I solved it. 

Just to be sure, I first moved the token directory from 
/var/lib/softhsm/tokens/ to /var/lib/opendnssec/tokens/. I did this by setting 
"directories.tokendir = /var/lib/opendnssec/tokens/" in 
/etc/softhsm/softhsm2.conf. Then I executed the migration steps described by 
Capser again.
Unfortunately this still resulted in the same error, when starting enforcer and 
signer.

Then I noticed that the permissions of the token directory and files within 
this directory. The were set to 'root' while opendnssec runs on user 
'opendnssec'. After changing the owner of the directory and the files within, I 
was able to succesfully start enforcer and signer. 

So it took me a couple of days, but it seems that opendnssec is running again. 
Casper, Yuri and Berry, thanks for your help!

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