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But it's not doing what quarantine normally does; quarantine says that the app cannot be opened because it was an unidentified developer, and you can work around that by ctrl-clicking the package and selecting "open." This was a completely different error message, about the app being damaged.

The really weird thing is that when I download it through the app vs the web and do a byte-for-byte comparison between the resulting .app bundles, though, it finds no differences.

Yeah, that's what I'd usually expect of quarantine. This is a different outcome so it was confusing, but it looks like the attribute quarantine sets on the app is breaking it if it's compiled with newer versions of GameMaker, than the one I previously used. The Terminal command restores it.