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They are defined in the Desktop Entry Specification. In fact there are also %f and %F options.
They tell the desktop manager how the program handles multiple files. Like if I select a bunch of files in Dolphin and select "Open with Firefox".
The small %u means that the program expects only a single argument at a time. So invoking with multiselect is actually launching the program several times, once with each file as an argument.
Capital %U means that the program can take them all as a list in a single go.
%f and %F is just the same thing but stands for local Files whereas U is for URL, which works for local files also. This is probably why you don't see much f's.
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