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

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 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.