2123202 - Upgrading user database to 15.7sp131 fails with error 2601 "insert duplicate key row in object 'sysusers'" - ASE

Upgrading a user database to ASE 15.7sp130 or higher by using 'in place upgrade' (binary overlay) or loading a dump fails.
ASE error log shows:

  Database '<user_db>': beginning upgrade step [ID   245]: Update system roles in system tables. 
  Initialization of sysusers in database <user_db> changed the system role name from '<group_name>' to 'dtm_tm_role'.

  Error: 2601, Severity: 14, State: 6
  Attempt to insert duplicate key row in object 'sysusers' with unique index 'ncsysusers1'

  Error: 3461, Severity: 20, State: 1
  Database '<user_db>': upgrade could not install required upgrade item '245'. 
  Please refer to previous error messages to determine the problem. Fix the problem, then try again.

  Error: 3462, Severity: 18, State: 1
  Database '<user_db>': upgrade could not install optional upgrade item '1720'. 
  Please refer to previous error messages to determine the problem. Upgrade will continue.

  Error: 3451, Severity: 20, State: 1
  Database '<user_db>': upgrade has failed for this database. 
  Please refer to previous messages for the cause of the failure, correct the problem and try again.

  Error: 3454, Severity: 20, State: 1
  Database '<user_db>': ASE could not completely upgrade this database; upgrade item 1720 could not be installed.
  The transaction log in the database '<user_db>' will use I/O size of 2 Kb.
  ASE could not completely upgrade database '<user_db>', but the database was online when upgrade began, so it will be left online.

SAP Adaptive Server Enterprise

KBA , dtm_tm_role , ha_role , webservices_role , keycustodian_rol , hadr_admin_role_ , replication_main , syssrvroles , sp_helprotects , BC-SYB-ASE , Sybase ASE Database Platform (non Business Suite) , BC-DB-SYB , Business Suite on Adaptive Server Enterprise , BC-SYB-ASE-CE , ASE Cluster Edition (Standalone) , BW-SYS-DB-SYB , BW on Adaptive Server Enterprise , Bug Filed