Yes, he has a hell of guillotine with a lot of variations; which probably comes from his strong wrestling background and working from the front headlock. Lindland doesn't really have great BJJ, but he does have a lot of guillotines as well.
I may be biased... but the answer is wrestling for sure. The reason is because if you are great at wrestling you have the opportunity to control where the fight takes place. If you are great at JJ but only good at wrestling... what happens when you are fighting someone better on their feet that you can't take down? Wrestling gives you the ability to keep it standing if someone is better in submissions off their back... or gives you the ability to take it to the ground if your opponent is better on their feet. Also... if you are already great at wrestling, and get good at JJ... it is rather easy to control the fight from the top position without ever getting into any real danger of a submission.
I remember Lindland submitting Lutter with some sort of powerful Guillotine but im not going to pop in the DVD to find out which. Do you remember ?
Yup, front headlock position to guillotine, stands up with it, sits down and throws a leg over. http://www.mma-core.com/videos/_Matt_Lindland_vs_Travis_Lutter_UFC_52_Round_2?vid=10003353 He also guillotined Carlos Newton: stood out of an attempted half guard sweep, got a mounted guillotine, end game. http://www.mma-core.com/videos/_Matt_Lindland_vs_Carlos_Newton_IFL?vid=10001122&tid=100
Yeah, Hughes is still evolving and improving his boxing and BJJ despite being comfortably past his physical peak. It sucks for certain guys where the game is evolving like it is. I'm glad to see Hughes finalise a legacy as many said he was rubbish, would never hang with this generate of complete fighters etc