On the question of "scent marking" being the explanation for the actual "learned" behavior, my understanding is that "footprint pheromone" would be the odor that one would expect to be left behind on a blossom, or on the surface of the turnstile that must be pushed to expose the food reward.

So, even if the bees were merely following an odor, they still learned to associate an odor that, in nature, means "This blossom has been foraged, nothing left here" and reverse that meaning to "This is where the food is!".  That's an interesting thing to be able to learn in itself, even if it is the only thing actually "learned" here.

The implications here make me wonder if the bumbles tested would be able to differ between the two contexts, and still correctly interpret the odor in each context (field vs lab), or if the experiment left them confused, and, thus, very poor foragers as a result of the "learned reversal" of the meaning of the odor.

But it is a shame to realize that a basic step taken in every feeder-dish experiment for a century or more was neglected here, as replacing the landing cards to eliminate any trace of odor has been a very standard practice.  (See COLOSS Vol 1 Sec 14.4, yes I looked it up)

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