Notice how towers don't like to fire diagonally; they fire much faster when they're shooting on a horizontal or vertical plane. I tried placing towers strictly on the longest stretches of straightaways where they could do just that, with good results:
Ah ha, that's your problem... Remember that leaf tower efficiency is pretty crappy until the max level, where it goes through the roof. Once you're in a safe enough position to do it (around wave 30 or so), build and upgrade leaf towers one at a time till they're maxed, then move on to another. Multiple non-max leaf towers at the end-game is what's slowing you down.
EDIT: Plus you've got an overmax water tower in there. The efficiency of that tower actually goes down at that level. The scribbles over the graphic and the fact that the cost doesn't go up at all leads me to believe that the author meant to remove that unbalanced watertower level from the game but hasn't done it yet.
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