The other thing it might be is a certain looseness in the spokes...
...or something as mundane as a bit of grot trapped inside the box section of the rim.
(Thread resurrection alert)
I'm beginning to think the issue could be spoke/eyelet/nipple related, so you could well be on the right track, MV.
Can those creak (e.g. when they get dirty?).
The spokes don't feel loose though. Wheel is true.
Since I first posted the question, the noise has become steadily more noticable. The sound is now not dissimilar to (but a fair bit quieter than) the popping/crackling sound of running studded ice tyres on tarmac.
Furthermore, I can now reproduce ( a quieter version of) the sound whilst walking the bike with a bit of weight pushing down on the saddle. I can also detect a light creaking which appears to come from the rim area if I push sideways on the tyre. I can't remember now if I also tried that pushing definitely on the rim alone to try and rule out the tyre/rim interface. I guess I could take off the tyre and tube and try again.
It sounds like the creak comes from the rim area. My supposition is that it's either the tyre creaking in the rim (Marathon+ 28mm in a rigida Sputnik rim, so it's about the narrowest tyre you'd want to put on there) or it's the spokes/nipples/eyelets.
(I've since rules out the inner-tube theory I proposed upthread (thanks to an East-Anglian flint). I've also greased all the spoke crossings in case it was those, with no effect.)
I guess it could still be something at the hub, e.g. bearings etc. (I know that it's often hard to pin down the source of noise since it can get transmitted through e.g. spokes to other areas which act as sounding boards...), but as I say there's no noticeable play in the hub.