lpr dos not see default printer - why?

I've set my only printer to be the default printer via CUPS, which lpstat confirms

$ lpstat -p -d
printer MFC-9840CDW is idle.  enabled since Fri 03 Jun 2016 09:05:46 AM
        Online
system default destination: MFC-9840CDW

Nevertheless, trying to print something via lpr does not work as it should:

$ lpr  2016129.pdf
lpr: Error - no default destination available.

It is only when I explicitly set the printer name for lpr, things work:

$ lpr  -P MFC-9840CDW 2016129.pdf

The obvious question is: why?

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