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I see them as a part of this tradition:

http://www.godecookery.com/macabre/macabre.htm

http://www.godecookery.com/macabre/holdod/holdod25.htm

I love these pictures, but it's all a bit medieval and therefore off-topic.

http://www.megalithomania.com/show/site/655/1867

Tomb shrines for the early saints were a big thing. A lot of these had holes in that you could reach through and touch the bones of the interred saint for a blessing. The symbols on the gravestones could relate to this practice, but I don't think so.

The symbols that most often occur side by side with the S&XB are the hour-glass, the bell and the coffin. The symbolism is pretty clear.

before I shut up about this altogether, though, here's some better links:

http://www.geocities.com/ppollefeys/dance.htm

http://www.geocities.com/ppollefeys/dm09a.htm

http://www.geocities.com/ppollefeys/dm10a.htm

notice the skull and crossbones in this one (and also the date, which is 1750):

http://www.geocities.com/ppollefeys/dm_ridinger.jpg