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I guess it happens on multiple levels. No reason it would be exclusive to one.

Certainly individual ants learn stuff

https://royalsocietypublishing.org/doi/10.1098/rsos.190778

But adaptions also happen by evolution and (very speculatively) might also happen at the colony level by it changing its responses somehow. E.g. I could imagine an ant colony suddenly faced with many attackers adapting by breeding more soldiers.



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