Created by the Black Wolf, Gaius van Baelsar, Crania Lupi were empowered by the XIVth Imperial Legion to carry out imperial will in all Ala Mhigan Territory, and are known for their cruelty and dealing out reprisals to any who would resist. Though Gaius had well-meaning intentions for the Crania Lupi, his Garlean peers were not so generous in their views, often scorning the Skulls as "savages". Their fellow Ala Mhigans likewise view the Crania Lupi with shame and anger, calling them traitors and kin slayers.
The regiment's commanding officer, the hardened Fordola rem Lupis, leads the Skulls with fierce determination with the goals to force the Garleans to accept her countrymen as equals. Though she and her Skulls serve loyally, the Garleans continue to treat the Crania Lupi as disposable cannon-fodder.
Following Gaius's defeat and the disbandment of the XIVth Legion, the Crania Lupi fell under the authority of the Zenos Galvus and the XIIth Imperial Legion. Though the Skulls' treatment under the XIIth was not any better, Fordola was able to gain the favor of Zenos for her brutality, and the two struck a crippling blow against the Ala Mhigan Resistance and Eorzean Alliance. This was not to last however, as after the Scions of the Seventh Dawn regain their strength (and liberate the nation of Doma), the Crania Lupi take losses. Fordola's lack of judgment spurs the Ananta Qaylana tribe into summoning a primal after murdering the broodmother's daughter. Though humiliated by these losses, Zenos allows Fordola a chance to make amends by allowing herself to be experimented on using the Resonant. She survives the procedure and is given new orders to stall the Eorzean Alliance's advance in Specula Imperatoris by using the main cannon in Castrum Abania to destroy the installation. This kills many imperial soldiers and her fellow Skulls.
With her soldiers dead by her own hand, Fordola is considered the last of the Crania Lupi, but she is subdued by the Scions and Lyse Hext. Fordola is imprisoned for her crimes as the resistance and Eorzean Alliance defeat Zenos and liberate Ala Mhigo from the empire.