes, it appears that Kate has been trampled to death. Upon her apparent death, Humphrey howls sadly, and the wolves of both packs also begin to howl in mourning, but she wakes up to the joy and relief of the packs. Now, at long last, Humphrey and Kate reveal their feelings for each other, taking turns whispering to one another. Winston and Tony then decide to unite the packs and retire. When they wonder what to do when they retire and grow old, Marcel says, "Gentleman, may I introduce you to golf?" and shoots a golf ball at the screen.
Later at the Moonlight Howl, all the wolves celebrate the marriages of Kate to Humphrey, and Garth to Lilly. The film ends with Humphrey and Kate singing a duet, and the camera pans upward to the full moon.
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