Ezio, residing in Italy during the Renaissance in the late 15th century, finds his family in Florence the victims of a political plot, and is unable to prevent the hanging of his father and two brothers. His father's final instructions lead Ezio to a chest containing Assassin garments and tools. Ezio's uncle Mario allows Ezio and his mother and sister to shelter in his countryside villa, and trains Ezio in the art of combat. Mario further provides leads to those in on his family's betrayal, and Ezio follows the trail of clues, leading from Florence to San Gimignano, Forlì, Venice and eventually to Rome. Along the way, he encounters several allies that assist in his pursuit, including Leonardo da Vinci, who is able to help decode pages from Altaïr's Codex, creating new equipment. After assassinating many people involved in the plot, Ezio is able to identify Rodrigo Borgia as the leader of the conspiracy, aimed to bring down the Medici family in Florence with the help of the Pazzi family and the Doge of Venice. Borgia later arrives in Venice with the "Apple," the Piece of Eden that Altaïr originally had found. Borgia believes himself to be the "Prophet" that would lead to the discovery of "The Vault", supposedly containing more Pieces of Eden and other knowledge the Templars seek. Ezio, joined by his allies, stop Borgia, who flees without the Apple. His allies, including Niccolò Machiavelli, reveal they are all members of the Assassins, and bring Ezio into their ranks, telling him that they believe him to be the Prophet that will lead them to the Vault.
During one break from using the Animus 2.0, Desmond suffers a hallucination that returns him to the body of Altaïr in Acre, and makes love with Maria, a Templar he had earlier spared (a decoy for Robert de Sable in Assassin's Creed whom Altaïr formed a relationship with in Bloodlines); when Altaïr departs, Desmond finds the memory staying with Maria, and leaves him confused when he recovers. He also learns of the case of Subject 16, one of several previous test subjects used by Abstergo, and the person responsible for writing the images he saw in his room at Abstergo. Subject 16 has managed to hack the Animus and place twenty glyphs that can only be seen and decoded by the Animus user, revealing a video of a man and a woman, claiming to be Adam and Eve, carrying the "Apple of Eden", running through a futuristic setting identified, by a flash of binary code, as Eden.
When Desmond returns to Ezio's memory after he fought off Borgia, they find several corrupted memories, and are only able to return Desmond to Ezio's memories in the year 1499 (The memories comprise of the fictional siege of Flori by the Orsini family and the Bonfire of the Vanities), where Borgia has since become Pope Alexander VI at Vatican City. Ezio's memories reveal that he and his allies have completed the Codex and discovered that "The Vault" lies in Rome, and realize that the Papal Staff is another Piece of Eden; Borgia's plan, upon deposition of the Medici family, would allow him to become Pope and gain access to the staff and the Vault.
Ezio travels to Rome; while his allies distract the rest of the city, Ezio discreetly enters the Vatican and attempts to assassinate Borgia inside the Sistine Chapel (while he preaches the Nicaean Creed in Latin). Borgia manages to escape with both the Apple and the Staff, and attempts to use them to open the Vault, but Ezio catches up and defeats him but does not kill Borgia, since revenge will not bring back his family. In Ezio's hands, the Apple and Staff open the Vault; inside, he finds a holographic figure that identifies him as The Prophet that has brought Desmond and the 21st century observers to the Vault, much to Ezio's confusion. The figure, calling herself "Minerva" and addressing the player directly, reveals that she and others of her kind were part of a far more advanced society (Those Who Came Before) that lived on Earth and created humans in their own image (it is hinted several times throughout the game that humans were created as a slave race). Eventually the two races went to war against one another, soon halted by a natural celestial catastrophe involving a massive solar flare which resulted in the destruction of most life on Earth. The remaining people of both races rebuilt society. The few remaining Ones Who Came Before constructed several "temples" around the world (one being located within the Vatican) that would help the human race to prevent the same disaster from reoccurring once the Ones Who Came Before had become extinct. Minerva insinuates that records of the existence of the Ones Who Came Before, as well as their warning, had been misunderstood and had, over time, evolved into various myths, legends and religions throughout the modern world. As the hologram vanishes, Minerva says "the rest is up to you, Desmond", confusing Ezio.
As the memory ends, Desmond wakes to find that Abstergo has discovered their hideout and, with his assimilated skills from Ezio and Altaïr, assists Lucy in dispatching their forces, led by Warren Vidic, while Shaun and Rebecca pack away their equipment. As the four escape, Lucy postulates that Minerva may have been talking about the Earth's geomagnetic reversal, Desmond realizes that the writing on his wall left by Subject 16 at the beginning of the game were actually warnings, and with time short, Desmond prepares to re-enter the Animus and continue searching his memories for more clues to the location of the remaining temples.
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