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Given the non-response on this question, it seems that others have not seen this problem.
The fix in my case was to add the following to each notify characteristic in the database, with a handle specific to each.
After CHARACTERISTIC_UUID128_WRITABLE section, add:
CHAR_DESCRIPTOR_UUID16_WRITABLE (HANDLE_CHARACTERISTIC_CHAR1_CLIENT_CONFIGURATION_DESCRIPTOR, UUID_DESCRIPTOR_CLIENT_CHARACTERISTIC_CONFIGURATION,
LEGATTDB_PERM_READABLE | LEGATTDB_PERM_WRITE_REQ, 2),