Though his official biography tells the story rather differently from how it actually happened, Starscream has led a life that has seen his every endeavour meet with failure, from his youth as a street hustler in Iacon—where he ran afoul of police officer Orion Pax for the first time—to conning his way into a job as a senator for Vos, only to be left drinking his sorrows away when he finds none of the appreciation or glory he'd hoped for. From there, he joined the Decepticons and witnessed Orion Pax's fall, and his rebirth as Optimus Prime.

Today, Starscream watches Optimus Prime fall once again—from Onyx Prime's mothership down into the ruins of Crystal City, where Optimus, and the Matrix of Leadership within him, are consumed by the miniature singularity at the city's heart. Confident that history has proven Optimus will eventually come back to life, Starscream realizes he has a limited time to act; the time has come for him to finally fulfill the prophecy Metrotitan spoke of, to become the savior of Cybertron destiny claims he is. Devastator believes he has deduced the true meaning of the prophecy: Starscream is the reincarnation of Megatronus, the Fallen Prime. Bumblebee dismisses the notion as absurd, but Starscream figures he can work with it nonetheless, and has Devastator carry him to where the body of the defeated Metrotitan lays.

Aboard his ship, Shockwave instructs Liege Maximo to take a squad of Maximals and finish off the Autobots who were aboard Metrotitan when he was shot down. Maximo's half-hearted protestations at being tasked with such gruntwork are silenced, and he does as ordered, leading the beast warriors down to locate and pick off the survivors: Arcee, Aileron, the Torchbearers, and the badly-wounded Jetfire. Maximo's forces arrive at Metrotitan at the same time as Starscream, Bumblebee, and Devastator, and Starscream orders Devastator go and join the battle as well. Bumblebee offers no rebuke; instead, he appears to be talking to someone else, who Starscream cannot see, and then suddenly disappears...

Windblade, the Mistress of Flame, and Marissa Faireborn watch events unfold from afar, amazed to see that anyone has survived the fall of Metrotitan. Faireborn contacts Trypticon, urging him to get the next generation of protoforms who live within his body to safety; the Dinobots want to join the fight, but Trypticon overrules them, spacebridging himself to Earth to get the protoforms out of the line of fire.

While Arcee holds off the beasts, and the Torchbearers merge into Victorion to engage Devastator, Aileron helps the injured Jetfire get to safety. Starscream, meanwhile, is left alone to clamber up to Metrotitan's head, where he discovers that the fallen Titan still lives. Metrotitan confirms Devatator's belief that Starscream is the reincarnation of Megatronus, and for one, shining moment, Starscream's life of failures and defeats makes sense to him, all his struggle part of some great destiny... until it is all ripped away a second later, when Shockwave appears and reveals that the prophecy is nothing more than a work of fiction, invented and sown into history by him, which Metrotitan only believes in because Shockwave used Regenesis ores to reprogram his mind. If Cybertron had an intelligent leader, Shockwave says, they might have been able to see through his machinations—but by manipulating history and "destiny" to place a fool like Starscream on the throne, Shockwave ensured the planet would become divided, left weak and vulnerable for the force of chaos that is soon to descend upon them all. Shockwave punctuates these revelations with a powerful blast that finishes Metrotitan off and leaves Starscream utterly shattered spiritually.

Elsewhere, Optimus Prime awakens within a shadow-shrouded mirror of Crystal City. He initially believes he has died and gone to the Afterspark, but as he picks his way through the ruins, he realizes he is, in fact, in Infraspace, the transitional limbo between l

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