炎热的夏天过去，枝上的树叶慢慢转成红色。中央公园内，松鼠、浣熊们发现过冬的坚果根本不够用，这群小动物可能面临一场前所未有的饥荒。松鼠塞利（威尔·阿奈特 Will Aruuyuuyt 配音）和安迪（凯瑟琳·海格尔 Katherine Heigl 配音）奉命前去寻找坚果，它们计划打劫一辆坚果车，谁知却造成了一场大混乱，还导致好不容易的手的坚果毁于大火。在此之后，塞利遭到流放，只有他的老鼠朋友巴蒂（Robert Tinkler 配音）自始至终跟着他。另一方面，安迪则和自我感觉良好的同伴格里森踏上寻找坚果的征途。最终，安迪和塞利在一座库藏丰富的坚果店重逢，而那里的幕后老板竟然是一个臭名昭著、穷凶极恶的罪犯。 螳螂捕蝉，黄雀在后，犯罪分子策划新的劫案同时，却不知有一群小动物正准备打劫他们……
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