Multiple bogosities in rend_cache_lookup_v2_desc_as_dir
rend_cache_lookup_v2_desc_as_dir misinterprets the return value of hid_serv_responsible_for_desc_id. As a consequence, when an HSDir is asked for a descriptor it does not consider itself responsible for, it returns error code 404 ('Not found') rather than error code 400 ('Bad request'), as it was apparently intended to.
To upload designs, you'll need to enable LFS and have admin enable hashed storage. More information