How do hippopotamuses use scent marks along paths to grasslands?

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Hippopotamuses use dung piles along paths leading inland to grasslands as olfactory signposts, which function as scent markers at night. Both territorial and subordinate male hippos create these piles, which help individuals recognize one another through scent. This behavior is significant because hippos often travel along these familiar paths in the dark, sometimes following one another in a nose-to-tail formation during their nightly treks to feed.1