This is first thing. The other is precursor of law and social norms. It is hard to explain to people, that they shouldn't steal, because "it's wrong" and it hurts society. It's easier, when you tell them they will burn in hell for that.
Though, I'm not sure if hive-mind intelligences or strictly hierarchical biologies (e.g. queens/workers/drones) would even need Law or guidance for social conduct, but they may still invent stopgap explanations (religions) on their way to understanding the weather and stars and such.
So I think the GP is correct, once a society encompasses multiple distinct tribes, it is easier to "guide" people with (threats of) in-group punishment than with out-group empathy.
The in-group out-group is some higher part of the brain that rationalised our values.
Wouldn't it make way more sense to explain why it's wrong? It has nothing to do with hurting society, but another individual.