Abe's Oddysee had the perfect tone in my opinion, the enemies were shadowy and formidable and the humour was always secondary to the story.
I don't want to downplay the humour in the games but my favourite levels and cutscenes were always the most serious, atmospheric parts; For instance, the intro scene to MO where Munch gets caught was one of the best scenes of the series, despite the rest of the game.
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And what happened to "Scrabs are fiercely territorial"?
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Scrabs are seen running around together in an AO cutscene as well to be fair, and you could argue that they're supposed to be a pack in the final chase sections in the temples. As far as I know the ones who fight each other are all supposed to be alpha males, there can only be one per pack and they fight to the death.
But yeah the Scrab/Paramite sections were IMO the biggest disappointment in MO... we're probably going off-topic ranting about that game anyway.