It was OK to run on Heroku with "whitenoise.storage.CompressedStaticFilesStorage", but Error500 raised when using "whitenoise.storage.CompressedManifestStaticFilesStorage".
Cause
The cause was in Dockerfile. APP_ENV=prod and DEBUG=false were set in Heroku, but these environment variables were not added in CircleCI. Therefore the correct container image didn't run on Heroku.