Just catching up on this thread - I was a summer worker in Yellowstone a very long time ago. Likely he was altitude adjusted as the whole park is pretty high and after 3 months living and hiking in the park he would be in good physical condition.
There are many peaks in the park that don't have established summit trails but rather routes and these are much easier to get yourself lost on especially in poor visibility. we had no GPS or phones when I was there and word of mouth\ paper maps were how you would hope to know where to go. It should be noted that unlike the Tetons there are virtually no true rock climbing routes in Yellowstone as quality of the rock is poor (especially as contrasted to the Tetons).
I'm surprised he was so completely off route since he had a phone and these days it is much easier to see where you are versus your intended path. But mapping apps drain phone batteries and they drain even quicker in the cold
It cannot be overstated how huge this park is and as mentioned elsewhere the backcountry of Yellowstone is incredibly remote.
All the best to his family and I hope they use extreme caution in continuing the search.