KeyError for list operation after upgrade

Today I tested an upgrade from Plone 3.1 to 4.3 and then on to 5.0.6.
When I edit a page and want to search for an image, I get a traceback:

Traceback (innermost last):
  Module ZPublisher.Publish, line 138, in publish
  Module ZPublisher.mapply, line 77, in mapply
  Module ZPublisher.Publish, line 48, in call_object
  Module plone.app.content.browser.vocabulary, line 101, in __call__
  Module plone.app.content.browser.vocabulary, line 323, in get_vocabulary
  Module plone.app.vocabularies.catalog, line 594, in __call__
  Module plone.app.vocabularies.utils, line 7, in parseQueryString
  Module plone.app.querystring.queryparser, line 29, in parseFormquery
  Module plone.registry.registry, line 38, in __getitem__
KeyError: u'plone.app.querystring.operation.list.contains.operation'

Looking in the configuration registry, I see lots of things starting with plone.app.querystring.operation, but nothing starting with plone.app.querystring.operation.list.

Going to portal_setup and applying the plone.app.querystring:default profile, fixes it. But it should work out of the box after an upgrade.
I don't know where this exactly goes wrong. There were no unapplied upgrade steps.

Start with the traceback entries in plone.app.content.browser.vocabulary, plone.app.vocabularies.catalog, plone.app.vocabularies.utils, and plone.app.querystring.queryparser, then compare the registry after the reported upgrade with the plone.app.querystring:default profile. Reproduce the image search and upgrade path, and verify that the list operation is available without manually applying the profile.