I forgot one – grit, mud or caked dry lube between cogs, but that tends to make it skip all the time. It would fit in with the immersion being the cause.
Cynic-al, if there is play in the freehub, and you rock the freehub, then the teeth of the larger cogs move more than those of the small cogs due to the different radius, meaning the alignment is further out on the big cogs.
The fact it does this with little pressure but not big stomping points to something getting aligned or being stopped from moving by the higher loads rather than chain/cog/chainring wear.
Having said all that I’d say it’s probably chain wear – buy a chain wear tool and be prepared to replace the chain (probably), cassette (probably) and the chainring that gets the most use ( probably not). 1500 miles is about 3 chains for me but my bikes do like mud.