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Each of the demon bosses is associated with a specific dungeon which may disappear after its defeat. The humans and animals are not associated with specific locations to that extent, and they do not remain opposed to Amaterasu for their entire duration of appearance within the game.
All of the human bosses fall into the same general category of demons. Each of them is armed with a sword for hand-to-hand combat and a forceful lunging attack, but can also launch a volley of glowing blades through the air. If Amaterasu can Power Slash all of the blades from a single attack before they strike her, the opponent greys out into Stunned mode and becomes more vulnerable until it wears off. Some of the bosses also have additional stun conditions.
- Main article: Waka
Waka introduces himself with the first boss-like battle in the game, soon after Amaterasu restores the Guardian Sapling in Agata Forest. He confronts her again in Taka Pass as she approaches the Guardian Sapling in that location. Multiple copies of Waka also reappear in the optional Kamui Devil Gate Trial.
Waka's aerial volley has one blade that travels in a straight line.
- Main article: Evil Rao
After Queen Himiko's demise, Evil Rao sheds her earlier pretense of friendship and partially reveals her true nature as a "fox-faced wench", as Issun calls her. Like Waka, she also reappears as multiple copies in the Kamui Devil Gate Trial.
Evil Rao has a deadly stabbing attack from above, but greys out if her sword misses Amaterasu and becomes stuck in the ground. Her aerial volley has five blades that follow Amaterasu's movements.
- Main article: Oki
Oki challenges Amaterasu to battle soon after she reaches Kamui. His battle has two separate rounds for his human and lupine forms.
In his lupine form, his aerial volley has five huge shards of ice that can cause extra freezing damage.
- Main article: Nagi
Mistaking her for Shiranui, whom he regards as a demonic familiar of Orochi, Nagi challenges Amaterasu to a battle on the other side of the Spirit Gate. He also reappears as multiple copies in the Kamui Devil Gate Trial.
His aerial volley has five blades that follow Amaterasu's movements.
All of the canine bosses have a glowing lunge attack, signaled by starting with a special howl. During these lunges, they are super-fast and nearly invulnerable. Some of them also have dirt-based attacks that can damage Amaterasu by hurling sprays of earth at her, temporarily trapping her in the hole if she falls in, or leaving brown lumps that can explode with the force of a Cherry Bomb.
There are four canine boss battles against individual members of the Canine Warriors, arguably a fifth one against the Tube Foxes (see below), and a last one against Oki's wolf form. If Amaterasu returns to the Gale Shrine for the optional training challenge against the Kusa 5, she faces all eight of the canine warriors in a three-part battle.
Each demon boss is associated with a specific dungeon, although some of the demons reappear on the Ark of Yamato at the end. Many bosses are particularly vulnerable to a specific Celestial Brush god that was acquired in earlier parts of their dungeon. After the boss's defeat, the dungeon may disappear and/or a new Divine Instrument may be granted to Amaterasu.
- Main article: Spider Queen
The Vine Celestial Brush technique peels open her abdomen and exposes her eyes to attack. However, if Amaterasu runs up to strike with a Divine Instrument, the Spider Queen may cause considerable damage when her abdomen snaps shut again. Brush attacks from a distance avoid that risk.
The three Bandit Spiders (one in each of the three Devil gate trial caves upon the first entry) are very similar to her and can be fought in similar ways. The defeat of each Bandit Spider causes the appearance of a treasure chest containing a Sun Fragment; also, rather than causing an area to close off, it actually opens up a new one by allowing the first Devil gate to appear in the cave.
- Main article: Crimson Helm
- Main article: Orochi
As each of his eight heads roars in frustration, it can be intoxicated by drawing sake up to its mouth with Waterspout. In the first round of the battle, all of his necks temporarily drop down when three heads are simultaneously intoxicated, allowing Amaterasu to run up them to strike his key vulnerability: the bell on the middle of his back. In the second round of the battle after the bell is destroyed, each head must be individually intoxicated and attacked. In the third and final round, all eight heads rise up in turn for a final attack, for which Amaterasu must assist Susano with Power Slash.
- Main article: Blight
Blight is the boss of the Issun-sized version of the Imperial Palace. It becomes inaccessible after his defeat, which bestows the third-tier Rosary Exorcism Beads. He later reappears on the Ark of Yamato.
The Tube Foxes are arguably the collective boss of the Water Dragon, as they are the last demons faced within the location itself and Amaterasu cannot return to it after their defeat. However, their defeat transforms their remains into the Fox Rods, leading directly into the combat with Evil Rao. As such, they may represent the converse of the battle with Oki: a separate first-round battle with Evil Rao in her vulpine form(s), shortly before facing her sword-wielding humanoid form.
They can be optionally fought once again in the North Ryoshima Coast Devil Gate Trials, but do not cause the reappearance of the Fox Rods.
Like the (other) canine bosses, they have a powerful lunging attack. Their other attacks include launching globes of foxfire through the air, and draining Amaterasu's ink.
- Main article: Ninetails
Ninetails is the final boss of the second major arc of the game. Its dungeon is Oni Island. The island becomes inaccessible after Ninetails' death, which bestows the third-tier Glaive Blade of Kusanagi. Ninetails later reappears on the Ark of Yamato.
Ninetails can switch between the single form of a gigantic multi-tailed fox and the separated forms of nine Rao-like humanoid entities.
As the multi-tailed fox, Ninetails has a powerful full-body lunge and can sink its tails into the ground to spear upward at Amaterasu from a distance. When Amaterasu's brush screen is up, the multi-tailed fox can interfere with her Celestial Brush techniques or even use its own. It is nearly invulnerable, but can be split apart into the humanoids by certain types of attack: extensive flame damage to its tails, and Thunderstorm to its sword when held aloft.
When split apart, Ninetails turns into multiple Rao-like humanoids. One pseudo-Rao always floats inaccessibly in the air; the others (initially eight, one for each extra tail) are scattered across the arena floor, each armed with a sword. The death of each pseudo-Rao destroys one of Ninetails' nine tails until only one tail is left, reducing Ninetails to a huge single-tailed fox for the last round of the battle.
- Main article: True Orochi
True Orochi is the boss of the original Moon Cave on the other side of the Spirit Gate. It becomes inaccessible after his defeat, which bestows the fifth-tier Glaive Thunder Edge. He later reappears on the Ark of Yamato.
Fighting him is very similar to fighting the weaker version of Orochi from the first arc of the game, except that each head now requires two draughts of sake for intoxication. On the other side of the Spirit Gate, the third round of the battle involves Nagi instead of Susano; on the Ark of Yamato, no third round is required.
Lechku and Nechku
- Main article: Lechku & Nechku
Lechku & Nechku are the final bosses of the third major arc of the game. Their dungeon is Wawku Shrine. It remains accessible after their defeat. Their battle is split into two rounds, both with another warrior fighting at Amaterasu's side.
The first round is with Shiranui against Nechku alone; as Shiranui's Celestial Brush Techniques are even more powerful than Amaterasu's, all that Amaterasu really needs to do is stay out of the way and not get hurt. At the end of this round, Lechku helps Nechku escape, but not without losing a small gear which Amaterasu can use as a key to reach a previously-inaccessible part of the shrine.
The second round is with Oki against both owls, who have a range of attacks signaled by objects that appear on top of their heads. Each object can be countered with an appropriate brush technique, temporarily confusing them so Amaterasu can fire Oki like an arrow from a bow. When struck by Oki, each owl greys out and becomes especially vulnerable for a short period of time.
- Main article: Yami
Yami is the final boss of the game. His dungeon is the Ark of Yamato. After his defeat, the game ends.