Unity on Mac upload to Oculus Developer Hub error "Please sign your APK then resubmit"?

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I've been through several forums and other questions addressing this issue and similar including this blog post http://blog.wafrat.com/publishing-an-app-on-the-oculus-quest-store-on-a-mac/ and following this tutorial verbatim https://youtu.be/Z3jlO76NDAY .

I am trying to upload my APK to the Oculus App Lab via Oculus Developer Hub. Despite generating a new keystore in Player Settings here:

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And generating a new manifest file via Oculus > Tools > Generate store ready manifest, I get a failed upload with the error

ERROR: Please sign your APK and then resubmit

The keystore also makes the build not load to my headset via ODH, leading me to believe that something is wrong with my APK signing in Unity.

I am in Unity 2020.8.3 with all Android modules installed. What can I do to properly sign my APK?

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