Gokuu Son and his friends are out camping when three mysterious enemies ambush them. After a quick fight, a fourth enemy named Cooler joins the fray. Seeking retribution for his younger brother Frieza after he was defeated, Cooler attacks Gokuu; however, the latter manages to escape with Gohan Son and goes into hiding. Unsatisfied and infuriated with the outcome, Cooler and his men begin the hunt to uncover Gokuu's location and kill him to carry out their plans of exacting revenge.

The Celestial Records speak of the existence of the Seraphim, a race of divine beings who give blessings to humanity and are offered prayers by them in return. Those who are anointed with the ability to interact with these spirits are known as Shepherds. Hailed as heroes for their prompt appearances in times of crisis, while also being feared for their power, the Shepherds are imprinted in common folklore along with the Seraphim. Sorey is a young human who has spent his entire life living in harmony alongside the Seraphim in the village of Elysia. Fascinated by the myths of the Celestial Records, he explores some nearby ruins with Mikleo—his childhood Seraphim companion—hoping to enlighten himself about the Seraphims' history with mankind. Unfortunately, they become trapped in the depths of the historical site during their investigation. While searching for an exit, they come across a mysterious girl who desperately seeks the help of a Shepherd to save the world, which is on the brink of being consumed by darkness. Despite Mikleo's warning about making contact with other humans, Sorey decides to help the stranger, which unknowingly leads him closer to the dream of peaceful coexistence between man and Seraphim.

In recent years, problems regarding atmospheric instability have greatly burdened the Hyland Kingdom, affecting the quality of crops and the citizens' health across the region. One day, the people of Guriel discover an eerie cloud of mist hovering over their town; its severity captures the attention of royal princess Alisha Diphda. She sends her subordinate Crem to investigate the situation, but receives no news from her for two weeks. Worried for Crem's safety, Alisha departs for Guriel herself with her attendants in tow. What awaits them there, however, is an issue of unprecedented scale.

The second season of Tales of Zestiria the Cross.